Ecology

Geography and climatology of study areaThe area of study, Ghana, is on the coastal edge of tropical West African, bounded in latitude 4.5° N and 11.5° N and longitude 3.5° W and 1.5° E, and characterized by a tropical monsoon climate system23,24. Figure 1 shows map of the study area indicating the selected twenty two (22) sunshine measurement stations distributed across the four main climatological zones and Table 1 summarizes the geographical positions of selected stations.Figure 1Adapted from Asilevi27.Map of the study area showing all twenty two (22) synoptic stations distributed in four main climatological zones countrywide.Full size imageTable 1 Geographical position and elevation for study sites.Full size tableAtmospheric clarity over the area is closely connected to cloud amount distribution and rainfall activities, largely determined by the oscillatory migration of the Inter-Tropical Discontinuity (ITD), accounting for the West African Monsoon (WAM)25,26.Owing to the highly variable spatiotemporal distribution of cloud amount vis-à-vis rainfall activities, resulting in contrasting climatic conditions in different parts of the region, the country is partitioned by the Ghana Meteorological Agency (GMet) into four main agro-ecological zones namely, the Savannah, Transition, Forest and Coastal zones as shown in Fig. 123. As a result, the region experiences an estimated Global solar radiation (GSR) intensity peaks in April–May and then in October–November, with the highest monthly average of 22 MJm−2 day−1 over the savannah climatic zone and the lowest monthly average of 13 MJm−2 day−1 over the forest climatic zone27.Research datasetsGround-based measurement dataDaily sunshine duration measurement datasets (n) spanning 1983–2018 where derived for estimating Global solar radiation (GSR). The measurements were taken by the Campbell-Stokes sunshine recorder, mounted at the 22 stations shown in Fig. 1, under unshaded conditions to ensure optimum sunlight exposure. The device concentrates sunlight onto a thin strip of sunshine card, which causes a burnt line representing the total period in hours during which sunshine intensity exceeds 120.0 Wm−2 according to World Meteorological Organization (WMO) recommendations27. The as-received daily records were quality control checked by ensuring 0 ≤ n ≤ N, where N is the astronomical day length representing the possible maximum duration of sunshine in hours determined by Eq. 1 from the latitude (ϕ) of the site of interest and the solar declination (δ) computed by Eq. 227:$$ {text{N}} = frac{2}{15}cos^{ – 1} left[ { – tan phi tan {updelta }} right] $$
(1)
$$ {updelta } = 23.45sin left[ {360^{{text{o}}} times frac{{284 + {text{J}}}}{365}} right] $$
(2)
where J represents the number for the Julian day of the year (first January is 1 and second January is 2).NASA-POWER Global solar radiation (GSR) reanalysis dataThe satellite-based Global solar radiation (GSR) dataset for specific longitudes and latitudes of all 22 stations, assessed in the study, were retrieved from the National Aeronautics and Space Administration-Prediction of Worldwide Energy Resources (NASA-POWER) reanalysis repository based on the Modern Era Retrospective-Analysis for Research and Applications (MERRA-2) assimilation model products, developed from Surface Radiation Budget, and spanning equal study period (1983–2018). The datasets are accessible on a daily and monthly temporal resolution scales at 0.5° × 0.5° spatial coverage via a user friendly web-based mapping portal: https://power.larc.nasa.gov/data-access-viewer/17. The advantage of the NASA-POWER reanalysis GSR, is the wide spatial coverage, and thus can be used to develop a high spatial resolution of solar radiation across the study area.The POWER Project analyzes, synthesizes and makes available surface radiation related parameters on a global scale, primarily from the World Climate Research Programme (WCRP), Global Energy and Water cycle Experiment (GEWEX), Surface Radiation Budget (SRB) project (Version 2.9), the Clouds and the Earth’s Radiant Energy System (CERES), FLASHFlux (Fast Longwave and Shortwave Radiative Fluxes from CERES and MODIS), and the Global Modeling and Assimilation Office (GMAO)17. Table 2 shows the source satellites and the corresponding temporal coverage used in the development of NASA-POWER GSR products.Table 2 Satellites providing the NASA-POWER GSR datasets20.Full size tableThe monthly average NASA-POWER all-sky shortwave surface radiation reanalysis products are statistically validated, showing reasonable biases of − 6.6–13%, against a global network of surface radiation measurement metadata in an integrated database from the Baseline Surface Radiation Network (BSRN) of the World Radiation Monitoring Center (WRMC)20,22. The datasets are widely used in renewable energy application16,22, agricultural modelling of crop yields28, crop simulation exercises29, and plant disease modelling30.Furthermore, in order to assess the suitability of the NASA-POWER surface solar radiation products for the study area, a synthetic sunshine duration based Global solar radiation (GSR) is developed from the Angstrom-Prescott sunshine duration model by Eq. 3 for comparisons27.$$ {text{GSR}} = left[ {{text{a}} + {text{b}}frac{{text{n}}}{{text{N}}}} right]{text{H}}_{{text{o}}} $$
(3)
were Ho (kWhm−2 day−1) is the daily extraterrestrial solar radiation on an horizontal surface, n is the daily sunshine duration measurements obtained from the Ghana Meteorological Agency (GMet), and N is the maximum possible daily sunshine duration or the day length in hours determined by Eq. 1. Generalized regression constants a = 0.25 and b = 0.5 for the study area were determined by Asilevi27 from experimental radiometric data based on correlation regression analysis between atmospheric clarity index (GSR/Ho) and atmospheric cloudlessness index (n/N), for estimating solar radiation over the study area, and compared with other satellite data retrieved from the National Renewable Energy Laboratory (NREL) and the German Aerospace Centre (DLR)27. Ho was calculated from astronomical parameters by Eq. 4:$$ {text{H}}_{0} = frac{{24{ } cdot { }60}}{pi } cdot {text{G}}_{{{text{sc}}}} cdot {text{d}}_{{text{r}}} left[ {omega_{{text{s}}} sin varphi sin delta + cos varphi cos delta sin omega_{{text{s}}} } right] $$
(4)
where Gsc is the Solar constant in MJm−2 min−1, dr is the relative Earth–Sun distance in meters (m), (omega_{s}) is the sunset hour angle (angular distance between the meridian of the observer and the meridian whose plane contains the sun), (delta) is the angle of declination in degrees (°) and (varphi) is the local latitude. A detailed presentation of the calculation was published in a previous work27.Statistical assessment analysisFor the purpose of assessing the NASA-POWER derived monthly mean GSR (GSRn) datasets in comparison with the estimated Global Solar Radiation (GSRe) datasets used in this paper, the following deviation and correlation methods in Eqs. 5–11, each showing a complimentary result were used: Standard deviation (({upsigma })), residual error (RE), Root mean square error (RMSE), Mean bias error (MBE), Mean percentage error (MPE), Pearson’s correlation coefficient (r), and Willmott index of agreement (d) for n observations31,32,33,34,35. GSRe, GSRn, and RE represent the estimated GSR, NASA-POWER GSR, and the residual error between GSRe and GSRn respectively. A positive RE indicates that sunshine-based estimated GSR is larger than the NASA-POWER reanalysis dataset, while a negative RE indicates that sunshine-based estimated GSR is smaller than the NASA-POWER reanalysis dataset. The arithmetic mean of any dataset is µ.The standard deviation (({upsigma })) was used to check the upper and lower limits of distribution around the mean deviations between GSRe and GSRn in order to ascertain violations between both datasets33. The RMSE is a standard statistical metric to quantify error margins in meteorology and climate research studies, and by definition is always positive, representing zero in the ideal case, plus a smaller value signifying a good marginal deviation31. The MBE is a good indicator for under-or overestimation in observations, with MBE values closest to zero being desirable. The MPE further indicates the percentage deviation between the GSRe and GSRn individual datasets35.$$ {upsigma } = sqrt {frac{1}{{{text{n}} – 1}}mathop sum limits_{{{text{i}} = 1}}^{{text{n}}} left( {{text{GSR}} – {upmu }} right)^{2} } $$
(5)
$$ {text{RE}} = {text{GSR}}_{{text{e}}} – {text{GSR}}_{{text{n}}} $$
(6)
$$ {text{RMSE}} = sqrt {frac{1}{{text{n}}}mathop sum limits_{{{text{i}} = 1}}^{{text{n}}} left( {{text{RE}}} right)^{2} } $$
(7)
$$ {text{MBE}} = frac{1}{{text{n}}}mathop sum limits_{{{text{i}} = 1}}^{{text{n}}} left( {{text{RE}}} right) $$
(8)
$$ {text{MPE}} = frac{1}{{text{n}}}mathop sum limits_{{{text{i}} = 1}}^{{text{n}}} left( {frac{{{text{RE}}}}{{{text{GSR}}_{{text{e}}} }} times 100{text{% }}} right) $$
(9)
$$ {text{r}} = frac{{mathop sum nolimits_{{{text{i}} = 1}}^{{text{n}}} left( {{text{GSR}}_{{text{e}}} – {upsigma }_{{text{e}}} } right)left( {{text{GSR}}_{{text{n}}} – {upsigma }_{{text{n}}} } right)}}{{left( {{text{n}} – 1} right){upsigma }_{{text{e}}} {upsigma }_{{text{n}}} }} $$
(10)
$$ {text{d}} = 1 – left[ {frac{{mathop sum nolimits_{{{text{i}} = 1}}^{{text{n}}} left( {{text{GSR}}_{{text{e}}} – {text{GSR}}_{{text{n}}} } right)^{2} }}{{mathop sum nolimits_{{{text{i}} = 1}}^{{text{n}}} left( {left| {{text{GSR}}_{{text{e}}} – {text{GSR}}_{{{text{nave}}}} left| + right|{text{GSR}}_{{text{n}}} – {text{GSR}}_{{{text{nave}}}} } right|} right)^{2} }}} right] $$
(11)
Further, as with other statistical studies in meteorology36, the Pearson’s correlation coefficient (r) was used to quantify the strength of correlation between GSRe and GSRn. Finally, the Willmott index of agreement (d) commonly used in meteorological literature computed from Eq. 7 is used to assess the degree of GSRe/GSRn agreement34. More

Soil sample collectionSoil samples were collected from three different crude oil reservoirs et al. Faisaliyyah, Al Sina’iyah, and Ghubairah located in Riyadh, Saudi Arabia. Briefly, 400 g of soil samples were collected at 0–10 cm depth, under aseptic conditions. Samples were sieved by 2.5 mm pore size sieves, homogenized, and stored at 4ºC until use.Sources of different hydrocarbonsDifferent samples of crude oil, kerosene, diesel, and used oil were collected in sterile flasks from the tankers of Saudi Aramco Company (Dammam, Saudi Arabia). Additionally, another flask was prepared by mixing 1% of each oil in MSM liquid media to make up the mixed oil. The oil samples were sterilized by Millex® Syringe Filters (Merck Millipore co., Burlington, MA, United States) and stored at 4 °C for further usage.Isolation and identification of fungal speciesThe fungal species in the soil contaminated by crude oil were identified using the dilution method. Briefly, 10% of each soil sample was dissolved in distilled water and vortexed thoroughly. Then, 0.2 ml of each sample was cultured on a sterile PDA plate incubated at 28 °C for three days until the growth of different fungal colonies. Carefully, each colony was isolated, re-cultured on new PDA McCartney bottles of PDA slant, and incubated at 28 °C for three days. The fungi were identified microscopically using standard taxonomic keys based on typical mycelia growth and morphological characteristics provided in the mycological keys54. Besides, the taxonomy of the isolated yeast strains was confirmed by the API 20 C AUX kit (Biomerieux Corp., Marcy-l’Étoile, France) (data not shown). The morphology of pure cultures was tested and identified under a light microscope as described before55.The incidence of each strain was calculated as follows:$$ Incidence ;(% ) = frac{{{text{Number }};{text{of }};{text{samples }};{text{showed }};{text{microbial }};{text{growth}}}}{{{text{Total }};{text{samples}}}} times 100 $$Hydrocarbon tolerance testThe growth rate of isolated strains was tested in a liquid medium of MSM mixed with 1% of either crude oil, used oil, diesel, kerosene, or mixed oil. Furthermore, a control sample of MSM liquid medium without any of the oils tested and all culture media were autoclaved at 121 °C for 30 min. After cooling, 1 ml of each isolate was inoculated with one of the above mixtures and incubated at 25 °C on an orbital shaker. The growth rate was measured every three days for a month for each treatment versus the control. All experiments were performed in triplicates.Scanning electron microscopy (SEM)The morphology of different strains of the isolated fungi was tested by SEM, as previously described56, with some modifications. Briefly, 1 ml of each growing strain, in the liquid media, was centrifuged at the maximum speed (14,000 rpm) for 1 min, followed by fixation with 2.5% glutaraldehyde, and overnight incubation at 5 °C. Later, the sample was pelleted, washed with distilled water, then dehydrated with different ascending concentrations of ethanol (30, 50, 70, 90, 100 (v/v)) for 15 min at room temperature. Finally, samples were examined in the Prince Naif Research Centre (King Saud University, Riyadh, Saudi Arabia) by the JEOL JEM-2100 microscope (JEOL, Peabody, MA, United States), according to the manufacturer instructions.Crude oil degradation assayA modified version of the DCPIP assay57 was employed to assess the oil-degrading ability of the fungal isolates. For each strain, 100 ml of the autoclaved MSM was mixed with 1% (V/V) of one of the hydrocarbons (crude oil, used oil, diesel, kerosene, or mixed oil), 0.1% (v/v) of Tween 80, and 0.6 mg/mL of the redox indicator (DCPIP). Then, 1–2 ml of different fungi growing in liquid media (24–48 h) add to the Crude Oil Degradation media, prepared previously, and incubated for two weeks in a shaking incubator at 25 °C. All flasks were covered and protected from light, aeration, or temperature exchanges to reduce the effects of oil weathering (evaporation, photooxidation). The surfactant Tween 80 was used for bio-stimulation and acceleration of the biosurfactant production by increasing metabolism58. A non-inoculated Crude Oil Degradation media was used as the negative control. Afterward, the colorimetric analysis for the change in DCPIP color was estimated, spectrophotometrically, at 420 nm. All experiments were performed in triplicates.Preparation of cell-free supernatant (CFS)To prepare the Cell-Free Supernatant (CFS), all isolates were grown in MSM broth medium with 1% of either crude oil, used oil, diesel, kerosene, or mixed oil for 30 days in a shaking incubator at 25 °C. After incubation, the cells were removed by centrifugation at 10,000 rpm for 30 min at 4 °C. The supernatant (CFS) was collected and filter-sterilized with a 0.45 μm pore size sterile membrane. CFS was screened for the production of different biosurfactants. All the experiments were carried out in triplicates, and the average values were calculated.Drop-Collapse assayThe Drop-Collapse assay was performed as previously described9, with some modifications. 100 µl of crude oil was applied on glass slides, then 10 µl of each CFS was added to the center of the slide surface and incubated for a minute at room temperature. The slides were imaged by a light microscope using the 10X objective lenses. The spreading on the soil surface was scored by either « + » to indicate the level of positive spreading, biosurfactant production, or «—» for negative spreading. Biosurfactant production was considered positive at the drop diameter ≥ 0.5 mm, compared to the negative control (treated with distilled water).Oil spreading assayAn amount of 20 ml of water was added to the Petri plate (size of 100 mm) and mixed with 20 µl of crude oil or mixed oil, which created a thin layer on the water surface. Then, 10 µl of CFS was delivered onto the surface of the oil, and the clear zone surrounding the CFS drop was observed. The results were compared to the negative control (without CFS) and positive control of 1% SDS41. We have measured the clear zones diameter from images and calculate the actual values in regards to the diameter of the Petri dish (10 cm). The assay was performed in triplicates.Emulsification activity assayThe emulsification activity of each isolate was assessed by mixing equal volumes of MSM broth medium of each isolate with different oils in separate tubes. The samples were homogenized by vortex at high speed for two minutes at room temperature (25 °C) and allowed to settle for 24 h. The tests were performed in duplicate. Then, the emulsification index was calculated as follows59:$$ Emulsification; activity; left( % right) = frac{{{text{Height }};{text{of }};{text{emulsion }};{text{layer}}}}{{{text{Total }};{text{height}}}} times 100 $$Recovery of biosurfactantsThe recovery of biosurfactants from CFS was tested through different assays:Acid precipitation assay3 ml of each CFS was adjusted by 6 N HCl to pH 2 and incubated for 24 h at 4 °C. Later, equal volumes of chloroform/methanol mixture (2:1 v/v) were added to each tube, vortexed, and incubated overnight at room temperature. Afterward, the samples were centrifuged for 30 min at 10,000 rpm (4 °C), the precipitate (Light brown colored paste) was air-dried in a fume hood, and weighed53.Solvent extraction assayThe CFS containing biosurfactant was treated with a mixture of extraction solvents (equal volumes of methanol, chloroform, and acetone). Then, the new mixture was incubated in a shaking incubator at 200 rpm, 30 °C for 5 h. The precipitate was separated into two layers, in which the lower layer (White) was isolated, dried, weighed, and stored60.Ammonium sulfate precipitation assayThe CFS containing biosurfactant was precipitated with 40% (w/v) ammonium sulfate and incubated overnight at 4 °C. The samples were centrifuged at 10,000 rpm for 30 min (4 °C). The precipitate was collected and extracted with an amount of acetone equal to the volume of the supernatant. After centrifugation, the precipitate (Creamy-white) was isolated, air-dried in a fume hood, and weighed53.Zinc sulfate precipitation methodSimilarly, 40% (w/v) zinc sulfate was mixed with the CFS containing biosurfactant. Then, the mixture was incubated at 4 °C, overnight. The precipitate (Light Brown) was collected by centrifugation at 10,000 rpm for 30 min (4 °C), air-dried in a fume hood, and weighed53.Statistical analysisAll experiments were performed in triplicate, and the results were expressed as the mean values ± standard deviation (SD). One-way ANOVA and Dunnett’s tests were used to estimate the significance levels at P  More

