Googling Trends in Conservation Biologycrie.uqtr.ca/pdfs/Proulx_ConsBiol_GoogleTrends_2014.pdf ·...

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Essay Googling Trends in Conservation Biology RAPHA ¨ EL PROULX, PHILIPPE MASSICOTTE, AND MARC P ´ EPINO Centre de recherche sur les interactions bassins versants—´ ecosyst` emes aquatiques (RIVE) and Groupe de recherche interuni- versitaire en limnologie (GRIL), Universit´ e du Qu´ ebec ` a Trois-Rivi` eres, C.P. 500, Trois-Rivi` eres, Qu´ ebec, G9A 5H7, Canada, email [email protected] Abstract: Web-crawling approaches, that is, automated programs data mining the internet to obtain in- formation about a particular process, have recently been proposed for monitoring early signs of ecosystem degradation or for establishing crop calendars. However, lack of a clear conceptual and methodological framework has prevented the development of such approaches within the field of conservation biology. Our objective was to illustrate how Google Trends, a freely accessible web-crawling engine, can be used to track changes in timing of biological processes, spatial distribution of invasive species, and level of public awareness about key conservation issues. Google Trends returns the number of internet searches that were made for a keyword in a given region of the world over a defined period. Using data retrieved online for 13 countries, we exemplify how Google Trends can be used to study the timing of biological processes, such as the seasonal recurrence of pollen release or mosquito outbreaks across a latitudinal gradient. We mapped the spatial extent of results from Google Trends for 5 invasive species in the United States and found geographic patterns in invasions that are consistent with their coarse-grained distribution at state levels. From 2004 through 2012, Google Trends showed that the level of public interest and awareness about conservation issues related to ecosystem services, biodiversity, and climate change increased, decreased, and followed both trends, respectively. Finally, to further the development of research approaches at the interface of conservation biology, collective knowledge, and environmental management, we developed an algorithm that allows the rapid retrieval of Google Trends data. Keywords: biodiversity, ecosystem services, Google Trends, phenology, public awareness, species distribution, web crawling Resumen: Los m´ etodos de navegaci´ on en la red, esto es, programas automatizados de miner´ ıa de datos para obtener informaci´ on de un proceso determinado, han sido propuestos recientemente para monitorear se˜ nales tempranas de la degradaci´ on de ecosistemas o para el establecimiento de calendarios de cosecha. Sin embargo, la falta de un marco conceptual y metodol´ ogico ha prevenido el desarrollo de tales m´ etodos en el campo de la biolog´ ıa de la conservaci´ on. Nuestro objetivo fue ilustrar como Google Trends, una plataforma de rastreo en la red accesible gratuitamente, puede ser utilizado para seguir los cambios de cronolog´ ıa en procesos biol´ ogicos, distribuci´ on espacial de especies invasoras y el nivel de conciencia p´ ublica acerca de temas clave de conservaci´ on. Google Trends reporta el n´ umero de b´ usquedas por internet realizadas para una palabra clave en una regi´ on determinada del mundo en un per´ ıodo definido. Mediante el uso de datos recuperados para 13 pa´ ıses, ejemplificamos como se puede usar Google Trends para estudiar la cronolog´ ıa de procesos biol´ ogicos, como la recurrencia estacional de liberaci´ on de polen o brotes de mosquitos en un gradiente latitudinal. Mapeamos la extensi´ on espacial de los resultados de Google Trends para cinco especies invasoras en Estados Unidos y encontramos patrones geogr´ aficos de invasiones que son consistentes con su distribuci´ on de grano grueso a nivel estatal. De 2004 a 2012 Google Trends mostr´ o que el nivel de inter´es y conciencia del p´ ublico sobre temas de conservaci´ on relacionados con servicios del ecosistema, biodiversidad y cambio clim´ atico incrementaron, disminuyeron y siguieron ambas tendencias, respectivamente. Finalmente, para promover el desarrollo de m´ etodos de investigaci´ on en la interfaz de la biolog´ ıa de la conservaci´ on, el conocimiento colectivo y la gesti´ on ambiental, desarrollamos un algoritmo que permite la r´ apida recuperaci´ on de datos de Google Trends. Paper submitted January 16, 2013; revised manuscript accepted April 12, 2013. 1 Conservation Biology, Volume 00, No. 00, 1–8 C 2013 Society for Conservation Biology DOI: 10.1111/cobi.12131

Transcript of Googling Trends in Conservation Biologycrie.uqtr.ca/pdfs/Proulx_ConsBiol_GoogleTrends_2014.pdf ·...

