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References & Scientific Sources

Fog, Climate, and Atmospheric Structure (Atacama Desert)

UBöhm, C., Garreaud, R., Rondanelli, R., et al. (2021).
Toward a climatology of fog frequency in the Atacama Desert via multispectral satellite data and machine learning techniques.
Journal of Applied Meteorology and Climatology, 60(7), 941–961.

  

Cáceres, L., Gómez-Silva, B., Garró, X., Rodríguez, V., Monardes, V., & McKay, C. P. (2007).
Relative humidity patterns and fog water precipitation in the Atacama Desert and biological implications.
Journal of Geophysical Research: Biogeosciences, 112, G04S14.

  

Cereceda, P., Osses, P., Larrain, H., Farías, M., & Schemenauer, R. S. (2008).
The occurrence of fog in the Atacama Desert (Chile).
Journal of Arid Environments, 72(11), 2043–2055.

  

Cereceda, P., & Larraín, H. (2008).
The spatial and temporal variability of fog and its relation to fog oases in the Atacama Desert, Chile.
Atmospheric Research, 87, 312–323.


Cordero, R. R., Damiani, A., Seckmeyer, G., et al. (2016). 

The Solar Spectrum in the Atacama Desert. 

Scientific Reports, 6, 22457. 


Cordero, R. R., Damiani, A., Jorquera, J., Sepúlveda, E., Caballero, M., Fernandez, S., Feron, S., Llanillo, P. J., Carrasco, J., Laroze, D., & Labbe, F. (2018).
Ultraviolet radiation in the Atacama Desert.
Antonie van Leeuwenhoek, 111, 1301–1312. 


Corvo, F., Pérez, A., Dzib, T., et al. (2008).
Atmospheric moisture availability and surface wetting processes in the Atacama Desert.
Corrosion Science, 50(8), 2202–2213. 


del Río, C., García, J.-L., Osses, P., et al. (2018).
ENSO influence on coastal fog-water yield in the Atacama Desert, Chile.
Atmospheric Research, 214, 162–175.


Fonseca, A., Ferreira, P. H., Nascimento, D. C., Fiaccone, R., Ulloa-Correa, C., García-Piña, A., & Louzada, F. (2021).

Water particles monitoring in the Atacama Desert: SPC approach based on proportional data.

Axioms, 10(3), 154.


Garreaud, R., Molina, A., Farias, M. (2010).
Andean uplift, ocean cooling and Atacama hyperaridity.
Climate of the Past, 6, 723–739.

  

Larraín Barrios, B. C. (2007).
Relaciones florísticas entre oasis de neblina del desierto costero del norte de Chile.
Memoria de Título, Universidad de Chile, Facultad de Ciencias Agronómicas.


Ibáñez, S. T., Muñoz-Schick, M., & Scherson, R. A. (2022). 

A new species of Diplostephium (Asteraceae, Astereae) from the Atacama Desert, Chile. 

PhytoKeys, 196, 105–125. 


Moat, J., Smith, P., Chilvers, M., et al. (2021).
Fog frequency, cloud cover, and vegetation patterns in the coastal Atacama Desert.
Global Ecology and Biogeography, 30(5), 1008–1022.

  

Muñoz, R. C., Falvey, M. J., Arancibia, M., Astudillo, V. I., 

Elgueta, J., Ibarra, M., Santana, C., & Vásquez, C. (2018).

Wind energy exploration over the Atacama Desert:

A numerical model–guided observational program.

Bulletin of the American Meteorological Society, 99(10), 2079–2092.


Rundel, P. W., Dillon, M. O., Palma, B., Mooney, H. A., Gulmon, S. L., & Ehleringer, J. R. (1991).

The phytogeography and ecology of the coastal Atacama and Peruvian Deserts.

Aliso: A Journal of Systematic and Evolutionary Botany, 13(1), 1–49.


Schulz, N., Aceituno, P., & Richter, M. (2011).

Phytogeographic divisions, climate change and plant dieback along the coastal desert of northern Chile. 

Erdkunde, 65(2), 169–187.

Geology, Mineralization, and Substrate Chemistry (Northern Chile)

Servicio Nacional de Geología y Minería (SERNAGEOMIN). (2003).
Mapa Geológico de Chile.
Government of Chile.


Arancibia, G., Clark, A. H., Farías, M., et al. (2014).
Supergene mineralization processes in the Coastal Cordillera of northern Chile.
Economic Geology, 109, 1907–1936.

  

Casanova, M., Salazar, O., Seguel, O., & Luzio, W. (2013).
The soils of Chile.
Springer Science & Business Media.


Lopez, B. R., Bashan, Y., Bacilio, M., De la Cruz-Agüero, G. (2009).
Rock-colonizing plants: abundance of the endemic cactus Mammillaria fraileana related to rock type in the southern Sonoran Desert.
Plant Ecology, 201, 575–588. 