Birenboim, A. The influence of urban environments on our subjective momentary experiences. Environ. Plan. B-Urban Anal. CIty Sci. 45, 915–932. https://doi.org/10.1177/2399808317690149 (2018).Article 

Google Scholar 
Flores, A., Pickett, S. T. A., Zipperer, W. C., Pouyat, R. V. & Pirani, R. Adopting a modern ecological view of the metropolitan landscape: The case of a greenspace system for the New York City region. Landsc. Urban Plan. 39, 295–308. https://doi.org/10.1016/S0169-2046(97)00084-4 (1998).Article 

Google Scholar 
Weijs-Perrée, M., Dane, G., Berg, P. V. D. & Dorst, M. V. A multi-level path analysis of the relationships between the momentary experience characteristics, satisfaction with urban public spaces, and momentary- and long-term subjective wellbeing. Int. J. Environ. Res. Public Health. https://doi.org/10.3390/ijerph16193621 (2019).Article 
PubMed 
PubMed Central 

Google Scholar 
Paulin, M. J. et al. Application of the natural capital model to assess changes in ecosystem services from changes in green infrastructure in Amsterdam. Ecosyst. Serv. 43, 101114. https://doi.org/10.1016/j.ecoser.2020.101114 (2020).Article 

Google Scholar 
Derkzen, M. L., van Teeffelen, A. J. A., Verburg, P. H. & Diamond, S. Quantifying urban ecosystem services based on high-resolution data of urban green space: An assessment for Rotterdam, the Netherlands. J. Appl. Ecol. 52, 1020–1032. https://doi.org/10.1111/1365-2664.12469 (2015).Article 

Google Scholar 
Leiva, M. A., Santibanez, D. A., Ibarra, S., Matus, P. & Seguel, R. A five-year study of particulate matter (PM2.5) and cerebrovascular diseases. Environ. Pollut. 181, 1–6. https://doi.org/10.1016/j.envpol.2013.05.057 (2013).CAS 
Article 

Google Scholar 
Venkataramanan, V. et al. Knowledge, attitudes, intentions, and behavior related to green infrastructure for flood management: A systematic literature review. Sci. Total Environ. 720, 137606. https://doi.org/10.1016/j.scitotenv.2020.137606 (2020).CAS 
Article 
PubMed 

Google Scholar 
Wang, G. Z., Han, Q. & De Vries, B. The multi-objective spatial optimization of urban land use based on low-carbon city planning. Ecol. Indic. 125, 107540. https://doi.org/10.1016/j.ecolind.2021.107540 (2021).CAS 
Article 

Google Scholar 
Cameron, R. W. F. et al. The domestic garden—Its contribution to urban green infrastructure. Urban For. Urban Green. 11, 129–137. https://doi.org/10.1016/j.ufug.2012.01.002 (2012).Article 

Google Scholar 
De la Sota, C., Ruffato-Ferreira, V. J., Ruiz-Garcia, L. & Alvarez, S. Urban green infrastructure as a strategy of climate change mitigation. A case study in northern Spain. Urban For. Urban Green. 40, 145–151. https://doi.org/10.1016/j.ufug.2018.09.004 (2019).Article 

Google Scholar 
Pongsakorn, S., Jiang, X. R. & Sullivan, W. C. Green infrastructure, green stormwater infrastructure, and human health a review. Curr. Landscape. Ecol. Rep. 2, 96–110. https://doi.org/10.1007/s40823-017-0028-y (2017).Article 

Google Scholar 
Liu, O. Y. & Russo, A. Assessing the contribution of urban green spaces in green infrastructure strategy planning for urban ecosystem conditions and services (Sust. Cities Soc., 2021). https://doi.org/10.1016/j.scs.2021.102772.Book 

Google Scholar 
McMahon, E. T. Green infrastructure. Plan. Commission. J. (2000).Mell, I. C. Green Infrastructure Concepts, Perceptions and Its Use in Spatial Planning. Doctor of Philosophy Thesis (Planning and Landscape Newcastle University, 2010).
Google Scholar 
Wang, J. X. & Banzhaf, E. Towards a better understanding of green infrastructure: A critical review. Ecol. Indic. 85, 758–772. https://doi.org/10.1016/j.ecolind.2017.09.018 (2018).Article 

Google Scholar 
Young, R., Zanders, J., Lieberknecht, K. & Fassman-Beck, E. A comprehensive typology for mainstreaming urban green infrastructure. J. Hydrol. 519, 2571–2583. https://doi.org/10.1016/j.jhydrol.2014.05.048 (2014).Article 

Google Scholar 
Wang, J. X., Xu, C., Pauleit, S., Kindler, A. & Banzhaf, E. Spatial patterns of urban green infrastructure for equity: A novel exploration. J. Clean Prod. 238, 117858. https://doi.org/10.1016/j.jclepro.2019.117858 (2019).Article 

Google Scholar 
Cook, E. A. Landscape structure indices for assessing urban ecological networks. Landsc. Urban Plan. 58, 269–280 (2002).Article 

Google Scholar 
Vogt, P. & Riitters, K. GuidosToolbox: Universal digital image object analysis. Eur. J. Remote Sens. 50, 352–361. https://doi.org/10.1080/22797254.2017.1330650 (2017).Article 

Google Scholar 
Vogt, P., Riitters, K. H., Estreguil, C., Kozak, J. & Wade, T. G. Mapping spatial patterns with morphological image processing. Landsc. Ecol. 22, 171–177. https://doi.org/10.1007/s10980-006-9013-2 (2007).Article 

Google Scholar 
Kuttner, M., Hainz-Renetzeder, C., Hermann, A. & Wrbka, T. Borders without barriers—Structural functionality and green infrastructure in the Austrian-Hungarian transboundary region of Lake Neusiedl. Ecol. Indic. 31, 59–72. https://doi.org/10.1016/j.ecolind.2012.04.014 (2013).Article 

Google Scholar 
Ma, Q. W., Li, Y. H. & Xu, L. H. Identification of green infrastructure networks based on ecosystem services in a rapidly urbanizing area. J. Clean Prod. 300, 126945. https://doi.org/10.1016/j.jclepro.2021.126945 (2021).Article 

Google Scholar 
Furberg, D., Ban, Y. & Mörtberg, U. Monitoring urban green infrastructure changes and impact on habitat connectivity using high-resolution satellite data. Remote Sens. 12, 3072. https://doi.org/10.3390/rs12183072 (2020).Article 

Google Scholar 
Barbati, A., Corona, P., Salvati, L. & Gasparella, L. Natural forest expansion into suburban countryside: Gained ground for a green infrastructure?. Urban For. Urban Green. 12, 36–43. https://doi.org/10.1016/j.ufug.2012.11 (2013).Article 

Google Scholar 
Fluhrer, T., Chapa, F. & Hack, J. A methodology for assessing the implementation potential for retrofitted and multifunctional urban green infrastructure in public areas of the global south. Sustainability https://doi.org/10.3390/su13010384 (2021).Article 

Google Scholar 
Carroll, C., McRae, B. H. & Brookes, A. Use of linkage mapping and centrality analysis across habitat gradients to conserve connectivity of gray wolf populations in western North America. Conserv. Biol. 26, 78–87. https://doi.org/10.1111/j.1523-1739.2011.01753.x (2012).Article 
PubMed 

Google Scholar 
Saura, S. & Torne, J. Conefor Sensinode 2.2: A software package for quantifying the importance of habitat patches for landscape connectivity. Environ. Modell. Softw. 24, 135–139 (2009).Article 

Google Scholar 
Jaworek-Jakubska, J., Filipiak, M., Michalski, A. & Napierała-Filipiak, A. Spatio-temporal changes of urban forests and planning evolution in a highly dynamical urban area: The case study of Wrocław, Poland. Forests 11, 17. https://doi.org/10.3390/f11010017 (2019).Article 

Google Scholar 
Ren, Z. B., He, X. Y., Zheng, H. F. & Wei, H. X. Spatio-temporal patterns of urban forest basal area under China’s rapid urban expansion and greening: Implications for urban green infrastructure management. Forests 9, 272. https://doi.org/10.3390/f9050272 (2018).Article 

Google Scholar 
Elliott, R. M. et al. Identifying linkages between urban green infrastructure and ecosystem services using an expert opinion methodology. Ambio 49, 569–583. https://doi.org/10.1007/s13280-019-01223-9 (2019).Article 
PubMed 
PubMed Central 

Google Scholar 
García, A. M., Santé, I., Loureiro, X. & Miranda, D. Green infrastructure spatial planning considering ecosystem services assessment and trade-off analysis. Application at landscape scale in Galicia region (NW Spain). Ecosyst. Serv. 43, 101115. https://doi.org/10.1016/j.ecoser.2020.101115 (2020).Article 

Google Scholar 
Tiwari, A. & Kumar, P. Integrated dispersion-deposition modelling for air pollutant reduction via green infrastructure at an urban scale. Sci. Total Environ. 723, 138078. https://doi.org/10.1016/j.scitotenv.2020.138078 (2020).CAS 
Article 
PubMed 

Google Scholar 
Zhang, Y. Q. et al. Unexpected air quality impacts from implementation of green infrastructure in urban environments: A Kansas City case study. Sci. Total Environ. 744, 140960. https://doi.org/10.1016/j.scitotenv.2020.140960 (2020).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Alizadehtazi, B., Gurian, P. L. & Montalto, F. A. Observed variability in soil moisture in engineered urban green infrastructure systems and linkages to ecosystem services. J. Hydrol. 590, 125381. https://doi.org/10.1016/j.jhydrol.2020.125381 (2020).Article 

Google Scholar 
Dennis, M., Cook, P. A., James, P., Wheater, C. P. & Lindley, S. J. Relationships between health outcomes in older populations and urban green infrastructure size, quality and proximity. BMC Public Health https://doi.org/10.1186/s12889-020-08762-x (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Van Oijstaeijen, W., Van Passel, S. & Cools, J. Urban green infrastructure: A review on valuation toolkits from an urban planning perspective. J. Environ. Manag. 267, 110603. https://doi.org/10.1016/j.jenvman.2020.110603 (2020).Article 

Google Scholar 
Majekodunmi, M., Emmanuel, R. & Jafry, T. A spatial exploration of deprivation and green infrastructure ecosystem services within Glasgow city. Urban For. Urban Green. 52, 126698. https://doi.org/10.1016/j.ufug.2020.126698 (2020).Article 

Google Scholar 
Liberalesso, T., Oliveira Cruz, C., Matos Silva, C. & Manso, M. Green infrastructure and public policies: An international review of green roofs and green walls incentives. Land Use Pol. 96, 104693. https://doi.org/10.1016/j.landusepol.2020.104693 (2020).Article 

Google Scholar 
Lin, H. Y., Qian, J., Yan, L. J. & Huang, S. R. Analysis of spatial-temporal pattern and scenario simulation of green infrastructure in Wuyi County based on morphological spatial pattern analysis and CA-Markov model. Acta Agricult. Zhejiangensis. https://doi.org/10.3969/j.issn.1004-1524.2019.07.21 (2019).Article 

Google Scholar 
Mitsova, D., Shuster, W. & Wang, X. H. A cellular automata model of land cover change to integrate urban growth with open space conservation. Landsc. Urban Plan. 99, 141–153. https://doi.org/10.1016/j.landurbplan.2010.10.001 (2011).Article 

Google Scholar 
Dennis, M. et al. Mapping urban green infrastructure: A novel landscape-based approach to incorporating land use and land cover in the mapping of human-dominated systems. Land 7, 17. https://doi.org/10.3390/land7010017 (2018).Article 

Google Scholar 
Hu, Y. J. et al. Urban expansion and farmland loss in Beijing during 1980–2015. Sustainability 10, 3927. https://doi.org/10.3390/su10113927 (2018).Article 

Google Scholar 
Li, W. J., Wang, Y., Xie, S. Y., Sun, R. H. & Cheng, X. Impacts of landscape multifunctionality change on landscape ecological risk in a megacity, China: A case study of Beijing. Ecol. Indic. 117 (2020).Song, W., Pijanowski, B. C. & Tayyebi, A. Urban expansion and its consumption of high-quality farmland in Beijing, China. Ecol. Indic. 54, 60–70. https://doi.org/10.1016/j.ecolind.2015.02.015 (2015).Article 

Google Scholar 
Li, Z. Z., Cheng, X. Q. & Han, H. R. Future impacts of land use change on ecosystem services under different scenarios in the ecological conservation area, Beijing, China. Forests https://doi.org/10.3390/f11050584 (2020).Article 