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Essay

Googling Trends in Conservation BiologyRAPHAEL PROULX, PHILIPPE MASSICOTTE, AND MARC PEPINOCentre de recherche sur les interactions bassins versants—ecosystemes aquatiques (RIVE) and Groupe de recherche interuni-versitaire en limnologie (GRIL), Universite du Quebec a Trois-Rivieres, C.P. 500, Trois-Rivieres, Quebec, G9A 5H7, Canada,email [email protected]

Abstract: Web-crawling approaches, that is, automated programs data mining the internet to obtain in-formation about a particular process, have recently been proposed for monitoring early signs of ecosystemdegradation or for establishing crop calendars. However, lack of a clear conceptual and methodologicalframework has prevented the development of such approaches within the field of conservation biology.Our objective was to illustrate how Google Trends, a freely accessible web-crawling engine, can be used totrack changes in timing of biological processes, spatial distribution of invasive species, and level of publicawareness about key conservation issues. Google Trends returns the number of internet searches that weremade for a keyword in a given region of the world over a defined period. Using data retrieved online for13 countries, we exemplify how Google Trends can be used to study the timing of biological processes, such asthe seasonal recurrence of pollen release or mosquito outbreaks across a latitudinal gradient. We mapped thespatial extent of results from Google Trends for 5 invasive species in the United States and found geographicpatterns in invasions that are consistent with their coarse-grained distribution at state levels. From 2004through 2012, Google Trends showed that the level of public interest and awareness about conservationissues related to ecosystem services, biodiversity, and climate change increased, decreased, and followed bothtrends, respectively. Finally, to further the development of research approaches at the interface of conservationbiology, collective knowledge, and environmental management, we developed an algorithm that allows therapid retrieval of Google Trends data.

Keywords: biodiversity, ecosystem services, Google Trends, phenology, public awareness, species distribution,web crawling

Resumen: Los metodos de navegacion en la red, esto es, programas automatizados de minerıa de datospara obtener informacion de un proceso determinado, han sido propuestos recientemente para monitorearsenales tempranas de la degradacion de ecosistemas o para el establecimiento de calendarios de cosecha. Sinembargo, la falta de un marco conceptual y metodologico ha prevenido el desarrollo de tales metodos en elcampo de la biologıa de la conservacion. Nuestro objetivo fue ilustrar como Google Trends, una plataformade rastreo en la red accesible gratuitamente, puede ser utilizado para seguir los cambios de cronologıa enprocesos biologicos, distribucion espacial de especies invasoras y el nivel de conciencia publica acerca detemas clave de conservacion. Google Trends reporta el numero de busquedas por internet realizadas parauna palabra clave en una region determinada del mundo en un perıodo definido. Mediante el uso de datosrecuperados para 13 paıses, ejemplificamos como se puede usar Google Trends para estudiar la cronologıade procesos biologicos, como la recurrencia estacional de liberacion de polen o brotes de mosquitos en ungradiente latitudinal. Mapeamos la extension espacial de los resultados de Google Trends para cinco especiesinvasoras en Estados Unidos y encontramos patrones geograficos de invasiones que son consistentes con sudistribucion de grano grueso a nivel estatal. De 2004 a 2012 Google Trends mostro que el nivel de interes yconciencia del publico sobre temas de conservacion relacionados con servicios del ecosistema, biodiversidad ycambio climatico incrementaron, disminuyeron y siguieron ambas tendencias, respectivamente. Finalmente,para promover el desarrollo de metodos de investigacion en la interfaz de la biologıa de la conservacion, elconocimiento colectivo y la gestion ambiental, desarrollamos un algoritmo que permite la rapida recuperacionde datos de Google Trends.

Paper submitted January 16, 2013; revised manuscript accepted April 12, 2013.