Sebastián-Perroud, J. (2015).
Geología y evolución tectónica de la Cuenca de Chañarcillo, Chile.
Undergraduate thesis, Universidad de Chile. 


Villalobos-Amador, E., et al. (2019).
Origen y naturaleza de la mineralización supergénica de cobre en el depósito Barreal, Chile.
Revista Geológica de Chile.

Vegetation Diversity, Endemism, and Conservation (Atacama Region)

Bobo-Pinilla, J., Salmerón-Sánchez, E., Mendoza-Fernández, A. J., Mota, J. F., & Peñas, J. (2022).

Conservation and phylogeography of plants: From the Mediterranean to the rest of the world.

Diversity, 14(2), 78.


Díaz-Siefer, P., Gutiérrez, J. R., & Squeo, F. A. (2023). 

Native plant seed systems and restoration bottlenecks in Chile: Implications for biodiversity conservation. 

Restoration Ecology, 31(4), e13782. 


Gómez-Silva, B., & Rojas-Pallero, J. (2005). 

Jardín Botánico del Desierto (JBD): Una herramienta de extensión e investigación sobre los recursos naturales renovables del Desierto de Atacama. 

Chloris Chilensis, 8(2).


Letelier, L., Squeo, F. A., Arancio, G., Marticorena, A., Muñoz-Schick, M., Arroyo, M. T. K., León-Lobos, P., Montecinos, S., & Gutiérrez, J. R. (2008).
Diversidad vegetal de la Región de Atacama, Chile.
In Libro Rojo de la Flora Nativa y de los Sitios Prioritarios para su Conservación: Región de Atacama (pp. 123–135).
Ediciones Universidad de La Serena, La Serena, Chile. 


Ortega-Baes, P., Galíndez, G., Sühring, S., Rojas-Aréchiga, M., & Cuevas, E. (2010). 

Thermal buffering capacity of the germination niche in cacti: Implications for species persistence under climate change. 

Journal of Arid Environments, 74(7), 810–816. 

CAM Photosynthesis and Desert Plant Physiology

Lüttge, U. (2004).
Ecophysiology of Crassulacean Acid Metabolism (CAM).
Annals of Botany, 93, 629–652.


Osmond, C. B. (1978).
Crassulacean acid metabolism: a curiosity in context.
Annual Review of Plant Physiology, 29, 379–414.


Winter, K., Smith, J. A. C. (1996).
Crassulacean Acid Metabolism: Biochemistry, Ecophysiology and Evolution.
Springer, Berlin. 

Light Measurement, Spectral Metrics, and Photobiology

Apogee Instruments.
Technical documentation on extended photosynthetically active radiation (ePAR) and quantum sensor spectral response (400–750 nm).


Bugbee, B. (Utah State University), Crop Physiology Laboratory.
Research on extended photosynthetically active radiation (ePAR) and far-red effects in plant lighting (c. 2020–present).

  

McCree, K. J. (1972).
The action spectrum, absorptance and quantum yield of photosynthesis in crop plants.
Agricultural Meteorology, 9, 191–216.


Taiz, L., Zeiger, E., Møller, I. M., & Murphy, A. (2015).
Plant Physiology and Development. 

Sinauer Associates, Sunderland, MA. 


Zhen, S., & Bugbee, B. (2020).
Far-red photons have equivalent efficiency to traditional photosynthetic photons: Implications for redefining photosynthetically active radiation.
Plant, Cell & Environment.

  

Zhen, S., & Bugbee, B. (2020)
Substituting far-red for traditionally defined photosynthetic photons results in equal canopy quantum yield for CO₂ fixation and increased photon capture during long-term studies.

Frontiers in Plant Science, 11:581156.  


Zhen, S., van Iersel, M., & Bugbee, B. (2021).
Why far-red photons should be included in the definition of photosynthetic photons and the measurement of horticultural fixture efficacy.
Frontiers in Plant Science, 12:693445.  

Cactus Morphology, Spines, and Functional Anatomy

Aliscioni, N. L., Delbón, N. E., & Gurvich, D. E. (2021).
Spine function in Cactaceae: a review.
Journal of the Professional Association for Cactus Development, 23, 1–24. 


de la Rosa-Manzano, E., Flores, J., Delgado-Sánchez, P. (2016).

Effects of spine shading on photosynthesis in three cactus species.

Botanical Sciences, 94(2), 301–310.


F.T. Malik, et al. (2016).
Hierarchical structures of cactus spines that aid in the directional movement of dew droplets.
Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences, 374(2073), 20160184.

  

Loik, M. E. (2008).

The effect of cactus spines on light interception and Photosystem II for three sympatric species of Opuntia from the Mojave Desert.

Physiologia Plantarum, 134, 87–98.