Google Scholar 
Liu, D. Y. et al. Interoperable scenario simulation of land-use policy for Beijing-Tianjin-Hebei region, China. Land Use Pol. 75, 155–165. https://doi.org/10.1016/j.landusepol.2018.03.040 (2018).Article 

Google Scholar 
Mo, W. B., Wang, Y., Zhang, Y. X. & Zhuang, D. F. Impacts of road network expansion on landscape ecological risk in a megacity, China: A case study of Beijing. Sci. Total Environ. 574, 1000–1011. https://doi.org/10.1016/j.scitotenv.2016.09.048 (2017).CAS 
Article 
PubMed 

Google Scholar 
Melgani, F. & Bruzzone, L. Classification of hyperspectral remote sensing images with support vector machines. IEEE Trans. Geosci. Remote Sens. 42, 1778–1790. https://doi.org/10.1109/Tgrs.2004.831865 (2004).Article 

Google Scholar 
Zhang, C., Wang, T. J., Atkinson, P. M., Pan, X. & Li, H. P. A novel multi-parameter support vector machine for image classification. Int. J. Remote Sens. 36, 1890–1906. https://doi.org/10.1080/01431161.2015.1029096 (2015).CAS 
Article 

Google Scholar 
Peterson, L. K., Bergen, K. M., Brown, D. G., Vashchuk, L. & Blam, Y. Forested land-cover patterns and trends over changing forest management eras in the Siberian Baikal region. For. Ecol. Manag. 257, 911–922. https://doi.org/10.1016/j.foreco.2008.10.037 (2009).Article 

Google Scholar 
Sang, L. L., Zhang, C., Yang, J. Y., Zhu, D. H. & Yun, W. J. Simulation of land use spatial pattern of towns and villages based on CA-Markov model. Math. Comput. Model. 54, 938–943. https://doi.org/10.1016/j.mcm.2010.11.019 (2011).Article 

Google Scholar 
Liu, D. Y., Zheng, X. Q. & Wang, H. B. Land-use Simulation and Decision-Support system (LandSDS): Seamlessly integrating system dynamics, agent-based model, and cellular automata. Ecol. Model. 417, 108924. https://doi.org/10.1016/j.ecolmodel.2019.108924 (2020).Article 

Google Scholar 
Kazak, J. K. The use of a decision support system for sustainable urbanization and thermal comfort in adaptation to climate change actions-The case of the Wroclaw larger urban zone (Poland). Sustainability https://doi.org/10.3390/su10041083 (2013).Article 

Google Scholar 
Sonnenberg, F. A. & Beck, J. R. Markov-models in medical decision-making—A practical guide. Med. Decis. Mak. 13, 322–338. https://doi.org/10.1177/0272989×9301300409 (1993).CAS 
Article 

Google Scholar 
Nadoushan, M. A., Soffianian, A. & Alebrahim, A. Modeling land use/cover changes by the combination of Markov chain and cellular automata Markov CA-Markov models. Int. J. Environ. Health Res. https://doi.org/10.4103/WKMP-0092.159922 (2015).Article 

Google Scholar 
Mansour, S., Al-Belushi, M. & Al-Awadhi, T. Monitoring land use and land cover changes in the mountainous cities of Oman using GIS and CA-Markov modelling techniques. Land Use Pol. 91, 104414. https://doi.org/10.1016/j.landusepol.2019.104414 (2020).Article 

Google Scholar 
Karimi, H., Jafarnezhad, J., Khaledi, J. & Ahmadi, P. Monitoring and prediction of land use/land cover changes using CA-Markov model: A case study of Ravansar County in Iran. Arab. J. Geosci. https://doi.org/10.1007/s12517-018-3940-5 (2018).Article 

Google Scholar 
Mondal, M. S., Sharma, N. C. P. K. G. & Kappas, M. Statistical independence test and validation of CA Markov land use land cover (LULC) prediction results. Egypt. J. Remote Sens. Space Sci. https://doi.org/10.1016/j.ejrs.2016.08.001 (2016).Article 

Google Scholar 
Liu, Q. et al. Multi-scenario simulation of land use change and its eco-environmental effect in Hainan Island based on CA-Markov model. Ecol. Environ. Sci. 30, 1522–1531. https://doi.org/10.16258/j.cnki.1674-5906.2021.07.021 (2021).Article 

Google Scholar 
Pontius, R. G. Statistical methods to partition effects of quantity and location during comparison of categorical maps at multiple resolutions. Photogramm. Eng. Remote Sens. 68, 1041–1049 (2002).
Google Scholar 
Soille, P. & Vogt, P. Morphological segmentation of binary patterns. Pattern Recognit. Lett. 30, 456–459 (2009).Article 

Google Scholar 
Chang, Q., Liu, X. W., Wu, J. S. & He, P. MSPA-based urban green infrastructure planning and management approach for urban sustainability: Case study of Longgang in China. J. Urban Plan. Dev. https://doi.org/10.1061/(asce)up.1943-5444.0000247 (2015).Article 

Google Scholar 
Li, K. M. et al. Spatiotemporal evolution characteristics of urban green infrastructure in central Liaoning urban agglomeration during the past 20 years based on landscape ecology and morphology. Acta Ecol. Sin. https://doi.org/10.5846/stxb202007221918 (2021).Article 

Google Scholar 
Ning, J. et al. Spatiotemporal patterns and characteristics of land-use change in China during 2010–2015. J. Geogr. Sci. 28, 547–562. https://doi.org/10.1007/s11442-018-1490-0 (2018).Article 

Google Scholar 
Sawyer, S. C., Epps, C. W. & Brashares, J. S. Placing linkages among fragmented habitats: Do least-cost models reflect how animals use landscapes?. J. Appl. Ecol. 48, 668–678. https://doi.org/10.1111/j.1365-2664.2011.01970.x (2011).Article 

Google Scholar 
Yin, G. Y., Liu, L. M. & Jiang, X. L. The sustainable arable land use pattern under the tradeoff of agricultural production, economic development, and ecological protection—An analysis of Dongting Lake basin, China. Environ. Sci. Pollut. Res. 24, 25329–25345. https://doi.org/10.1007/s11356-017-0132-x (2017).Article 

Google Scholar  More

Shiklomanov, I. A. & Rodda, J. C. World water resources at the beginning of the twenty-first century. (Cambridge University Press, 2003).Biggs, J., von Fumetti, S. & Kelly-Quinn, M. The importance of small waterbodies for biodiversity and ecosystem services: implications for policy makers. Hydrobiologia 793, 3–39 (2017).Article 

Google Scholar 
Heino, J. et al. Lakes in the era of global change: moving beyond single-lake thinking in maintaining biodiversity and ecosystem services. Biol. Rev. 96, 89–106 (2021).PubMed 
Article 

Google Scholar 
Janssen, A. B. G. et al. Shifting states, shifting services: linking regime shifts to changes in ecosystem services of shallow lakes. Freshw. Biol. 66, 1–12 (2021).Article 

Google Scholar 
Knoll, L. B. et al. Consequences of lake and river ice loss on cultural ecosystem services. Limnol. Oceanogr. Lett. 4, 119–131 (2019).Article 

Google Scholar 
Sterner, R. W. et al. Ecosystem services of Earth’s largest freshwater lakes. Ecosyst. Serv. 41, 101046 (2020).Article 

Google Scholar 
Reynaud, A. & Lanzanova, D. A global meta-analysis of the value of ecosystem services provided by lakes. Ecol. Econ. 137, 184–194 (2017).PubMed 
PubMed Central 
Article 

Google Scholar 
Cooley, S. W., Ryan, J. C. & Smith, L. C. Human alteration of global surface water storage variability. Nature 591, 78–81 (2021).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Downing, J. A. Global limnology: up-scaling aquatic services and processes to planet Earth. SIL Proceedings, 1922–2010 30, 1149–1166 (2009).Article 

Google Scholar 
Tranvik, L. J., Cole, J. J. & Prairie, Y. T. The study of carbon in inland waters—from isolated ecosystems to players in the global carbon cycle. Limnol. Oceanogr. Lett. 3, 41–48 (2018).Article 

Google Scholar 
Balsamo, G. et al. On the contribution of lakes in predicting near-surface temperature in a global weather forecasting model. Tellus A Dyn. Meteorol. Oceanogr. 64, 15829 (2012).Article 

Google Scholar 
DelSontro, T., Beaulieu, J. J. & Downing, J. A. Greenhouse gas emissions from lakes and impoundments: upscaling in the face of global change. Limnol. Oceanogr. Lett. 3, 64–75 (2018).CAS 
Article 

Google Scholar 
Beaulieu, J. J. et al. Methane and carbon dioxide emissions from reservoirs: controls and upscaling. J. Geophys. Res. Biogeosciences 125, e2019JG005474 (2020).ADS 
CAS 
Article 

Google Scholar 
Slater, J. A. et al. The SRTM data “finishing” process and products. Photogramm. Eng. Remote Sens. 72, 237–247 (2006).Article 

Google Scholar 
Pekel, J.-F., Cottam, A., Gorelick, N. & Belward, A. S. High-resolution mapping of global surface water and its long-term changes. Nature 540, 418–422 (2016).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Verpoorter, C., Kutser, T., Seekell, D. A. & Tranvik, L. J. A global inventory of lakes based on high-resolution satellite imagery. Geophys. Res. Lett. 41, 6396–6402 (2014).ADS 
Article 

Google Scholar 
Pickens, A. H. et al. Mapping and sampling to characterize global inland water dynamics from 1999 to 2018 with full Landsat time-series. Remote Sens. Environ. 243, 111792 (2020).ADS 
Article 

Google Scholar 
Messager, M. L., Lehner, B., Grill, G., Nedeva, I. & Schmitt, O. Estimating the volume and age of water stored in global lakes using a geo-statistical approach. Nat. Commun. 7, 13603 (2016).ADS 
CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Tickner, D. et al. Bending the curve of global freshwater biodiversity loss: an emergency recovery plan. Bioscience 70, 330–342 (2020).PubMed 
PubMed Central 
Article 

Google Scholar 
Downing, J. A., Polasky, S., Olmstead, S. M. & Newbold, S. C. Protecting local water quality has global benefits. Nat. Commun. 12, 1–6 (2021).Article 
CAS 

Google Scholar 
Hill, R. A., Weber, M. H., Debbout, R. M., Leibowitz, S. G. & Olsen, A. R. The Lake-Catchment (LakeCat) Dataset: characterizing landscape features for lake basins within the conterminous USA. Freshw. Sci. 37, 208–221 (2018).PubMed 
PubMed Central 
Article 

Google Scholar 
Soranno, P. A. et al. LAGOS-NE: a multi-scaled geospatial and temporal database of lake ecological context and water quality for thousands of US lakes. Gigascience 6, 1–22 (2017).PubMed 
PubMed Central 
Article 

Google Scholar 
Toptunova, O., Choulga, M. & Kurzeneva, E. Status and progress in global lake database developments. Adv. Sci. Res. 16, 57–61 (2019).Article 

Google Scholar 
Meyer, M. F., Labou, S. G., Cramer, A. N., Brousil, M. R. & Luff, B. T. The global lake area, climate, and population dataset. Sci. Data 7, 174 (2020).PubMed 
PubMed Central 
Article 

Google Scholar 
Kling, G. W., Kipphut, G. W., Miller, M. M. & O’Brien, W. J. Integration of lakes and streams in a landscape perspective: the importance of material processing on spatial patterns and temporal coherence. Freshw. Biol. 43, 477–497 (2000).Article 

Google Scholar 
Fergus, C. E. et al. The freshwater landscape: lake, wetland, and stream abundance and connectivity at macroscales. Ecosphere 8, e01911 (2017).Article 

Google Scholar 
Lehner, B., Messager, ML., Korver, MC. & Linke, S. LakeATLAS Version 1.0, figshare, https://doi.org/10.6084/m9.figshare.19312001 (2022).Linke, S. et al. Global hydro-environmental sub-basin and river reach characteristics at high spatial resolution. Sci. data 6, 283 (2019).PubMed 
PubMed Central 
Article 

Google Scholar 
Fergus, C. E. et al. National framework for ranking lakes by potential for anthropogenic hydro-alteration. Ecol. Indic. 122, 107241 (2021).Article 

Google Scholar 
Bracht-Flyr, B., Istanbulluoglu, E. & Fritz, S. A hydro-climatological lake classification model and its evaluation using global data. J. Hydrol. 486, 376–383 (2013).ADS 
Article 

Google Scholar 
Soranno, P. A. et al. Using landscape limnology to classify freshwater ecosystems for multi-ecosystem management and conservation. Bioscience 60, 440–454 (2010).Article 

Google Scholar 
McCullough, I. M., Skaff, N. K., Soranno, P. A. & Cheruvelil, K. S. No lake left behind: how well do U.S. protected areas meet lake conservation targets? Limnol. Oceanogr. Lett. 4, 183–192 (2019).Article 

Google Scholar 
Stanley, E. H. et al. Biases in lake water quality sampling and implications for macroscale research. Limnol. Oceanogr. 64, 1572–1585 (2019).ADS 
CAS 
Article 

Google Scholar 
Hanson, P. C., Weathers, K. C. & Kratz, T. K. Networked lake science: how the Global Lake Ecological Observatory Network (GLEON) works to understand, predict, and communicate lake ecosystem response to global change. Inl. Waters 6, 543–554 (2016).Article 

Google Scholar 
Lottig, N. R. & Carpenter, S. R. Interpolating and forecasting lake characteristics using long-term monitoring data. Limnol. Oceanogr. 57, 1113–1125 (2012).ADS 
Article 

Google Scholar 
Filazzola, A. et al. A database of chlorophyll and water chemistry in freshwater lakes. Sci. Data 2020 71 7, 1–10 (2020).
Google Scholar 
Lehner, B. & Messager, M. L. HydroLAKES – Technical Documentation Version 1.0. https://data.hydrosheds.org/file/technical-documentation/HydroLAKES_TechDoc_v10.pdf (2016).Natural Resources Canada. CanVec Hydrography: Waterbody Features. Version 12.0. https://ftp.maps.canada.ca/pub/nrcan_rncan/vector/canvec (2013).Lehner, B., Verdin, K. & Jarvis, A. New global hydrography derived from spaceborne elevation data. Eos, Trans. AGU 89, 93–94 (2008).ADS 
Article 

Google Scholar 
Farr, T. G. & Kobrick, M. Shuttle radar topography mission produces a wealth of data. Eos, Trans. AGU 81, 583–585 (2000).ADS 
Article 

Google Scholar 
Müller Schmied, H. et al. The global water resources and use model WaterGAP v2.2d: model description and evaluation. Geosci. Model Dev. 14, 1037–1079 (2021).ADS 
Article 

Google Scholar 
Beck, H. E. et al. Global evaluation of runoff from 10 state-of-the-art hydrological models. Hydrol. Earth Syst. Sci. 21, 2881–2903 (2017).ADS 
Article 

Google Scholar 
Alcamo, J. et al. Development and testing of the WaterGAP 2 global model of water use and availability. Hydrol. Sci. J. 48, 317–338 (2003).Article 