1Conservation Biology, Volume 00, No. 00, 1–8C© 2013 Society for Conservation BiologyDOI: 10.1111/cobi.12131

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2 Trends in Conservation Biology

Palabras Clave: Biodiversidad, conciencia publica, distribucion de especies, fenologıa, rastreo en la red, servi-cios del ecosistema, Tendencias de Google

Introduction

With the recent advents of highly distributed mobilenetworks and online social platforms, combined withthe establishment of online search engines, access toinformation has never been so extensive and immediate(e.g., Barroso et al. 2003; Butler 2006; Aanensen et al.2009). Paradoxically, gathering data on the distributionand abundance of species at high spatial and temporal res-olution is still a major shortcoming of current ecosystems,or species, monitoring programs (Morisette et al. 2009;Cleland et al. 2012). Online data streams are increasinglybeing used by economists (Vosen & Schmidt 2011; Choi& Varian 2012), politicians (Relly et al. 2012), and epi-demiologists (Carneiro & Mylonakis 2009; Ginsberg et al.2009; Dugas et al. 2012) alike to provide data on marketand public opinion trends or the spread of human infec-tious diseases. However, this continuous stream of freelyavailable data remains underexploited by conservationbiologists, perhaps because the link between biologicalprocesses in nature and data driven by human behavioris not as obvious as in other disciplines (e.g., Martin et al.2012). We used Google Trends, a freely accessible searchengine, to track changes in the temporal pattern (phenol-ogy) of biological processes and the spatial distributionof invasive species.

Google Trends

Google is currently the most-used search engine on theWorld Wide Web; nearly 5 billion queries are submit-ted every day. As a part of the array of Google onlineproducts, Google Trends returns the usage volume of aparticular search term for a specific region of the worldover a defined period. Search-term hits are recorded atthe spatial resolution of individual cities within a region(e.g., France > Bretagne > Brest) and at the temporal res-olution of a week. A query in Google Trends first returnsa world map of the search-term hits per country and amonthly time series of the search-term hits dating backto 2004. By default, the results returned by Google Trendsare rescaled by dividing the search-term hits obtained fora given week by the maximum number of hits obtainedat any moment over the period of interest. Query resultsare accessed by logging into a Google account and down-loading a csv file of the data. Manually downloading themany files generated by entering separate search-termqueries is cumbersome. Hence, we have developed anR package that allows for the rapid retrieval of GoogleTrends data (Supporting Information).

Timing of Biological Processes

Phenology is the study of the causes and consequences ofadvancing or delaying the timing of biological processes,such as plant greening and flowering, pest outbreak, an-imal migration, or breeding time. For example, trendsin the phenology of hundreds of plant species showedearlier spring onset in Europe between 1971 and 2000,advancing at a rate of 2.5 d/decade in response to in-creased air temperature (Menzel et al. 2006). Despitethe importance of phenological records for studying theeffects of climate change on biological processes, cur-rent monitoring programs often only actively follow alimited number of biological processes at a low temporaland spatial resolution. With these limitations in mind,Google Trends may be viewed as a source of up-to-datecollective knowledge about biological processes. Theprecision of Google Trend data for assessing the tim-ing of biological processes was recently demonstratedby Dugas et al. (2012), who reported a high correlation(Pearson’s r = 0.87) between postprocessed GoogleTrends data and clinical cases of confirmed influenza inadults and no apparent time lags between the 2 sources ofinformation.

To illustrate how Google Trends may be used to trackthe phenology of biological processes, we queried thesearch terms mosquitoes and pollen for 4 English speak-ing countries: Australia, Canada, England, and the UnitedStates. Moreover, to provide a geographically compre-hensive picture, we entered the same search terms trans-lated into the official languages of 9 additional countries(in parentheses): mosquitoes and pollen (Brazil, Mexico,Spain); �� and �� (China); moustique and pollen(France); mucken and bluten (Germany); zanzare andpolline (Italy); �� and � (Japan); and and(Thailand). We obtained the timing of biological pro-cesses associated with keywords pollen and mosquitoesin each country by extracting for each year from 2008to 2012, the week associated with the maximum num-ber of search-term hits. Weekly time series of the rel-ative number of search-term hits returned by GoogleTrends revealed recurring temporal patterns for pollenand mosquitoes (Fig. 1). The seasonal timing of these bi-ological processes at the country level is captured by thebroad latitudinal gradient of environmental conditions,at least, for temperate countries of Europe and NorthAmerica (Fig. 2). The temperate countries of Canada,Germany, England, France, United States, and Australiadisplay clear cyclical patterns of search-term hits. In con-trast, such seasonality is difficult to detect in subtropical

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Figure 1. Weekly time series (2008–2012) of the relative number of search-term hits returned by country afterquerying in Google Trends the keywords pollen and mosquitoes (terms were translated into the official languageof the country). A loess smoothing (span of 0.05) was applied to each time series.

or tropical countries such as Brazil, Thailand, and Mex-ico, which are characterized by a large interannual vari-ability in the timing date of both biological processes(large error bars in Fig. 2). Finally, the seasonal inver-sion of pollen release or mosquitoes outbreak eventsin southern (e.g., Australia) versus northern hemispherecountries (e.g., Untied States) is also manifest (Figs.1 & 2).