Martinez, Stewart & Szeto (2017)
The Relationship Between Structural Parameters and Mechanical Properties of Cactus Spines.
B.S. Materials Engineering thesis, Cal Poly 


Mauseth, J. D. (2005).
Anatomical adaptations to xeric conditions in cacti.
Cactus and Succulent Journal (U.S.), 77(4), 198–208. 


Mauseth, J. D. (2006).
Structure–function relationships in highly modified shoots of Cactaceae.
Annals of Botany, 98, 901–926.


Mauseth, J. D. (2013).
Spines and glochids of cacti: structure, development, and function.
Bradley Journal of Botany, 2, 45–67.


Miesen, F., de Porras, M. E., & Maldonado, A. (2015).

Pollen morphology of Cactaceae in Northern Chile.

Gayana Botánica, 72(2), 258–271.


Nobel, P. S. (2002).
Cactus Biology and Uses.
University of California Press.

  

Nolasco, H., Vega-Villasante, F., Díaz-Rondero, A., & Flores, J. (1997).
Spine and trichome density in cacti as an adaptation to drought.
Journal of Arid Environments, 35(2), 251–258.


Nyffeler & Eggli (1997)

Comparative Stem Anatomy and Systematics of Eriosyce sensu lato

Annals of Botany

  

Riglet, L., Gatti, S., & Moyroud, E. (2021). 

Sculpting the surface: Structural patterning of plant epidermis. 

iScience, 24, 103346.


Reyes-Rivera J, Solano E, Terrazas T, Soto-Hernández M, Arias S, Almanza-Arjona YC, Polindara-García LA

Classification of lignocellulosic matrix of spines in Cactaceae by Py-GC/MS combined omic tools and multivariate analysis: a chemotaxonomic approach. Journal of Analytical and Applied Pyrolysis (2020) 


Warren, S. D., Aguilera, L. E., & Baggett, L. S. (2016).

Directional orientation of reproductive tissue of Eulychnia breviflora (Cactaceae) in the hyperarid Atacama Desert. 

Revista Chilena de Historia Natural, 89, 10.

Thermal Regulation, Albedo, and Microclimate

Geller, G. N., Nobel, P. S. (1984).
Influence of trichomes and spines on boundary layer properties of desert plants.
Oecologia, 64, 1–8.


Nobel, P. S. (1988).
Environmental Biology of Agaves and Cacti.
Cambridge University Press.

Microbial Ecology, Lithic Hydration, and Endolithic Systems

Castro et al. 2022. 

Soil microbiome influences on seedling establishment and growth of Prosopis chilensis and Prosopis tamarugo from northern Chile 

Plants 2022, 11, 2717


Crits-Christoph, A., et al. (2013).
Colonization patterns of lithic microbial communities in the Atacama Desert.
Frontiers in Microbiology, 4, 1–14.


Huang, S., Li, Y., Wu, X., et al. (2020).
Microbial extraction of structural water from gypsum in hyper-arid environments.
Proceedings of the National Academy of Sciences (PNAS), 117(19), 10602–10608.

  

Prisa, D. (2021). 

Biochar effects in the growing and control of biotic and abiotic stress in Astrophytum myriostigma and Astrophytum capricorne. 

GSC Biological and Pharmaceutical Sciences, 16(1), 186–194.


Sánchez-Reinoso, A. D., Ávila-Pedraza, E. A., & Restrepo-Díaz, H. (2020). 

Use of biochar in agriculture. 

Acta Biológica Colombiana, 25(2), 327–338.


Santiago, I. F., Gonçalves, V. N., Gómez-Silva, B., Galetovic, A., & Rosa, L. H. (2018).
Fungal diversity in the Atacama Desert.
Antonie van Leeuwenhoek, 111, 1345–1361. 


Vásquez-Dean, J., et al. (2020).
Microbial and mineralogical patterns in the hyper-arid Atacama Desert.
Scientific Reports, 10(1), Article 1–13.


Wierzchos, J., Ascaso, C., McKay, C. P. (2012).
Endolithic microbial habitats as refuges in hyper-arid deserts.
Antonie van Leeuwenhoek, 101, 1–16.


Yadav, S. P. S., Bhandari, S., Bhatta, D., Poudel, A., Bhattarai, S., Yadav, P., Ghimire, N., Paudel, P., Paudel, P., Shrestha, J., & Oli, B. (2023). 

Biochar application: A sustainable approach to improve soil health. 

Journal of Agriculture and Food Research, 11, 100498. 

Fog-Dependent Nutrient Cycling and Atmospheric Inputs

Ewing, S. A., et al. (2008).
Fog deposition and nutrient cycling in hyper-arid ecosystems.
Biogeochemistry, 87, 53–64.

  

Fletcher, L. A., Parro, V., Gómez, F., et al. (2012).
Variability of organic material in surface horizons of hyper-arid Atacama Desert soils.
Geochimica et Cosmochimica Acta, 87, 30–44.