Google Scholar 
Döll, P., Kaspar, F. & Lehner, B. A global hydrological model for deriving water availability indicators: model tuning and validation. J. Hydrol. 270, 105–134 (2003).ADS 
Article 

Google Scholar 
Lehner, B. & Grill, G. Global river hydrography and network routing: baseline data and new approaches to study the world’s large river systems. Hydrol. Process. 27, 2171–2186 (2013).ADS 
Article 

Google Scholar 
Fick, S. E. & Hijmans, R. J. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).Article 

Google Scholar 
Hengl, T. et al. SoilGrids250m: Global gridded soil information based on machine learning. PLoS One 12, e0169748 (2017).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Zhang, X. et al. GLC_FCS30: Global land-cover product with fine classification system at 30 m using time-series Landsat imagery. Earth Syst. Sci. Data 13, 2753–2776 (2021).ADS 
Article 

Google Scholar 
Buchhorn, M. et al. Copernicus Global Land Service: Land Cover 100m: Collection 3: epoch 2019: Globe, Zenodo, https://doi.org/10.5281/zenodo.3939050 (2020).ESRI. ArcGIS Desktop: Release 10.4.1 (Environmental Systems Research Institute, Redlands, CA, USA, 2016).Soranno, P. A., Cheruvelil, K. S., Wagner, T., Webster, K. E. & Bremigan, M. T. Effects of land use on lake nutrients: the importance of scale, hydrologic connectivity, and region. PLoS One 10, e0135454 (2015).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Su, Z. H., Lin, C., Ma, R. H., Luo, J. H. & Liang, Q. O. Effect of land use change on lake water quality in different buffer zones. Appl. Ecol. Environ. Res. 13, 639–653 (2015).
Google Scholar 
Brakebill, J. W., Schwarz, G. E. & Wieczorek, M. E. An enhanced hydrologic stream network based on the NHDPlus medium resolution dataset. Scientific Investigations Report https://doi.org/10.3133/sir20195127 (2020).Carroll, M., Townshend, J., DiMiceli, C., Noojipady, P. & Sohlberg, R. Global raster water mask at 250 meter spatial resolution, Collection 5: MOD44W MODIS Water Mask. College Park, Maryland: University of Maryland (2009).Carroll, M. L., Townshend, J. R., DiMiceli, C. M., Noojipady, P. & Sohlberg, R. A. A new global raster water mask at 250 m resolution. Int. J. Digit. Earth 2, 291–308 (2009).ADS 
Article 

Google Scholar 
European Environment Agency (EEA). European Catchments and Rivers Network System (ECRINS), https://www.eea.europa.eu/data-and-maps/data/european-catchments-and-rivers-network (2012).Ouellet Dallaire, C., Lehner, B., Sayre, R. & Thieme, M. A multidisciplinary framework to derive global river reach classifications at high spatial resolution. Environ. Res. Lett. 14, 024003 (2019).ADS 
Article 

Google Scholar 
Global Runoff Data Centre (GRDC). River discharge data. Federal Institute of Hydrology, 56068 Koblenz, Germany, https://www.bafg.de/GRDC (2014).Openshaw, S. The modifiable areal unit problem. In Quantitative Geography: A British View (eds. Wrigley, N. & Bennett, R.) 60–69 (Routledge and Kegan Paul, Andover, 1981).United States Census Bureau. 2010 Census. ftp://ftp2.census.gov/geo/tiger (2010).Center for International Earth Science Information Network (CIESIN) & NASA Socioeconomic Data and Applications Center (SEDAC). Gridded Population of the World, Version 4 (GPWv4): Population Count and Density. https://doi.org/10.7927/H4JW8BX5 (2016).Grill, G. et al. Mapping the world’s free-flowing rivers. Nature 569, 215–221 (2019).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Allen, D. J. et al. The Diversity of Life in African Freshwaters: Under Water, Under Threat: an Analysis of the Status and Distribution of Freshwater Species Throughout Mainland Africa. (IUCN, 2011).Markovic, D. et al. Europe’s freshwater biodiversity under climate change: distribution shifts and conservation needs. Divers. Distrib. 20, 1097–1107 (2014).Article 

Google Scholar 
Fluet-Chouinard, E., Lehner, B., Rebelo, L.-M., Papa, F. & Hamilton, S. K. Development of a global inundation map at high spatial resolution from topographic downscaling of coarse-scale remote sensing data. Remote Sens. Environ. 158, 348–361 (2015).ADS 
Article 

Google Scholar 
Lehner, B. et al. High‐resolution mapping of the world’s reservoirs and dams for sustainable river‐flow management. Front. Ecol. Environ. 9, 494–502 (2011).Article 

Google Scholar 
Fan, Y., Li, H. & Miguez-Macho, G. Global patterns of groundwater table depth. Science 339, 940–943 (2013).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Robinson, N., Regetz, J. & Guralnick, R. P. EarthEnv-DEM90: A nearly-global, void-free, multi-scale smoothed, 90m digital elevation model from fused ASTER and SRTM data. ISPRS J. Photogramm. Remote Sens. 87, 57–67 (2014).ADS 
Article 

Google Scholar 
Metzger, M. J. et al. A high-resolution bioclimate map of the world: a unifying framework for global biodiversity research and monitoring. Glob. Ecol. Biogeogr. 22, 630–638 (2013).Article 

Google Scholar 
Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G. & Jarvis, A. Very high resolution interpolated climate surfaces for global land areas. Int. J. Climatol. 25, 1965–1978 (2005).Article 

Google Scholar 
Zomer, R. J., Trabucco, A., Bossio, D. A. & Verchot, L. V. Climate change mitigation: a spatial analysis of global land suitability for clean development mechanism afforestation and reforestation. Agric. Ecosyst. Environ. 126, 67–80 (2008).Article 

Google Scholar 
Trabucco, A., Zomer, R. J., Bossio, D. A., van Straaten, O. & Verchot, L. V. Climate change mitigation through afforestation/reforestation: a global analysis of hydrologic impacts with four case studies. Agric. Ecosyst. Environ. 126, 81–97 (2008).Article 

Google Scholar 
Trabucco, A. & Zomer, R. J. Global soil water balance geospatial database. CGIAR Consortium for Spatial Information, https://cgiarcsi.community/data/global-high-resolution-soil-water-balance (2010).Hall, D. K., Riggs, G. A. & Salomonson, V. MODIS/Terra snow cover daily L3 global 500m grid, version 5, 2002–2015, https://doi.org/10.5067/MODIS/MOD10A1.006 (2016).Bartholomé, E. & Belward, A. S. GLC2000: a new approach to global land cover mapping from Earth observation data. Int. J. Remote Sens. 26, 1959–1977 (2005).Article 

Google Scholar 
Ramankutty, N. & Foley, J. A. Estimating historical changes in global land cover: Croplands from 1700 to 1992. Global Biogeochem. Cycles 13, 997–1027 (1999).ADS 
CAS 
Article 

Google Scholar 
Lehner, B. & Döll, P. Development and validation of a global database of lakes, reservoirs and wetlands. J. Hydrol. 296, 1–22 (2004).ADS 
Article 

Google Scholar 
Ramankutty, N., Evan, A. T., Monfreda, C. & Foley, J. A. Farming the planet: 1. Geographic distribution of global agricultural lands in the year 2000. Global Biogeochem. Cycles 22, (2008).Siebert, S. et al. A global data set of the extent of irrigated land from 1900 to 2005. Hydrol. Earth Syst. Sci. 19, 1521–1545 (2015).ADS 
Article 

Google Scholar 
GLIMS & NSIDC. Global land ice measurements from space (GLIMS) glacier database, v1. National Snow and Ice Data Center (NSIDC), https://doi.org/10.7265/N5V98602 (2012).Gruber, S. Derivation and analysis of a high-resolution estimate of global permafrost zonation. Cryosphere 6, 221–233 (2012).ADS 
Article 

Google Scholar 
UNEP-WCMC & IUCN. The World Database on Protected Areas, http://www.protectedplanet.net (2014).Dinerstein, E. et al. An ecoregion-based approach to protecting half the terrestrial realm. Bioscience 67, 534–545 (2017).PubMed 
PubMed Central 
Article 

Google Scholar 
Abell, R. et al. Freshwater ecoregions of the world: a new map of biogeographic units for freshwater biodiversity conservation. Bioscience 58, 403–414 (2008).Article 

Google Scholar 
Hengl, T. et al. SoilGrids1km—global soil information based on automated mapping. PLoS One 9, e105992 (2014).ADS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Hartmann, J. & Moosdorf, N. The new global lithological map database GLiM: a representation of rock properties at the Earth surface. Geochem. Geophys. Geosyst. 13, Q12004 (2012).ADS 
Article 

Google Scholar 
Williams, P. W. & Ford, D. C. Global distribution of carbonate rocks. Zeitschrift für Geomorphologie Suppl. 147, 1–2 (2006).
Google Scholar 
Borrelli, P. et al. An assessment of the global impact of 21st century land use change on soil erosion. Nat. Commun. 8, 1–13 (2017).ADS 
CAS 
Article 

Google Scholar 
Pesaresi, M. & Freire, S. GHS Settlement grid following the REGIO model 2014 in application to GHSL Landsat and CIESIN GPW v4-multitemporal (1975-1990-2000-2015). European Commission, Joint Research Centre (JRC), https://data.europa.eu/data/datasets/jrc-ghsl-ghs_smod_pop_globe_r2016a (2016).Doll, C. N. H. CIESIN thematic guide to night-time light remote sensing and its applications. CIESIN http://sedac.ciesin.columbia.edu/binaries/web/sedac/thematic-guides/ciesin_nl_tg.pdf (2008).Meijer, J. R., Huijbregts, M. A. J., Schotten, K. C. G. J. & Schipper, A. M. Global patterns of current and future road infrastructure. Environ. Res. Lett. 13, 64006 (2018).Article 

Google Scholar 
Venter, O. et al. Global terrestrial Human Footprint maps for 1993 and 2009. Sci. data 3, 160067 (2016).PubMed 
PubMed Central 
Article 

Google Scholar 
University of Berkeley. Database of global administrative areas (GADM). University of Berkeley, Museum of Vertebrate Zoology and the International Rice Research Institute, http://www.gadm.org (2012).Kummu, M., Taka, M. & Guillaume, J. H. A. Gridded global datasets for gross domestic product and Human Development Index over 1990–2015. Sci. data 5, 180004 (2018).PubMed 
PubMed Central 
Article 

Google Scholar  More

Study speciesCane toads (Rhinella marina) are large (to  > 1 kg) bufonids (Fig. 1a). Although native to north-eastern South America, these toads have been translocated to many countries worldwide to control insect pests12. Adult cane toads forage at night for insect prey and retreat to moist shelter-sites per day13. Small body size (and thus, high desiccation rate) restricts young toads to the margins of natal ponds14, but adult toads can survive even in highly arid habitats if they have access to water13,15. Cane toads prefer open habitats for foraging12, and thus can thrive in post-fire landscapes16,17. Cane toads in post-fire landscapes tend to have lower parasite burdens, probably because free-living larvae of their lungworm parasites cannot survive either the fire or the more sun-exposed post-fire landscape18.Figure 1taken from study sites between Casino, Grafton, and surrounds, NSW, by S.W. Kaiser.The cane toad Rhinella marina (a), and unburned, (b) and burned (c) habitats in which toads were collected and radio-tracked. Photographs were Full size imageStudy areaEast of the Great Dividing Range, near-coastal Clarence Dry Sclerophyll Forests of north-eastern New South Wales (NSW) are dominated by Spotted gum (Corymbia variegata) and Pink bloodwood (Corymbia intermedia)19. Fires are common, but typically cover relatively small areas before they are extinguished. In the summer of 2019–2020, however, prolonged drought followed by an unusually hot summer resulted in massive fires across this region, burning almost 100,000 km2 of vegetation9. In the current study, the toads we measured and dissected came from several sites within 75 km of the city of Casino (for site locations, see Fig. 2, Table 1, and18). The impacts of fire on faunal abundance and attributes shift with time since fire; for example, the abundance of a particular species may be reduced by fire (due to mortality from flames) but then increase as individuals from surrounding areas migrate to the recently-burned site to exploit new ecological opportunities provided by that landscape8. We chose to study this system 1-year post-fire, to allow time for such longer-term effects to be manifested.Figure 2Sampling sites relative to fire history. Sample sites are burned (red circles), and unburned (green squares). See Table 1 for key to sites. The legend shows the extent of burn a year prior to our study. Map created in QGIS 3.22.3. Fire history available from https://datasets.seed.nsw.gov.au/dataset/fire-extent-and-severity-mapping-fesm CC BY 4.0.Full size imageTable 1 Sampling sites and sample sizes for dissected and radio-tracked cane toads (Rhinella marina) in New South Wales, Australia.Full size tableSurveys of toad abundanceTo quantify toad abundance in burned and unburned sites, one observer (MJG) walked 100-m transects along roads at night (N = 23 and 8 respectively), recording all toads and native frogs (both adult and juvenile). The smaller number of unburned sites reflects the massive spatial scale of the wildfires, which made it difficult to find unburned areas. The transect sites were not the same as those sampled by “toad-busters” (below). We sampled both burned and unburned sites on each night, to de-confound effects of weather conditions with fire treatment. We scored frogs as well as toads to provide an estimate of overall anuran abundance and activity, and so that we could examine toad abundance relative to frog abundance as well as absolute toad numbers.“Toad-buster” sampleBecause of their ecological impact on native fauna, cane toads are culled by community groups as well as by government authorities12,20. We asked “toad-buster” groups to record whether the sites at which they collected toads had been burned during the 2019–2020 fires, or had remained unburned (Table 1). The toads were humanely euthanized (cooled-then-pithed: see21). The euthanasia method is brief (a few hours in the refrigerator, followed by pithing) and thus should not have affected any of the traits that we measured. For all of these toads, we measured body length (snout-urostyle length = SUL) and mass, and determined sex based on external morphology (skin colour and rugosity, nuptial pads: see22). A subset of toads (chosen to provide relatively equal numbers of males and females, and with equal numbers from burned and unburned sites) was dissected to provide data on mass of internal organs (fat bodies, liver, ovaries), reproductive condition (state of ovarian follicle development) and diet (mass and identity of prey items). To select the subsample of toads for dissection, we took relatively equal numbers of male and female toads from each bag of toads that was provided to us by the “toad-busters”. For logistical reasons, we were unable to dissect all of the toads that had been collected. Overall, we obtained data on morphology, diets and other traits from 481 fully dissected and 1443 partially dissected cane toads.Radio-trackingTo explore habitat use and movement patterns, we radio-tracked 57 toads over the course of two fieldtrips (0900–1800 h from 20 Nov 2021 to 6 Dec 2021 and 25 Jan 2022 to 10 Feb 2022). We selected seven sites (4 burned, 3 unburned) within 28 km of Tabbimoble, NSW (see Table 1 for locations and sample sizes of tracked toads). We hand-captured toads found active at night. These were measured, and their sex determined by external morphology (see above) and behaviour (release calls, given only by males: see23). We then fitted the toads with radio-transmitters (PD-2; Holohil Systems, Ontario, Canada; weighing ≤ 3.8 g) on cotton waist-belts, and released them at the site of capture. Tracked toads were 88.2–160.9 mm SUL (mass 70.1–546.3 g); thus, transmitters weighed  20 mm thick) within the quadrat, and estimated exposure of the toad within its refuge (the percentage of the animal’s body exposed to the naked eye). We then selected a compass bearing at random and walked 20 m in that direction where we rescored all of the above habitat attributes, to quantify habitat features in the broader environment (i.e., not just in microhabitats used by toads). We used those “random” sites to quantify overall habitat attributes of burned and unburned sites. Temperature was recorded by directing a temperature gun (Digitech QM7221) on (or otherwise close-to) toads and at a random point on the ground for random replicates. In total, we gathered radio-tracking data on movements and habitat variables from 57 cane toads, each of which was tracked for 5 days. Recaptured toads were euthanized by cooling-then-pithing.Morphological traitsTo obtain an index of body condition of toads, we regressed ln mass against ln SUL, and used the residual scores from that general linear regression as our estimate of body condition. Negative residual scores show an individual that weighs less-than-expected based on its body length. Likewise, we regressed mass of the fat bodies, liver and stomach against body mass to obtain indices of energy stores and stomach-content volumes relative to body mass. We scored male secondary sexual characteristics using the system of Bowcock et al.22. In their system, three sexually dimorphic traits (nuptial pad size, skin roughness and skin colouration) are scored from 0 to 2, and the scores from those three traits are summed to create a final value (on a 6-point scale) for the degree of elaboration of male-specific secondary sexual characteristics. We scored reproductive condition in adult female toads based on whether or not egg masses were visible during dissection, based on dissected toads from both “toad-buster” and telemetry samples.Statistical methodsData were analysed in R version 4.2.025. We used Linear Mixed Models (LMMs), Generalised Linear Mixed Models (GLMMs) and logistic regressions for our analyses. The R packages ‘tidyverse’26, ‘lmerTest’27, and ‘performance’28 were used.Habitat dataWe compared habitat variables between burned and unburned sites, and attributes of toads in burned versus unburned sites, using GLMMs (with negative binomial distribution) for count data (models were checked for overdispersion29) and LMMs on distance data, using ln-transformations where required to achieve normality. LMMs were used on non-normal percentage data, which were ln- and then logit-transformed (using log[(P + e)/(1 − P + e)], where e is the lowest non-zero number, halved)30. We used toad id, site (sampling location) and sampling trip (2019 versus 2020) as random factors.Anuran transect dataCounts of toads in burned versus unburned areas were compared both directly via GLMMs with a negative binomial distribution and relative to the numbers of frogs sighted along the same transects (binding the columns in R as ‘number of toads, number of amphibians – number of toads’ and using a GLMM with a binomial distribution). We used site as a random factor.Telemetry dataFor telemetry data, we analysed response variables via LMMs, and ln-transformed data where relevant to achieve normality.Dissection dataWe used LMMs for SUL, body mass, body condition and organ mass residuals (e.g., fat body mass relative to body mass). For prey item data, we used a poisson distribution with row number as a random factor, as the negative binomial and beta distribution GLMMs were overdispersed (see31). We used LMM for number of prey items and number of prey groups, with site as a random factor. Where models failed to converge, we reduced or removed the error term(s). Analyses were restricted to toads ≥ 70 mm SUL, because animals below this size were difficult to sex. We also performed nominal logistic regression to explore variation in sex ratio and male secondary sexual traits.Reproductive conditionWe used LMM for male secondary sexual characteristic display, using site as a random factor. For ovary presence, we used a binomial GLMM with a logit link, using site as a random factor. We used a LMM of the residual values from ovary mass relative to body mass (ln-transformed), using site as a random factor.Ethics declarationsAll procedures were performed in accordance with the relevant guidelines and regulations approved by Macquarie University Animal Ethics Committee (ARA Number: 2019/040-2) and in accordance with ARRIVE guidelines. More