The phenological trends in a country may be ex-plained by the feedback between people’s physiologi-cal responses to mosquito bites (cutaneous itching) orpollen exposure (allergic reaction) and their motivation

to search for a remedy on the internet. To further in-vestigate this point, we correlated the query results weobtained for mosquitoes to the search-term hits returnedby querying deet and citronella, the 2 main active ingre-dients in most commercial insect-repellent lotions. Wealso correlated the query results returned for pollen withthose for Zyrtec, Claritin, and Reactin, the 3 main allergy-treating drugs sold in Canada and the United States.Using the data from all weeks since 1 January 2008(n = 253), we obtained high Pearson’s coefficients (r)of 0.90 and 0.91 for the United States and 0.88 and 0.87(for Canada) for the mosquitoes and pollen correlations,

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4 Trends in Conservation Biology

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Figure 2. Latitudinal variation in the annual peak ofsearch-term hits returned after querying in GoogleTrends the keywords pollen and mosquitoes for theperiod 2008–2012. For each country, the annual peakis expressed as the mean date (SD) of search-hitmaximum reported each year starting on 1November. Latitude coordinates were obtained onlinefrom the NationMaster database, with the exception ofCanada, which was assigned a value of 45◦N to reflectthe strong southward asymmetry in the repartition ofhuman population centers.

respectively, a result that suggests a causal relationbetween the seasonal recurrence of pollen release ormosquito outbreak and people’s behavioral responses tothese 2 processes.

Spatial Distribution of Invasive Species

Biodiversity and economic losses caused by newly intro-duced, rapidly spreading non-native species, is an issue ofgrowing concern in conservation biology, most notablybecause globalization, and in particular increased inter-

national trading, accelerates the dispersion of speciesaround the world (Puth & Post 2005; Crowl et al. 2008;Pysek et al. 2010). Early detection of invasive non-nativespecies is therefore a fundamental component of pre-venting their establishment and spread. Because at theirintroduction populations are generally composed of fewindividuals, the initial stage of dispersion is a critical steptoward the establishment of a non-native invasive species(Puth & Post 2005; Blackburn et al. 2011). However,governmental agencies often lack the resources to detectspecies introductions at their early stage of dispersion.Moreover, in cases of established invasive species, a con-siderable amount of public and scientific resources areinvested annually to monitor the distribution of thesepopulations. Galaz et al. (2010) proposed combiningweb-crawling and expert-knowledge approaches for theearly detection of ecosystem change or degradation. Inthe context of our study, a first step in that directionis to determine whether web crawlers, such as GoogleTrends, can be used to map the spatial distribution ofinvasive species at the country level.

Google Trends spatially disaggregates the volume ofreturned search-term hits at the level of cities or regionswithin a country. To illustrate this feature and its potentialto address the shortfalls of species detection and tracking,we mapped the distribution of search-term hits for 5 in-vasive species in the United States. We entered in GoogleTrends the following search-term queries: ash borer(Agrilus planipennis), Asian carp (Cyprinus carpio,Hypophthalmichthys molitrix, Hypophthalmichthys no-bilis, Mylopharyngodon piceus), fire ants (Solenopsisinvicta), Africanized bees (Apis mellifera scutellata),and pine beetle (Dendroctonus ponderosae). The result-ing maps show the relative volume of search terms re-turned by state for each of the search terms we queried(Fig. 3). The large number of search-term hits in statesbordering the Great Lakes for the emerald ash borer(Haack et al. 2002) and in the upper Mississippi basinfor the Asian carp (Koel et al. 2000) reflects their re-spective areas of origin and current dispersion range inthe United States. In the case of Asian carp, the elevatednumber of search-term hits in the northernmost statesalso implies apprehension about the invasion of GreatLakes by these fish. The search-term distribution of fireants and Africanized bees reflects their introduction anddispersal patterns; they were first imported to southernUnited States in the mid-1990s and have since dispersedto neighboring states (Woodward & Quinn 2011). Fi-nally, it is well documented that current pine beetleoutbreaks have occurred and continue to spread east-ward across the Rocky Mountain states such as Montana,Wyoming, and Colorado (Evangelista et al. 2011) (Fig. 3).A more thorough validation of Google Trends maps isbeyond the scope of this essay, but will be needed infuture applications (see also “Google Trends Limitations”below).