  

González, A. L., Farina, J. M., Pinto, R., Pérez, C., Weathers, K. C., Armesto, J. J., & Marquet, P. A. (2011).
Bromeliad growth and stoichiometry: responses to atmospheric nutrient supply in fog-dependent ecosystems of the hyper-arid Atacama Desert, Chile. 

Oecologia, 167, 835–845.

  

Pinto, R., Barría, I., Marquet, P. A., & Jaksic, F. M. (2006).
The role of fog in the maintenance of Tillandsia landbeckii populations in the Atacama Desert, Chile.
Journal of Arid Environments, 65(4), 558–567.


Weathers, K. C., et al. (2010).
Atmospheric deposition and canopy interactions in fog-dominated systems.
Ecosystems, 13, 767–780.

Copiapoa-Specific, Historical, and Cultivation Sources

Cayo, M., Solís-Cornejo, F., Santos, A., Zamorano, P., & Valenzuela, B. (2025).
The abundance and distribution of the acdS gene in microbial communities from the rhizosphere of Copiapoa solaris, a native cactus in the arid coastal region of Antofagasta, Chile.
Microorganisms, 13, 1547. 


Charles, G. (1998).
Copiapoa and their cultivation.
British Cactus & Succulent Society.


Doweld, A. B. (2001). 

On the phylogeny and systematics of the genus Copiapoa Britton et Rose. Succulents (CYKKY JIEHTbl), 1–2, 46–56.


Ehleringer, J. R., Mooney, H. A., Gulmon, S. L., & Rundel, P. W. (1980).

Orientation and its consequences for Copiapoa (Cactaceae) in the Atacama Desert.

Oecologia, 46, 63–67. 


Flores, J., Jurado, E., & Chapa-Vargas, L. (2011). 

Germination of Cactaceae: A review of germination ecology, seed traits, and environmental requirements. 

Seed Science Research, 21(3), 1–15. 

  

Ritter, F. (1980).
Kakteen in Südamerika, Band 3: Chile.
Spangenberg Verlag.


Sarnes, J. (2025).
Copiapoa: The Complete Monograph.
Self-published / Private edition. 


Schulz, R., Kapitany, A. (1994).
Copiapoa.
Cactus & Co.


Señoret, F., & Acosta, J. P. (2013).
Cactáceas Nativas de Chile: Guía de Campo.
Corporación Chilena de la Madera (CORMA), Concepción, Chile. 

ISBN 978-956-8398-06-4. 

  

Stone, G. (2014).
Copiapoa: The extreme end of the scale.
Cactus and Succulent Journal (U.S.), 86(3).


Panco, D. (2013).
The Stone Eaters.
Xerophilia, Issue 2.


Tkachenko, H. (2020).
Latent time characteristic of some species of Cactaceae.
Ukrainian Journal of Ecology, 10(3), 114–120.

  

Evolutionary and Phylogenetic Context  


Hernández-Hernández, T., Brown, J. W., Schlumpberger, B. O., Eguiarte, L. E., & Magallón, S. (2014).
Beyond aridification: multiple explanations for the elevated diversification of cacti in the New World Succulent Biome.
New Phytologist, 202(4), 1382–1397.


Larridon, I., Walter, H. E., Eggli, U., Ogburn, R. M., & Moore, M. J. (2014).
Is there a future for the Cactaceae genera Copiapoa, Eriosyce and Eulychnia?
Biodiversity and Conservation, 23, 1249–1287.


Larridon, I., Walter, H. E., Eggli, U., Ogburn, R. M., & Moore, M. J. (2015).
An integrative approach to understanding the evolution and diversity of Copiapoa.
American Journal of Botany, 102(11), 1853–1867.


Larridon, I., et al. (2018).

Investigating taxon boundaries and extinction risk in endemic Chilean cacti (Copiapoa subsection Cinerei, Cactaceae) using chloroplast DNA sequences, microsatellite data and 3D mapping. 

Kew Bulletin, 73: 55. 

Documentation, Provenance, and Collection Data Integrity

Davis, T. J., & Pillet, M. D. (2023).
Don’t Tell Me, Show Me: The Importance of Managing Collection Data.
Cactus and Succulent Journal (U.S.), 95(3).

  

Korotkova, N., Borsch, T., Arias, S., Barthlott, W., Hunt, D. R., Taylor, N. P., & Liede-Schumann, S. (2021).
Cactaceae at Caryophyllales.org: A dynamic online species-level taxonomic backbone for the family.
Willdenowia, 51(2), 251–270.

Interpretation Note

This site synthesizes peer-reviewed research across climatology, geology, plant physiology, and desert ecology. Where Copiapoa-specific experimental data are limited, broader cactus and fog-desert literature is used conservatively to interpret recurring morphological and ecological patterns. 

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