Thiel, M. et al. The Humboldt Current System of northern and central Chile—Oceanographic processes, ecological interactions and socioeconomic feedback. Oceanogr. Mar. Biol. Annu. Rev. 45, 195–344 (2007).
Google Scholar 
FAO. The State of World Fisheries and Aquaculture 2020. Sustainability in action 2020.Castilla, J. C. & Camus, P. A. The Humboldt-El Niño scenario: Coastal benthic resources and anthropogenic influences, with particular reference to the 1982/83 ENSO. S. Afr. J. Mar. Sci. 12, 703–712. https://doi.org/10.2989/02577619209504735 (1992).Article 

Google Scholar 
Alheit, J. & Niquen, M. Regime shifts in the Humboldt Current ecosystem. Prog. Oceanogr. 60, 201–222. https://doi.org/10.1016/j.pocean.2004.02.006 (2004).Article 

Google Scholar 
González, H. E. et al. Carbon fluxes within the epipelagic zone of the Humboldt Current System off Chile: The significance of euphausiids and diatoms as key functional groups for the biological pump. Prog. Oceanogr. 83, 217–227. https://doi.org/10.1016/j.pocean.2009.07.036 (2009).Article 

Google Scholar 
Quiñones, R. A., Levipan, H. A. & Urrutia, H. Spatial and temporal variability of planktonic archaeal abundance in the Humboldt Current System off Chile. Deep Sea Res. Part II 56, 1073–1082. https://doi.org/10.1016/j.dsr2.2008.09.012 (2009).Article 

Google Scholar 
Antezana, T. Euphausia mucronata: A keystone herbivore and prey of the Humboldt Current System. Deep Sea Res. Part II 57, 652–662. https://doi.org/10.1016/j.dsr2.2009.10.014 (2010).Article 

Google Scholar 
Anguita, C., Gelcich, S., Aldana, M. & Pulgar, J. Exploring the influence of upwelling on the total allowed catch and harvests of a benthic gastropod managed under a territorial user rights for fisheries regime along the Chilean coast. Ocean Coast. Manag. 195, 105256. https://doi.org/10.1016/j.ocecoaman.2020.105256 (2020).Article 

Google Scholar 
González, J. E., Yannicelli, B. & Stotz, W. The interplay of natural variability, productivity and management of the benthic ecosystem in the Humboldt Current System: Twenty years of assessment of Concholepas concholepas fishery under a TURF management system. Ocean Coast. Manag. 208, 105628. https://doi.org/10.1016/j.ocecoaman.2021.105628 (2021).Article 

Google Scholar 
Canales, T. M. et al. Endogenous, climate, and fishing influences on the population dynamics of Small Pelagic Fish in the Southern Humboldt Current Ecosystem. Front. Mar. Sci. https://doi.org/10.3389/fmars.2020.00082 (2020).Article 

Google Scholar 
González, J. E., Ortiz, M. Exploring harvest strategies in a benthic habitat in the Humboldt Current System (Chile): A study case. In Marine Coastal Ecosystems Modelling and Conservation: Latin American Experiences 127–141 (Springer International Publishing, 2021). https://doi.org/10.1007/978-3-030-58211-1_6.Ortiz, M. Pre-image population indices for anchovy and sardine species in the Humboldt Current System off Peru and Chile: Years decaying productivity. Ecol. Ind. 119, 106844. https://doi.org/10.1016/j.ecolind.2020.106844 (2020).Article 

Google Scholar 
Tognelli, M. F., Silva-Garcia, C., Labra, F. A. & Marquet, P. A. Priority areas for the conservation of coastal marine vertebrates in Chile. Biol. Conserv. 126, 420–428. https://doi.org/10.1016/j.biocon.2005.06.021 (2005).Article 

Google Scholar 
Bustamante, C., Vargas-Caro, C. & Bennett, M. B. Not all fish are equal: Functional biodiversity of cartilaginous fishes (Elasmobranchii and Holocephali) in Chile. J. Fish Biol. 85, 1617–1633. https://doi.org/10.1111/jfb.12517 (2014).CAS 
Article 
PubMed 

Google Scholar 
Sarmiento-Devia, R. A., Harrod, C. & Pacheco, A. S. Ecology and Conservation of Sea Turtles in Chile. Chelonian Conserv. Biol. 14, 21–33. https://doi.org/10.2744/ccab-14-01-21-33.1 (2015).Article 

Google Scholar 
Pérez-Álvarez, M. J., Alvarez, E., Aguayo-Lobo, A. & Olavarría, C. Occurrence and distribution of Chilean dolphin (Cephalorhynchus eutropia) in coastal waters of central Chile. N.Z. J. Mar. Freshw. Res. 41, 405–409. https://doi.org/10.1080/00288330709509931 (2007).Article 

Google Scholar 
Pacheco, A. S. et al. Cetacean diversity revealed from whale-watching observations in Northern Peru. Aquat. Mamm. 45, 116–122. https://doi.org/10.1578/AM.45.1.2019.116 (2019).Article 

Google Scholar 
Buchan, S. J., Vásquez, P., Olavarría, C. & Castro, L. R. Prey items of baleen whale species off the coast of Chile from fecal plume analysis. Mar. Mamm. Sci. 37, 1116–1127 (2021).Article 

Google Scholar 
Hucke-Gaete, R. et al. From Chilean Patagonia to Galapagos, Ecuador: Novel insights on blue whale migratory pathways along the Eastern South Pacific. PeerJ 6, e4695. https://doi.org/10.7717/peerj.4695 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Llapapasca, M. A. et al. Modeling the potential habitats of dusky, commons and bottlenose dolphins in the Humboldt Current System off Peru: The influence of non-El Niño vs. El Niño 1997–98 conditions and potential prey availability. Prog. Oceanogr. 168, 169–181. https://doi.org/10.1016/j.pocean.2018.09.003 (2018).Article 

Google Scholar 
Sepúlveda, M. et al. From whaling to whale watching: Identifying fin whale critical foraging habitats off the Chilean coast. Aquat. Conserv. Mar. Freshw. Ecosyst. 28, 821–829. https://doi.org/10.1002/aqc.2899 (2018).Article 

Google Scholar 
Williams, R. et al. Chilean blue whales as a case study to illustrate methods to estimate abundance and evaluate conservation status of rare species. Conserv. Biol. 25, 526–535. https://doi.org/10.1111/j.1523-1739.2011.01656.x (2011).Article 
PubMed 

Google Scholar 
Moore, J. E. & Barlow, J. Bayesian state-space model of fin whale abundance trends from a 1991–2008 time series of line-transect surveys in the California Current. J. Appl. Ecol. 48, 1195–1205. https://doi.org/10.1111/j.1365-2664.2011.02018.x (2011).Article 

Google Scholar 
Campbell, G. S. et al. Inter-annual and seasonal trends in cetacean distribution, density and abundance off southern California. Deep Sea Res. Part II 112, 143–157. https://doi.org/10.1016/j.dsr2.2014.10.008 (2015).Article 

Google Scholar 
Nichol, L. M., Wright, B. M., O’Hara, P. & Ford, J. K. B. Risk of lethal vessel strikes to humpback and fin whales off the west coast of Vancouver Island, Canada. Endanger. Species Res. 32, 373–390. https://doi.org/10.3354/esr00813 (2017).Article 

Google Scholar 
Pennino, M. G. et al. A spatially explicit risk assessment approach: Cetaceans and marine traffic in the Pelagos Sanctuary (Mediterranean Sea). PLoS One 12, e0179686. https://doi.org/10.1371/journal.pone.0179686 (2017).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Van Waerebeek, K. & Reyes, J. C. Catch of small cetaceans at Pucusana Port, central Peru, during 1987. Biol. Conserv. 51, 15–22. https://doi.org/10.1016/0006-3207(90)90028-N (1990).Article 

Google Scholar 
Mangel, J. C. et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Conserv. 143, 136–143. https://doi.org/10.1016/j.biocon.2009.09.017 (2010).Article 

Google Scholar 
Campbell, E., Pasara-Polack, A., Mangel, J. C. & Alfaro-Shigueto, J. Use of small cetaceans as bait in small-scale fisheries in Peru. Front. Mar. Sci. https://doi.org/10.3389/fmars.2020.534507 (2020).Article 

Google Scholar 
Reyes, J. C. & Oporto, J. A. Gillnet fisheries and cetaceans in the southeast Pacific. Report of the International Whaling Commission 467–474 (1994).Aguayo-Lobo, A. Los cetáceos y sus perspectivas de conservación. Estudios Oceanológicos 18, 35–43 (1999).
Google Scholar 
Félix, F., Muñoz, M., Falconí, J., Botero, N., Haase, B., et al. Entanglement of humpback whales in artisanal fishing gear in Ecuador. J. Cetacean. Res. Manag. 283–290 (2020).Félix, F. et al. Challenges and opportunities for the conservation of marine mammals in the Southeast Pacific with the entry into force of the U.S. Marine Mammal Protection Act. Reg. Stud. Mar. Sci. 48, 102036. https://doi.org/10.1016/j.rsma.2021.102036 (2021).Article 

Google Scholar 
García-Cegarra, A. M. & Pacheco, A. S. Collision risk areas between fin and humpback whales with large cargo vessels in Mejillones Bay (23°S), northern Chile. Mar. Policy 103, 182–186. https://doi.org/10.1016/j.marpol.2018.12.022 (2019).Article 

Google Scholar 
Santos-Carvallo, M. et al. Impacts of whale-watching on the short-term behavior of Fin Whales (Balaenoptera physalus) in a marine protected area in the southeastern pacific. Front. Mar. Sci. https://doi.org/10.3389/fmars.2021.623954 (2021).Article 

Google Scholar 
Villagra, D., García-Cegarra, A., Gallardo, D. I. & Pacheco, A. S. Energetic effects of whale-watching boats on humpback whales on a breeding ground. Front. Mar. Sci. https://doi.org/10.3389/fmars.2020.600508 (2021).Article 

Google Scholar 
Buckland, S., Anderson, D., Burnham, K., Laake, J., Borchers, D., Thomas, L. Introduction to Distance Sampling Estimating Abundance of Biological Populations. (Oxford University Press, 2001).Hedley, S. L. & Buckland, S. T. Spatial models for line transect sampling. JABES 9, 181–199. https://doi.org/10.1198/1085711043578 (2004).Article 

Google Scholar 
Williams, R., Hedley, S. L., Hammond, P. S. Modeling distribution and abundance of Antarctic baleen whales using ships of opportunity (2006).DoniolValcroze, T., Berteaux, D., Larouche, P. & Sears, R. Influence of thermal fronts on habitat selection by four rorqual whale species in the Gulf of St. Lawrence. Mar. Ecol. Prog. Ser. 335, 207–216. https://doi.org/10.3354/meps335207 (2007).Article 

Google Scholar 
Scales, K. L. et al. Should I stay or should I go? Modelling year-round habitat suitability and drivers of residency for fin whales in the California Current. Divers. Distrib. 23, 1204–1215. https://doi.org/10.1111/ddi.12611 (2017).Article 

Google Scholar 
Bedriñana-Romano, L. et al. Integrating multiple data sources for assessing blue whale abundance and distribution in Chilean Northern Patagonia. Divers. Distrib. https://doi.org/10.1111/ddi.12739 (2018).Article 