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Figure 3. Spatial distribution of the relative number of search-term hits returned for each U.S. state of theconterminous United States after querying 5 invasive species keywords in Google Trends: Africanized bees, Ashborer, Asian carp, Fire ants, and Pine beetle. The numbers have been scaled such that the state with the maximumof search-term hits has a value of 100. The white line was drawn from the reference spatial distribution of eachspecies available at the following websites: Africanized bees, www.nationalatlas.gov/mld/afrbeep; ash borer,www.aphis.usda.gov/plant_health/plant_pest_info/emerald_ash_b; Asian carp,nas.er.usgs.gov/queries/speciesmap.aspx?SpeciesID=551; fire ants,www.aphis.usda.gov/plant_health/plant_pest_info/fireants; pine beetle,www.barkbeetles.org/mountain/fidl2.htm.

Public Awareness of Conservation Issues

A central objective of conservation biology is to ensurethat best management practices, or environmental threatsto biodiversity, are efficiently communicated to decisionmakers and stakeholders (Malcevschi et al. 2012). Pub-lic awareness is often a key component of conservationagendas because the public may not only be stakeholdersthemselves, but they may also have the power to influ-ence decision makers. In this light, 2 international panelshave been established by the United Nations to improvecommunication among the public, conservation biolo-gists, and policy makers: the Intergovernmental Panel onClimate Change (IPCC) and the Intergovernmental Panelof Biodiversity and Ecosystem Services (IPBES). Thesepanels provide an interface between scientists and pol-icy makers in order to better inform the larger commu-nity (e.g., parties involved, stakeholders, and the public)through the publication of periodic reports. Thus, if pub-lic interest and awareness is a key ingredient in achievingconservation goals, it leads to the question: are climatechange-, biodiversity-, and ecosystem-related issues pro-

gressively garnering more public attention as they be-come more important, or is interest waning over time?

To illustrate how Google Trends can be used to trackchanges in the level of public interest and awarenessabout key conservation issues, we entered the keywordsclimate change, biodiversity, and ecosystem services andgraphed their temporal trends between 2004 and 2012(Fig. 4). Recent conservation issues, such as the ecosys-tem services concept, are on an overall increasing searchtrend in English-speaking countries, whereas relativelyolder conservation issues, such as those related to climatechange, are attracting less attention since 2008 (Fig. 4).Moreover, the relative search-term volume of biodiversitystopped declining after 2010, which incidentally was de-clared the international year of biodiversity. Although in-terpreting the human-driven motives behind such broadtemporal trends is far from trivial, the sole existence ofa trend should be seriously considered because GoogleTrends data are routinely corrected for the total num-ber of web queries made over a given week. Hence,the observed temporal trend in the number of searchhits cannot be attributed to baseline changes in the total

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6 Trends in Conservation Biology

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Figure 4. Weekly time series (2004–2012) of the relative number of search-term hits returned after querying inGoogle Trends the keywords biodiversity, climate change, and ecosystem services. A loess smoothing (span of 30)was applied to each time series to extract the long-term trends of these conservation issues.

number of people searching the web. For instance, al-though the number of persons actively searching the webhas substantially increased since 2004, we verified thatcommon search terms such as weather and news do notshow temporal trends over the 2004–2012 period. Weconducted this verification by entering the 2 keywords(weather and news) in Google Trends and observed notendency.