Google Scholar 
Bedriñana-Romano, L. et al. Defining priority areas for blue whale conservation and investigating overlap with vessel traffic in Chilean Patagonia, using a fast-fitting movement model. Sci. Rep. 11, 2709. https://doi.org/10.1038/s41598-021-82220-5 (2021).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pirotta, E., Matthiopoulos, J., MacKenzie, M., Scott-Hayward, L. & Rendell, L. Modelling sperm whale habitat preference: A novel approach combining transect and follow data. Mar. Ecol. Prog. Ser. 436, 257–272. https://doi.org/10.3354/meps09236 (2011).Article 

Google Scholar 
Mendelssohn, R. rerddapXtracto: Extracts Environmental Data from “ERDDAP” Web Services. (2020).Lau-Medrano, W. grec: Gradient-Based Recognition of Spatial Patterns in Environmental Data. (2020).Belkin, I. M. & O’Reilly, J. E. An algorithm for oceanic front detection in chlorophyll and SST satellite imagery. J. Mar. Syst. 78, 319–326. https://doi.org/10.1016/j.jmarsys.2008.11.018 (2009).Article 

Google Scholar 
Hijmans, R. J., van Etten, J., Cheng, J., Sumner, M., Mattiuzzi, M., Greenberg, J. A., et al. raster: Geographic Data Analysis and Modeling. (2018).Royle, J. A. N-mixture models for estimating population size from spatially replicated counts. Biometrics 60, 108–115. https://doi.org/10.1111/j.0006-341X.2004.00142.x (2004).MathSciNet 
Article 
PubMed 
MATH 

Google Scholar 
Chelgren, N. D., Samora, B., Adams, M. J. & McCreary, B. Using spatiotemporal models and distance sampling to map the space use and abundance of newly metamorphosed Western Toads (Anaxyrus boreas). Herpetol. Conserv. Biol. 6, 16 (2011).
Google Scholar 
Hartig, F., Lohse, L. DHARMa: Residual Diagnostics for Hierarchical (Multi-Level/Mixed) Regression Models. (2022).Gelman, A., Meng, X.-L. & Stern, H. Posterior predictive assessment of model fitness via realized discrepancies. Stat. Sin. 6, 733–760. https://doi.org/10.2307/24306036 (1996).MathSciNet 
Article 
MATH 

Google Scholar 
Kery, M. & Royle, J. A. Applied Hierarchical Modeling in Ecology: Analysis of Distribution, Abundance and Species Richness in R and BUGS: Volume 1: Prelude and Static Models. (Academic Press, 2015).R DCT. R: A Language and Environment for Statistical Computing. (R Foundation for Statistical Computing, 2015).Plummer, M. JAGS: A program for analysis of Bayesian graphical models using Gibbs sampling. (2003).Fonnesbeck, C. J., Garrison, L. P., Ward-Geiger, L. I. & Baumstark, R. D. Bayesian hierarchichal model for evaluating the risk of vessel strikes on North Atlantic right whales in the SE United States. Endanger. Species Res. 6, 87–94. https://doi.org/10.3354/esr00134 (2008).Article 

Google Scholar 
Vanderlaan, A. S. M., Taggart, C. T., Serdynska, A. R., Kenney, R. D. & Brown, M. W. Reducing the risk of lethal encounters: Vessels and right whales in the Bay of Fundy and on the Scotian Shelf. Endanger. Species Res. 4, 283–297. https://doi.org/10.3354/esr00083 (2008).Article 

Google Scholar 
Warren, D. L., Glor, R. E. & Turelli, M. Environmental niche equivalency versus conservatism: Quantitative approaches to niche evolution. Evolution 62, 2868–2883. https://doi.org/10.1111/j.1558-5646.2008.00482.x (2008).Article 
PubMed 

Google Scholar 
Hijmans, R. J., Phillips, S., Leathwick, J., Elith, J. & Hijmans, M. R. J. Package ‘dismo’. Circles 9, 1–68 (2017).
Google Scholar 
Daneri, G. et al. Primary production and community respiration in the Humboldt Current System off Chile and associated oceanic areas. Mar. Ecol. Prog. Ser. 197, 41–49. https://doi.org/10.3354/meps197041 (2000).Article 

Google Scholar 
Montecino, V. & Lange, C. B. The Humboldt Current System: Ecosystem components and processes, fisheries, and sediment studies. Prog. Oceanogr. 83, 65–79. https://doi.org/10.1016/j.pocean.2009.07.041 (2009).Article 

Google Scholar 
Escribano, R., Hidalgo, P. & Krautz, C. Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000. Deep Sea Res. Part II 56, 1083–1094. https://doi.org/10.1016/j.dsr2.2008.09.009 (2009).Article 

Google Scholar 
Perez-Alvarez, M. et al. Fin whales (Balaenoptera physalus) feeding on Euphausia mucronata in nearshore waters off North-Central Chile. Aquat. Mamm. 32, 109–113. https://doi.org/10.1578/AM.32.1.2006.109 (2006).Article 

Google Scholar 
Riquelme-Bugueño, R. et al. Fatty acid composition in the endemic Humboldt Current krill, Euphausia mucronata (Crustacea, Euphausiacea) in relation to the phytoplankton community and oceanographic variability off Dichato coast in central Chile. Prog. Oceanogr. 188, 102425. https://doi.org/10.1016/j.pocean.2020.102425 (2020).Article 

Google Scholar 
Escribano, R., Marin, V. & Irribarren, C. Distribution of Euphausia mucronata at the upwelling area of Peninsula Mejillones, northern Chile: The influence of the oxygen minimum layer. Sci. Mar. 64, 69–77. https://doi.org/10.3989/scimar.2000.64n169 (2000).Article 

Google Scholar 
Riquelme-Bugueno, R., Escribano, R. & Gomez-Gutierrez, J. Somatic and molt production in Euphausia mucronata off central-southern Chile: The influence of coastal upwelling variability. Mar. Ecol. Prog. Ser. 476, 39–57 (2013).Article 

Google Scholar 
Savoca, M. S. et al. Baleen whale prey consumption based on high-resolution foraging measurements. Nature 599, 85–90. https://doi.org/10.1038/s41586-021-03991-5 (2021).CAS 
Article 
PubMed 

Google Scholar 
Roman, J. & McCarthy, J. J. The whale pump: Marine mammals enhance primary productivity in a coastal basin. PLoS One 5, e13255. https://doi.org/10.1371/journal.pone.0013255 (2010).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Hucke-Gaete, R. Whales might also be an important component in patagonian fjord ecosystems: Comment to Iriarte et al. Ambio 40, 104–105. https://doi.org/10.1007/s13280-010-0110-8 (2011).Article 
PubMed 
PubMed Central 

Google Scholar 
Lavery, T. J. et al. Whales sustain fisheries: Blue whales stimulate primary production in the Southern Ocean. Mar. Mamm. Sci. https://doi.org/10.1111/mms.12108 (2014).Article 

Google Scholar 
Roman, J. et al. Whales as marine ecosystem engineers. Front. Ecol. Environ. 12, 377–385. https://doi.org/10.1890/130220 (2014).Article 

Google Scholar 
Hucke-Gaete, R., Osman, L. P., Moreno, C. A., Findlay, K. P. & Ljungblad, D. K. Discovery of a blue whale feeding and nursing ground in southern Chile. Proc. R. Soc. Lond. B 271, S170–S173. https://doi.org/10.1098/rsbl.2003.0132 (2004).Article 

Google Scholar 
Buchan, S. J. & Quiones, R. A. First insights into the oceanographic characteristics of a blue whale feeding ground in northern Patagonia, Chile. Mar. Ecol. Prog. Ser. 554, 183–199. https://doi.org/10.3354/meps11762 (2016).CAS 
Article 

Google Scholar 
Findlay, K., Pitman, R., Tsurui, T., Sakai, K., Ensor, P., Iwakami, H., et al. IWC-southern whale and ecosystem research (IWC/SOWER) blue whale Cruise, Chile. Documento Técnico, IWC 1998 (1998).Branch, T. A. et al. Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean. Mamm. Rev. 37, 116–175. https://doi.org/10.1111/j.1365-2907.2007.00106.x (2007).Article 

Google Scholar 
Barlow, D. R., Klinck, H., Ponirakis, D., Garvey, C. & Torres, L. G. Temporal and spatial lags between wind, coastal upwelling, and blue whale occurrence. Sci. Rep. 11, 6915. https://doi.org/10.1038/s41598-021-86403-y (2021).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Galletti-Vernazzani, B., Jackson, J. A., Cabrera, E., Carlson, C. A. Jr. & RLB.,. Estimates of abundance and trend of chilean blue whales off Isla de Chiloé, Chile. PLoS One 12, e0168646. https://doi.org/10.1371/journal.pone.0168646 (2017).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Friedlaender, A. S., Goldbogen, J. A., Hazen, E. L., Calambokidis, J. & Southall, B. L. Feeding performance by sympatric blue and fin whales exploiting a common prey resource. Mar. Mamm. Sci. 31, 345–354. https://doi.org/10.1111/mms.12134 (2015).Article 

Google Scholar 
Abrahms, B. et al. Memory and resource tracking drive blue whale migrations. PNAS 116, 5582–5587 (2019).CAS 
Article 

Google Scholar 
Clarke, R., Aguayo, A. & Basulto, S. Whale observation and whale marking off the coast of Chile in 1964. Sci. Rep. Whales Res. Inst. Tokyo 30, 117–178 (1978).
Google Scholar 
Allison, C. IWC individual and summary catch databases Version 5.5 (12 February 2013). Available from the International Whaling Commission 135 (2013).Pastene, L. A., Acevedo, J. & Branch, T. A. Morphometric analysis of Chilean blue whales and implications for their taxonomy. Mar. Mamm. Sci. 36, 116–135. https://doi.org/10.1111/mms.12625 (2020).Article 

Google Scholar 
Rendell, L., Whitehead, H. & Escribano, R. Sperm whale habitat use and foraging success off northern Chile: Evidence of ecological links between coastal and pelagic systems. Mar. Ecol. Prog. Ser. 275, 289–295. https://doi.org/10.3354/meps275289 (2004).Article 

Google Scholar 
Jaquet, N. & Whitehead, H. Scale-dependent correlation of sperm whale distribution with environmental features and productivity in the South Pacific. Mar. Ecol. Prog. Ser. 135, 1–9. https://doi.org/10.3354/meps135001 (1996).Article 

Google Scholar 
O’Hern, J. E., Biggs, D. C. Sperm whale (Physeter macrocephalus) habitat in the Gulf of Mexico: Satellite observed ocean color and altimetry applied to small-scale variability in distribution. Aquat. Mamm. 35 (2009).Koen Alonso, M., Crespo, E. A., García, N. A., Pedraza, S. N. & Coscarella, M. A. Diet of dusky dolphins, Lagenorhynchus obscurus, in waters off Patagonia, Argentina. Fish. Bull. 96, 366–374 (1998).
Google Scholar 
García-Godos, I., Waerebeek, K. V., Reyes, J. C., Alfaro-Shigueto, J. & Arias-Schreiber, M. Prey occurrence in the stomach contents of four small cetacean species in Peru. Latin Am. J. Aquat. Mamm. 6, 171–183. https://doi.org/10.5597/lajam00122 (2007).Article 

Google Scholar 
Dans, S. L., Crespo, E. A., Koen-Alonso, M., Markowitz, T. M., Berón Vera, B., Dahood, A. D. Chapter 3—Dusky dolphin trophic ecology: Their role in the food web. In The Dusky Dolphin (eds. Würsig, B., Würsig, M.) 49–74 (Academic Press, 2010). https://doi.org/10.1016/B978-0-12-373723-6.00003-5.Romero, M. A. et al. Feeding habits of two sympatric dolphin species off North Patagonia, Argentina. Mar. Mamm. Sci. 28, 364–377 (2012).Article 

Google Scholar 
Loizaga de Castro, R. et al. Feeding ecology of dusky dolphins Lagenorhynchus obscurus: Evidence from stable isotopes. J. Mammal. 97, 310–320. https://doi.org/10.1093/jmammal/gyv180 (2016).Article 

Google Scholar 
Cipriano, F. W. Behavior and occurrence patterns, feeding ecology, and life history of dusky dolphins (Lagenorhynchus obscurus) off Kaikoura, New Zealand. (1992).Benoit-Bird, K. J., Würsig, B. & Mfadden, C. J. Dusky dolphin (lagenorhynchus obscurus) foraging in two different habitats: Active acoustic detection of dolphins and their prey. Mar. Mamm. Sci. 20, 215–231. https://doi.org/10.1111/j.1748-7692.2004.tb01152.x (2004).Article 

Google Scholar 
Van Waerebeek, K. Records of dusky dolphins Lagenorhynchus obscurus (Gray, 1828) in the eastern South Pacific. Beaufortia (1992).Selzer, L. A. & Payne, P. M. The distribution of white-sided (Lagenorhynchus acutus) and common dolphins (Delphinus delphis) vs. Environmental features of the continental shelf of the Northeastern United States. Mar. Mamm. Sci. 4, 141–153. https://doi.org/10.1111/j.1748-7692.1988.tb00194.x (1988).Article 

Google Scholar 
Neumann, D. R. Seasonal movements of short-beaked common dolphins (Delphinus delphis) in the north-western Bay of Plenty, New Zealand: Influence of sea surface temperature and El Niño/La Niña. N.Z. J. Mar. Freshw. Res. 35, 371–374. https://doi.org/10.1080/00288330.2001.9517007 (2001).Article 

Google Scholar 
Peters, K. J. et al. Foraging ecology of the common dolphin Delphinus delphis revealed by stable isotope analysis. Mar. Ecol. Prog. Ser. 652, 173–186. https://doi.org/10.3354/meps13482 (2020).CAS 
Article 

Google Scholar 
Brand, D. et al. Common dolphins, common in neritic waters off southern Israel, demonstrate uncommon dietary habits. Aquat. Conserv. Mar. Freshw. Ecosyst. 31, 15–21. https://doi.org/10.1002/aqc.3165 (2021).Article 

Google Scholar 
Barlow, J. & Taylor, B. L. Estimates of sperm whale abundance in the Northeastern temperate pacific from a combined acoustic and visual survey. Mar. Mamm. Sci. 21, 429–445. https://doi.org/10.1111/j.1748-7692.2005.tb01242.x (2005).Article 

Google Scholar 
Cañadas, A., Desportes, G. & Borchers, D. Estimation of g (0) and abundance of common dolphins (Delphinus delphis) from the NASS-95 Faroese survey. J. Cetac. Res. Manag. 6, 191–198 (2004).
Google Scholar 
Miller, D. L., Burt, M. L., Rexstad, E. A. & Thomas, L. Spatial models for distance sampling data: Recent developments and future directions. Methods Ecol. Evol. 4, 1001–1010. https://doi.org/10.1111/2041-210X.12105 (2013).Article 

Google Scholar 
Sigourney, D. B. et al. Developing and assessing a density surface model in a Bayesian hierarchical framework with a focus on uncertainty: Insights from simulations and an application to fin whales (Balaenoptera physalus). PeerJ 8, e8226. https://doi.org/10.7717/peerj.8226 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Panigada, S. et al. Mediterranean fin whales at risk from fatal ship strikes. Mar. Pollut. Bull. 52, 1287–1298. https://doi.org/10.1016/j.marpolbul.2006.03.014 (2006).CAS 
Article 
PubMed 

Google Scholar 
Ribeiro, S., Viddi, F. A. & Freitas, T. R. Behavioural responses of Chilean dolphins (Cephalorhynchus eutropia) to boats in Yaldad Bay, southern Chile. Aquat. Mamm. 31, 234 (2005).Article 

Google Scholar 
Bearzi, G. et al. Overfishing and the disappearance of short-beaked common dolphins from western Greece. Endanger. Species Res. 5, 1–12. https://doi.org/10.3354/esr00103 (2008).Article 

Google Scholar 
Reeves, R. R., McClellan, K. & Werner, T. B. Marine mammal bycatch in gillnet and other entangling net fisheries, 1990 to 2011. Endanger. Species Res. 20, 71–97. https://doi.org/10.3354/esr00481 (2013).Article 

Google Scholar 
van der Hoop, J. M. et al. Vessel strikes to large whales before and after the 2008 Ship Strike Rule. Conserv. Lett. 8, 24–32. https://doi.org/10.1111/conl.12105 (2015).Article 

Google Scholar 
Erbe, C., Reichmuth, C., Cunningham, K., Lucke, K. & Dooling, R. Communication masking in marine mammals: A review and research strategy. Mar. Pollut. Bull. 103, 15–38. https://doi.org/10.1016/j.marpolbul.2015.12.007 (2016).CAS 
Article 
PubMed 

Google Scholar 
González-But, J. C. & Sepúlveda, M. Captura incidental del delfín común (Delphinus delphis) en la pesquería industrial de cerco, norte de Chile. Rev. Biol. Mar. Oceanogr. 51, 429–433. https://doi.org/10.4067/S0718-19572016000200019 (2016).Article 

Google Scholar 
Alvarado-Rybak, M. et al. Pathological findings in cetaceans sporadically stranded along the Chilean Coast. Front. Mar. Sci. 7, 684. https://doi.org/10.3389/fmars.2020.00684 (2020).Article 

Google Scholar 
Dans, S. L., Koen, A. M., Pedraza, S. & Crespo, E. A. Incidental catch of dolphins in trawling fisheries off Patagonia, Argentina: Can populations persist?. Ecol. Appl. 13, 754–762. https://doi.org/10.1890/1051-0761(2003)013[0754:ICODIT]2.0.CO;2 (2003).Article 

Google Scholar 
Childerhouse S, Baxter A. Human interactions with dusky dolphins: A management perspective, Chapter 12. In The Dusky Dolphin (eds. Würsig, B. & Würsig, M.) 245–275 (Academic Press, 2010). https://doi.org/10.1016/B978-0-12-373723-6.00012-6.Mannocci, L. et al. Assessing the impact of bycatch on dolphin populations: The case of the common dolphin in the Eastern North Atlantic. PLoS One 7, e32615. https://doi.org/10.1371/journal.pone.0032615 (2012).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Thompson, F. N., Abraham, E. R. & Berkenbusch, K. Common dolphin (Delphinus delphis) Bycatch in New Zealand commercial trawl fisheries. PLoS One 8, e64438. https://doi.org/10.1371/journal.pone.0064438 (2013).Article 
PubMed 
PubMed Central 

Google Scholar  More

Proteobacteria are major contributors to marine microbial phosphonate cyclingDatabases for all putative sequences of genes for phosphonate production (pepM, aepY, phpC, mpnS, hepD), substrate-specific catabolism (phnAWXYZ, palA), and broad-specificity catabolism (phnIJM) were created using available public genomes from JGI IMG/MER and GORG-Tropics. Gene identity was verified by the presence of catalytically essential residues (Supplementary Table S2). Phosphonate genes were identified in 10,337 genomes of bacteria and archaea spanning over 100 unique classes, suggesting a wide variety of microorganisms mediate phosphonate production and catabolism (Fig. 2, Supplementary Dataset S1). A high proportion of all collected sequences affiliated with Proteobacteria (Gamma, Alpha, and Beta classes), averaging 52% of the production genes, 78% of substrate-specific catabolism genes, and 88% of broad-specificity catabolism genes before dereplication (Fig. 2).Fig. 2: Phosphonate gene and genome count with taxonomic distribution.Number of sequences and genomes collected for study (A, D, G) with distribution of class-level taxa for all redundant sequences (B, E, H) and marine redundant (C, F, I) sequences. Results are shown for selected genes representing phosphonate (A–C) production, (D–F) substrate-specific catabolism, and (G–I) broad-specificity catabolism. The taxa shown are the 15 classes with the highest representation across all databases.Full size imageOf the 10,337 genomes, 1556 (15%) were confirmed to be marine organisms from 35 different classes (Fig. 2, Supplementary Dataset S1). Proteobacteria had even greater representation in the subset of marine genomes, averaging 65% of marine production genes, 88% of marine substrate-specific catabolism genes, and 96% of marine broad-specificity catabolism genes from the redundant databases (Fig. 2). The dominance of Alphaproteobacteria in the marine subset may be attributed to the wide variety of Pelagibacterales bacterium captured in the database, making up 426 (27%) of the 1556 genomes involved in all three categories of phosphonate cycling. Rhodobacterales (Ascidiaceihabitans sp., Roseovarius sp., Sulfitobacter sp., Labrenzia sp., and Phaeobacter sp.) alongside Rhodospirillales (Thalassobaculum sp., Thalassospira sp., Roseospira sp., Varunaivibrio sp., and Oceanibaculum sp.) were also highly represented among the marine subset with 214 (14%) and 251 (16%) genomes, respectively (Supplemental Dataset S1), though these taxa primarily show potential for phosphonate catabolism rather than production. Vibrionales were well represented in the JGI IMG/MER marine genome subset with 107 (7%) genomes spanning 59 different species including Vibrio lentus, Vibrio breoganii, and Vibrio splendidus.Diverse taxa encode the capacity to produce phosphonate derivativesPhosphonate production is widespread and distributed throughout many different bacteria and archaea. Genes responsible for the first two steps in phosphonate production, pepM and aepY, had the broadest taxonomic distribution within the redundant databases (Shannon indices of 2.66 and 2.76) for all genes in this study, distributed with 0.59 and 0.61 evenness from 70 and 72 unique, verified classes, respectively. Their broad distribution further highlights the ubiquity and necessity of phosphonate compounds to microbial life and function across all environments. Within the marine setting, both pepM and aepY have reference sequences from 22 unique, verified classes which is the second highest class representation in the marine genome subset (Fig. 2). The marine subset of pepM and aepY also have the highest Shannon indices (1.76 and 1.92) distributed with 0.53 and 0.58 evenness, respectively. A majority (87%) of the Alphaproteobacteria phosphonate producers are Pelagibacterales bacterium with other notable taxa including Bacteria: Candidatus Actinomarinaceae, Prochlorococcus sp., Synechococcus sp., Nitrosococcus sp., and MG-I Archaea: Candidatus Nitrosomarinus catalina, Nitrosopumilus maritimus, alongside other unidentified Crenarchaeota and Thaumarchaeota genomes.The gene phpC was found in less than half the number of genomes than pepM and aepY, and encoded by fewer classes in both the general database (47) and marine subset (10). In the full databases, the distribution of retrieved phpC sequences are similar to pepM and aepY with respect to taxonomic ranking, Shannon index (2.49), and evenness (0.60) (Fig. 2A–C, Supplementary Table 5). Within the marine subset, phpC has less Shannon index (1.61) but greater evenness (0.61) than the marine subset of pepM and aepY. All three upstream phosphonate production genes (pepM, aepY, phpC) are found together within Pelagibacterales bacterium, Prochlorococcus sp., Thaumarchaeota, and Crenarchaoeta alongside other taxa such as Oceanospirillales sp., Arenimonas donghaensis, Desulfuromusa kysingii, and Cellulosilyticum lentocellum.We further investigated the relationship between pepM, aepY, and phpC by examining co-occurrence in genomes and synteny with the general, redundant databases. The first two steps in phosphonate biosynthesis are intimately linked (Fig. 3). Out of all genomes with pepM, 86% have aepY, and out of all genomes with aepY, 90% have pepM. By contrast, phpC is not as closely tied to pepM and phosphonate production. We found phpC in just over 20% of genomes with the capability of phosphonate production (Fig. 3), implying that a majority of bacterial and archaeal phosphonate production stops at the production of phosphonoacetaldehyde or 2-AEP (Fig. 1A). Furthermore, half of the phpC genes were not associated with phosphonate production, given 53% of genomes with phpC did not have pepM and 54% did not have aepY (Fig. 3). In these instances, microbes may use phpC within a 2-AEP substrate-specific catabolism operon (Fig. 3) that allows phosphonate compounds to be synthesized by transforming 2-AEP with phnW and phpC into 2-HEP (Figs. 1A and 3). By repurposing 2-AEP, individuals can still create the specific compound needed while bypassing the energetically unfavourable first step of phosphonate production.Fig. 3: Co-occurrence of phosphonate cycling genes within the same genome and examples of genetic organization of phosphonate cycling genes.The heatmap displays co-occurrence of phosphonate cycling genes. Each column represents the subset of all genomes which contain the source gene and the heatmap value represents the fraction of the source genomes which also contain the co-occurring gene. Heatmap values are not symmetrical due to differing number of genomes represented in each column, database size listed above each column. Examples for phosphonate cycling genomic neighbourhoods were chosen to maximize diversity in synteny with examples from both Bacteria and Archaea where applicable. Several phosphonate-specific ABC transport system clusters are labelled as follows: phnC = phosphonate transport system ATPase; phnD = phosphonate transport system substrate-binding; phnE = phosphonate transport system permease; phnS = 2-AEP transport system substrate-binding; phnT = 2-AEP transport system ATP-binding; phnV = 2-AEP transport system permease; palC = transport system permease; palD = transport system ATP-binding; palE = transport system permease. Genes are colour coded by: red = lyase; orange = transcriptional regulator; yellow = hydrolase; green = transferase; light blue = oxidoreductase; dark blue = transaminase; purple = kinase; pink = isomerase; brown = transport; white = synthase; black = uncharacterized protein; grey = unknown.Full size imageA narrow but diverse selection of taxa encoded MpnS, the marker gene for Mpn production and a key determinant in marine methane production. We observed distinct clades of this enzyme in autotrophic archaea and heterotrophic bacteria (Fig. 2B, C). Within the marine ecosystems, Pelagibacterales, Rhodospirillales, Rickettsiales, Oceanospirillales, Flavobacteriales, and Synechococcales are bacterial candidates for MPn production alongside Thaumarchaeota and Crenarchaeota archaeon (Fig. 2B, C). While six of the bacterial genomes with MpnS also encoded genes for phosphonate catabolism, none of the archaeal MPn producers showed capacity for catabolism (Supplementary Dataset S1). The genomic neighbourhoods for general phosphonate production (pepM, aepY, phpC) and MPn production (mpnS) in both bacteria and archaea include genes such as glycosyltransferase, lipopolysaccharide choline phosphotransferase, choline kinase, adenylyltransferase, and arylsulfatase A (Fig. 3) suggesting the potential for synthesis of (methyl)phosphonate esters [93]. This is consistent with previous analysis [29] of the Nitrosopumilus maritimus SCM1 MPn production genomic neighbourhood and biophysical evidence that MPn producing archaea synthesize an exopolysaccharide modified with MPn similar to 2-AEP modified polymers.Contrary to the diversity of the other phosphonate production databases, the hepD database has low Shannon index (0.62) and evenness (0.45) with 79% of sequences mapping to Actinomycetia including Streptomycetales and Corynebacteriales (Fig. 2B, C). The marine subset has lower Shannon index (0.28) and evenness (0.41) where all sequences derive from Pelagibacterales except one from Prochlorococcus sp. The genomic neighbourhood of HMP production may contain genes for cell surface modification such as acetyltransferase, peptidoglycan biosynthesis, and adenylylsulfate transferase, suggesting that some organisms may use HMP as a conjugate for membrane-associated or exported macromolecules similar to theories on MPn utilization. Other examples of hepD synteny contain more specific genes such as the HMP dehydrogenase or other enzymes for downstream modification (Fig. 3).Marine proteobacteria encode genes for substrate-specific and broad-specificity phosphonate catabolismGenes for marine substrate-specific phosphonate catabolism were widespread among Proteobacterial classes, and to a lesser extent amongst other classes including Bacilli, Planctomycetes, and Synechococcus (Fig. 2E,F). Marine substrate-specific catabolism has lower average Shannon index (1.00) and evenness (0.43) than the three general production genes (pepM, aepY, phpC). The most widespread of these genes was phnW, likely due to its pivotal role in 2-AEP transformations as a precursor reaction to phnAY or phnX (Fig. 1, Supplementary Table 5). Marine hydrolases for 2-AEP catabolism, phnA, phnX, and phnZ, have similar Shannon indices (mean: 1.11 ± 0.05) and evenness (mean: 0.41 ± 0.03) (Fig. 2E, F, Supplementary Table 5).While not exclusive, sequenced references demonstrate a strong taxonomic partition between Proteobacterial classes for 2-AEP catabolism pathways phnAWY and phnWX. Over 74% of marine genomes with phnAWY are Alphaproteobacteria, in particular Rhodobacterales species such as Roseovarius nubinhibens, Marivita geojedonensis, and Pelagicola litoralis. On the contrary, ~80% of marine genomes with phnWX are Gammaproteobacteria, specifically of Vibrionales, Oceanospirillales, and Alteromonadales including a wide range of species from Vibrio, Photobacterium, Marinobacterium, Halomonas, and Pseudoalteromonas.Taxonomic distribution for marine phnZ was 72% Alphaproteobacteria with Pelagibacterales making up 45% of marine phnZ sequences. Note that phnZ has the most (17) reference sequences from marine Cyanobacteriia, specifically Prochlorococcus sp., than any other phosphonate catabolizing gene. Lack of marine sequence representatives for catabolism of phosphonopyruvate by palA suggests that either the substrate is uncommon, therefore the function unnecessary, or marine microbes have other methods of catabolizing phosphonopyruvate, perhaps by the C-P lyase. Overall taxonomic distribution of phosphonate substrate-specific catabolism, specifically targeting 2-AEP, suggests said function is essential to many marine heterotrophs within Alphaproteobacteria and Gammaproteobacteria. However 2-AEP catabolism appears to be less universally important than phosphonate production to marine microbial life since the required genes are found in a less diverse selection of taxa.Genetic organization for substrate-specific catabolism genes, particularly those targeting 2-AEP, varied widely in line with the numerous options for 2-AEP catabolism (Fig. 3). Though some bacteria specialize in a single 2-AEP degradation pathway such as only containing phnWAY, others contained multiple hydrolases for 2-AEP catabolism with some incorporating phpC into a 2-AEP specific catabolism operon (Fig. 3). When a genome has two hydrolases for phosphonate catabolism, often phnZ was paired with either phnA or phnX. Co-occurrence between phnZ and either phnA or phnX ranged between 30-50%, whereas co-occurrence between phnA and phnX was between 6-12% (Fig. 3). This discrepancy in co-occurrence may be due to the metabolic similarity between phnA and phnX, where having both may be redundant. Both of these enzymes rely on phnW for 2-AEP catabolism and produce carbon metabolites, whereas phnZ does not need phnW and produces the amino acid glycine (Fig. 1B).C-P lyase genes representing substrate non-specific catabolism were overwhelmingly attributed to Alphaproteobacteria which consisted over 75% of all collected marine sequences for phnIJM (Fig. 2H, I). A wide variety of Rhodobacterales, spanning 55 different genus are the most numerous representatives, followed by Pelagibacterales and Rhodospirillales. The genes in all three databases have very high genome co-occurrence, 89–99%, as expected given all three operate within the same enzyme complex (Fig. 3). Gene co-occurrence, Shannon index, and evenness is lower for phnM than the other two C-P lyase components, phnI and phnJ, likely due to instances of organisms containing two copies of phnM where one copy lies outside the C-P lyase operon [94]. C-P lyase gene databases have lower Shannon index (mean 0.81 ± 0.11) than phosphonate production and 2-AEP substrate-specific catabolism genes (phnAWXZ) (Fig. 3D, G), suggesting broad-specificity phosphonate catabolism by the C-P lyase is a narrowly distributed function (Supplementary Table 5). Organization of C-P lyase operons held the most consistency between example genomes, likely due to the high number of genes simultaneously utilized for lyase construction. These operons encoded a consecutive string of lyase subunits, including a generic phosphonate transporter (phnCDE) and GntR transcriptional regulator (Fig. 3). C-P lyase genes had low genomic co-occurrence with all other phosphonate cycling genes with notable co-occurrence between phnW at 26%, phnZ at 21%, and phnX and 18% (Fig. 3). The low rate of co-occurrence may be due to redundancy in function for P harvesting between the C-P lyase and substrate-specific catabolism. In some cases there are instances of a substrate-specific hydrolase gene located within the C-P lyase operon (Fig. 3).Phosphonate biosynthesis genes are globally prevalent in oceans and increase in mesopelagic watersFollowing curation of phn-gene databases, we analysed 121 metagenomes and 91 metatranscriptomes from the publicly available TARA Oceans expedition (spanning samples from the Atlantic, Indian, Pacific, and Southern Ocean and Red Sea) to investigate the global potential for marine phosphonate cycling. Measuring the proportion of the community capable of performing specific tasks through metagenomics indicates the long-term selective pressures that shape P-cycling and microbial communities.Potential for phosphonate production (pepM) was globally ubiquitous across all depths, with 14–17% of the community encoding in the surface waters and deep chlorophyll maximum (DCM), increasing to 45% in mesopalagic waters (Fig. 4A, Supplementary Tables S6 and S7), highlighting the importance of phosphonate compounds to marine microbial communities. Relative abundance of phpC was 64–76% that of pepM and aepY across all depths (Fig. 4A). We observed significant increase in relative abundance between the surface and mesopelagic for pepM (ANOVA: F = 1262, p  More