Google Trends Limitations

First, online keyword queries in Google Trends within acountry are sent from highly populated cities, which donot form a representative (spatially extensive, random,unbiased) sample of a region. Second, one cannot knowthe real motives behind each internet search recorded byGoogle Trends. For example, we did not know whethersearch-term queries returned for Asian carp were en-

tered by anglers, researchers studying the topic, or byweb surfers looking for a popular Asian carp video.Third, temporal or spatial patterns may be mistakenlyinterpreted as being driven by biological processes. Forexample, the Google Trends search-term volume in theUnited States for pollen correlated identically (Person’sr = 0.85) to both plant flowers and pine straw searchhits. Although there may be a direct causal associationbetween pollen and flowering plants, the link betweenpollen and pine straw is more tenuous. Although substan-tial, some of these limitations could be counterbalanced ifkeyword queries are crossvalidated so that they all relateto the same process (e.g., correlating search hits betweenpollen and plant flowers) or if search trends of irrelevantkeywords were removed (e.g., Asian carp youtube). Thisis what the search engine Google Flu does. In Google Flu,a list of associated search terms are used to estimate sea-sonal trends in the progression of influenza cases (Dugaset al. 2012).

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Table 1. Main advantages of using Google Trends over conventionalfield-monitoring programs for tracking changes in the timing of bio-logical processes (T), distribution of invasive species (D), and level ofpublic awareness about conservation issues (P).

Added value of Google Trends Associated examples

Cost-effective P, D, TRapid assessment P, D, THigh temporal resolution P, D, TStandardized protocol P, D, TMultiple spatial scales (local,

regional, global)P, D, T

Google Trend Advantages

A thoughtful selection and thorough consideration ofkeywords apropos of a research question, cultural her-itage, and regional differences are therefore the mostfundamental steps in this analytical approach (Al-Eroudet al. 2011; Al-Kabi et al. 2012). Once the keywords as-sociated with a particular biological process are defined,temporal and spatial trends for a region can be validatedby the governmental agencies or research laboratoriesthat own the data. For example, pollen release is mea-sured as part of public-health monitoring programs, andscientific protocols are routinely established by firmsspecialized in mosquito control. However, in an era ofopen access, validation, testing, and use of web-crawlingapproaches is not limited to data owners. Because theinformation returned by Google Trends is disaggregatedat the city level (Supporting Information), integrating itsresults with global or regional data sets is a straightfor-ward operation. If one uses cities’ geographic coordi-nates as an anchor point, Google Trends results can bematched to georeferenced data sets on, for example, cli-mate (Climate Research Unit), topography (Shuttle RadarTopography Mission), land cover, and land use (NationalAeronautics and Space Administration’s [NASA] EarthObserving System), species and ecosystem conservationstatus (World Wildlife Fund; International Union for Con-servation of Nature), and socioeconomic data (NASA’sSocioeconomic Data and Applications Center).

Google Trends offers several advantages over conven-tional field-monitoring programs for tracking changes intiming of biological processes, distribution of invasivespecies, and level of public awareness about conservationissues (Table 1). The list of questions that conservationbiologists may tackle with results obtained through in-ternet searches of specific keywords is potentially end-less. Questions associated with the effect of climate (e.g.,climate warming, drought severity) on advancing or de-laying the timing of biological processes could be ad-dressed with such data (Sherman-Morris et al. 2011; vander Velde et al. 2012). The mismatch hypothesis (Durantet al. 2005; Thackeray et al. 2010; Donnelly et al. 2011),which stipulates that the functioning of populations and

communities is impaired if biological processes becomeless synchronized over time, could also be tested moreextensively. Moreover, governmental agencies may haveinterest in monitoring the invasion front of a recentlyintroduced species. What is more, they may want topublish maps showing where the temporal trends of aspecies search term is increasing, decreasing, or remainsstable and use this information to take actions. Increasingpublic awareness would also lead to an increased volumeof internet searches, thus strengthening the use of web-crawling approaches for studying conservation issues.

Acknowledgments

We thank all the graduate students and scientists at theRIVE who enjoy thinking outside the box. We thank I.Seiferling, Y. Paradis, and students from the Geomaticsand Landscape Ecology Laboratory (Carleton University)for providing engaged comments on the manuscript. R.P.,P.M., and M.P. participated in the writing of this essay.R.P. and M.P. contributed the figures, and P.M. has devel-oped the R package for automatically retrieving GoogleTrends data. This research was supported by a grant fromThe Natural Sciences and Engineering Research Councilof Canada (NSERC).

Supporting Information

Googling Trends in Conservation Biology Using R(Appendix S1) is available online. The authors are solelyresponsible for the content and functionality of thesematerials. Queries (other than absence of the material)should be directed to the corresponding author.

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