Foote, M. The evolution of morphological diversity. Annu. Rev. Ecol. Syst. 28, 129–152 (1997).Article 

Google Scholar 
Félix, M. A. Phenotypic evolution with and beyond genome evolution. Curr. Top. Dev. Biol. 119, 291–347 (2016).PubMed 
Article 
CAS 

Google Scholar 
Carroll, S. B. Evo-devo and an expanding evolutionary synthesis: A genetic theory of morphological evolution. Cell 134, 25–36 (2008).CAS 
PubMed 
Article 

Google Scholar 
Harvey, P. & Pagel, M. The Comparative Method in Evolutionary Biology. (Oxford University Press, 1991).Huxley, J. S. & Teissier, G. Terminology of relative growth. Nature 137, 780–781 (1936).ADS 
Article 

Google Scholar 
Klingenberg, C. P. Size, shape, and form: Concepts of allometry in geometric morphometrics. Dev. Genes Evol. 226, 113–137 (2016).PubMed 
PubMed Central 
Article 

Google Scholar 
Russell, E. S. Form and Function: A Contribution to the History of Animal Morphology. (John Murray, 1916).Goswami, A. & Polly, P. D. Methods for studying morphological integration and modularity. Paleontol. Soc. Pap. 16, 213–243 (2010).Article 

Google Scholar 
Vidal-García, M., Byrne, P. G., Roberts, J. D. & Keogh, J. S. The role of phylogeny and ecology in shaping morphology in 21 genera and 127 species of Australo-Papuan myobatrachid frogs. J. Evol. Biol. 27, 181–192 (2014).PubMed 
Article 

Google Scholar 
Erwin, D. H. Disparity: Morphological pattern and developmental context. Palaeontology 50, 57–73 (2007).Article 

Google Scholar 
Fišer, C., Robinson, C. T. & Malard, F. Cryptic species as a window into the paradigm shift of the species concept. Mol. Ecol. 27, 613–635 (2018).PubMed 
Article 

Google Scholar 
Wilson, D. E. & Mittermeier, R. A. Handbook of the Mammals of the World: Volume 8: Insectivores. vol. 8 (Lynx Edicions, 2018).Jacquet, F. et al. Phylogeography and evolutionary history of the Crocidura olivieri complex (Mammalia, Soricomorpha): From a forest origin to broad ecological expansion across Africa. BMC Evol. Biol. 15, 71. https://doi.org/10.1186/s12862-015-0344-y (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Ceríaco, L. M. P. et al. Description of a new endemic species of shrew (Mammalia, Soricomorpha) from PrÍncipe Island (Gulf of Guinea). Mammalia 79, 325–341 (2015).Article 

Google Scholar 
Nicolas, V. et al. Multilocus phylogeny of the Crocidura poensis species complex (Mammalia, Eulipotyphla): Influences of the palaeoclimate on its diversification and evolution. J. Biogeogr. 46, 871–883 (2019).Article 

Google Scholar 
Konečný, A., Hutterer, R., Meheretu, Y. & Bryja, J. Two new species of Crocidura (Mammalia: Soricidae) from Ethiopia and updates on the Ethiopian shrew fauna. J. Vertebr. Biol. 69, 20064.1. https://doi.org/10.25225/jvb.20064 (2020).Article 

Google Scholar 
Couvreur, T. L. P. et al. Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biol. Rev. 96, 16–51 (2021).PubMed 
Article 

Google Scholar 
Mayr, E. & O’Hara, R. J. The biogeographic evidence supporting the Pleistocene forest refuge hypothesis. Evolution 40, 55–67 (1986).PubMed 
Article 

Google Scholar 
Wiens, J. J. & Graham, C. H. Niche conservatism: Integrating evolution, ecology, and conservation biology. Annu. Rev. Ecol. Evol. Syst. 36, 519–539 (2005).Article 

Google Scholar 
Smith, T. B., Wayne, R. K., Girman, D. J. & Bruford, M. W. A role for ecotones in generating rainforest biodiversity. Science 276, 1855–1857 (1997).CAS 
Article 

Google Scholar 
Needham, A. E. & Hardy, A. C. The form-transformation of the abdomen of the female pea-crab, Pinnotheres pisum Leach. Proc. R Soc. Lond. Ser. B Biol. Sci. 137, 115–136 (1950).ADS 
CAS 

Google Scholar 
Hanken, J. & Hall, B. K. The Skull, Volume 3: Functional and Evolutionary Mechanisms. (University of Chicago Press, 1993).Hautier, L., Lebrun, R. & Cox, P. G. Patterns of covariation in the masticatory apparatus of hystricognathous rodents: Implications for evolution and diversification. J. Morphol. 273, 1319–1337 (2012).PubMed 
Article 

Google Scholar 
Aristide, L. et al. Multiple factors behind early diversification of skull morphology in the continental radiation of New World monkeys. Evolution 72, 2697–2711 (2018).PubMed 
Article 

Google Scholar 
Hardin, G. The competitive exclusion principle. Science 131, 1292–1297 (1960).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Denys, C. et al. Shrews (Mammalia, Eulipotyphla) from a biodiversity hotspot, Mount Nimba (West Africa), with a field identification key to species. Zoosystema 43, 729–757 (2021).Article 

Google Scholar 
Estevo, C. A., Nagy-Reis, M. B. & Nichols, J. D. When habitat matters: Habitat preferences can modulate co-occurrence patterns of similar sympatric species. PLoS One 12, e0179489. https://doi.org/10.1371/journal.pone.0179489 (2017).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Spaeth, P. A. Morphological convergence and coexistence in three sympatric North American species of Microtus (Rodentia: Arvicolinae). J. Biogeogr. 36, 350–361 (2009).Article 

Google Scholar 
Adams, D. C., Berns, C. M., Kozak, K. H. & Wiens, J. J. Are rates of species diversification correlated with rates of morphological evolution?. Proc. R. Soc. B Biol. Sci. 276, 2729–2738 (2009).Article 

Google Scholar 
Caumul, R. & Polly, P. D. Phylogenetic and environmental components of morphological variation: Skull, mandible, and molar shape in marmots (marmota, Rodentia). Evolution 59, 2460–2472 (2005).PubMed 
Article 

Google Scholar 
Da Silva, F. O. et al. The ecological origins of snakes as revealed by skull evolution. Nat. Commun. 9, 376. https://doi.org/10.1038/s41467-017-02788-3 (2018).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Hirano, T., Kameda, Y., Kimura, K. & Chiba, S. Substantial incongruence among the morphology, taxonomy, and molecular phylogeny of the land snails Aegista, Landouria, Trishoplita, and Pseudobuliminus (Pulmonata: Bradybaenidae) occurring in East Asia. Mol. Phylogenet. Evol. 70, 171–181 (2014).PubMed 
Article 

Google Scholar 
Ge, D., Yao, L., Xia, L., Zhang, Z. & Yang, Q. Geometric morphometric analysis of skull morphology reveals loss of phylogenetic signal at the generic level in extant lagomorphs (Mammalia: Lagomorpha). Contrib. Zool. 84, 267–284 (2015).Article 

Google Scholar 
Zou, Z. & Zhang, J. Morphological and molecular convergences in mammalian phylogenetics. Nat. Commun. 7, 12758. https://doi.org/10.1038/ncomms12758 (2016).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Ananjeva, N. B. Current state of the problems in the phylogeny of squamate reptiles (Squamata, Reptilia). Biol. Bull. Rev. 9, 119–128 (2019).Article 

Google Scholar 
Revell, L. J., Harmon, L. J. & Collar, D. C. Phylogenetic signal, evolutionary process, and rate. Syst. Biol. 57, 591–601 (2008).PubMed 
Article 

Google Scholar 
Klingenberg, C. P. & Marugán-Lobón, J. Evolutionary covariation in geometric morphometric data: Analyzing integration, modularity, and allometry in a phylogenetic context. Syst. Biol. 62, 591–610 (2013).PubMed 
Article 

Google Scholar 
Cardini, A. & Polly, P. D. Larger mammals have longer faces because of size-related constraints on skull form. Nat. Commun. 4, 2458. https://doi.org/10.1038/ncomms3458 (2013).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Esquerré, D., Sherratt, E. & Keogh, J. S. Evolution of extreme ontogenetic allometric diversity and heterochrony in pythons, a clade of giant and dwarf snakes. Evolution 71, 2829–2844 (2017).PubMed 
Article 

Google Scholar 
Marroig, G. & Cheverud, J. M. Size as a line of least evolutionary resistance: Diet and adaptive morphological radiation in New World monkeys. Evolution 59, 1128–1142 (2005).PubMed 
Article 

Google Scholar 
Cornette, R., Tresset, A., Houssin, C., Pascal, M. & Herrel, A. Does bite force provide a competitive advantage in shrews? The case of the greater white-toothed shrew. Biol. J. Linn. Soc. 114, 795–807 (2015).Article 

Google Scholar 
Rodgers, G. M., Downing, B. & Morrell, L. J. Prey body size mediates the predation risk associated with being “odd”. Behav. Ecol. 26, 242–246 (2015).Article 

Google Scholar 
Damuth, J. Population density and body size in mammals. Nature 290, 699–700 (1981).ADS 
Article 

Google Scholar 
Verschuren, D. Decadal and century-scale climate variability in tropical Africa during the past 2000 years. In Past Climate Variability Through Europe and Africa (eds. Battarbee, R. W., Gasse, F. & Stickley, C. E.) 139–158 (Springer Netherlands, 2004). https://doi.org/10.1007/978-1-4020-2121-3_8.Smith, T. B., Schneider, C. J. & Holder, K. Refugial isolation versus ecological gradients. Genetica 112, 383–398 (2001).PubMed 
Article 

Google Scholar 
Brown, W. L. Jr. & Wilson, E. O. Character displacement. Syst. Biol. 5, 49–64 (1956).
Google Scholar 
Vogel, P. et al. Genetic identity of the critically endangered Wimmer’s shrew Crocidura wimmeri. Biol. J. Linn. Soc. 111, 224–229 (2014).Article 

Google Scholar 
Esselstyn, J. A. et al. Fourteen new, endemic species of shrew (genus Crocidura) from Sulawesi reveal a spectacular island radiation. Bull. Am. Mus. Nat. Hist. 454, 1–108 (2021).Article 

Google Scholar 
Evin, A., Bonhomme, V. & Claude, J. Optimizing digitalization effort in morphometrics. Biol. Methods Protoc. 5, bpaa023. https://doi.org/10.1093/biomethods/bpaa023 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Blomberg, S. P., Garland, T. & Ives, A. R. Testing for phylogenetic signal in comparative data: Behavioral traits are more labile. Evolution 57, 717–745 (2003).PubMed 
Article 

Google Scholar 
Adams, D. C. A generalized K statistic for estimating phylogenetic signal from shape and other high-dimensional multivariate data. Syst. Biol. 63, 685–697 (2014).PubMed 
Article 

Google Scholar 
Kembel, S. W. et al. Picante: R tools for integrating phylogenies and ecology. Bioinformatics 26, 1463–1464 (2010).CAS 
PubMed 
Article 

Google Scholar 
Revell, L. J. phytools: Phylogenetic Tools for Comparative Biology (and Other Things). (2021).Fick, S. E. & Hijmans, R. J. WorldClim 2: New 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).Article 

Google Scholar 
Pebesma, E. Simple features for R: Standardized support for spatial vector data. R J. 10, 439 (2018).Article 

Google Scholar 
Oksanen, J. et al. vegan: Community Ecology Package. (2020).Dray, S., Legendre, P. & Peres-Neto, P. R. Spatial modelling: A comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecol. Model. 196, 483–493 (2006).Article 

Google Scholar 
Borcard, D., Gillet, F. & Legendre, P. Numerical Ecology with R. (Springer, 2018).Dray, S. et al. adespatial: Multivariate Multiscale Spatial Analysis. (2021).Collyer, M. & Adams, D. RRPP: Linear Model Evaluation with Randomized Residuals in a Permutation Procedure. (2021).Kassambara, A. rstatix: Pipe-Friendly Framework for Basic Statistical Tests. (2021).Borcard, D., Legendre, P. & Drapeau, P. Partialling out the spatial component of ecological variation. Ecology 73, 1045–1055 (1992).Article 

Google Scholar 
Rohlf, F. J. & Corti, M. Use of two-block partial least-squares to study covariation in shape. Syst. Biol. 49, 740–753 (2000).CAS 
PubMed 
Article 

Google Scholar 
Schlager, S., Jefferis, G. & Ian, D. Morpho: Calculations and Visualisations Related to Geometric Morphometrics. (2020). More