Research Paper

INTRODUCTION

The order Procellariiformes is among the most endangered seabird groups with the highest number of threatened species of marine birds globally (Croxall et al. 2012, as cited in Rodríguez et al. 2019, Norambuena et al. 2021). Procellariiformes, in general, are highly adapted to the marine environment and spend most of their lives in the open oceans (Rodríguez et al. 2019). They travel long distances for foraging, migration, and dispersal, mostly returning to land only for breeding or at night (Silva et al. 2016, Sausner et al. 2016, Rodríguez et al. 2019).

Within the Procellariiformes order, storm-petrels (Hydrobatidae family) are among the most poorly-known seabird groups (Rodríguez et al. 2019). The limited knowledge of the biology and ecology of storm-petrels, combined with the absence of accurate population estimates, is largely due to their nocturnal, cryptic, and burrow-nesting behavior, as well as the unknown or inaccessible locations of their colonies, which are often found on cliffs or remote islands (Rodríguez et al. 2017, 2019). Of the 22 identified species within the Hydrobatidae family (Nunn and Stanley 1998, Tree of Life Web Project 2008), 12 have declining population trends, and four prove to be Data Deficient (IUCN 2022). Thus, it is important to identify threats and their impact on storm-petrel populations (Norambuena et al. 2021).

Among the documented land-based and marine threats for storm-petrels (e.g. invasive alien species, human disturbance, residential and community development, pollution, mining, and energy farms; Dias et al. 2019; Norambuena et al. 2021), one of the globally increasing threats is light pollution (Kyba et al. 2017). Light pollution is defined as the use of artificial illumination that disrupts natural darkness and the visibility of the night sky (Gaston et al. 2013, Falchi et al. 2016). With the global rise in artificial light due to expanding human settlements driven by population growth (Kyba et al. 2017, United Nations 2019), the ongoing expansion of population centers is expected to escalate light pollution and its effects on storm-petrel populations.

The presence of Artificial Lights at Night (ALAN) may result in the light-induced disorientation of seabirds (i.e., “fallout”), particularly among the youngest. On their first flight away from the colony, fledglings become disoriented by the presence of anthropogenic sources of light (Miles et al. 2010, Wilhelm et al. 2021). As fledglings approach and fly over light sources, they can fall to the ground due to physical exhaustion or collision with man-made structures (Troy et al. 2011), hereby referred to as “grounding.” Storm-petrel fledglings that survive the fall may still die due to an increased risk of predation, starvation, dehydration, drowning, or even collisions with motorized vehicles (Rodríguez et al. 2012, Rodríguez et al. 2014, Rodríguez et al. 2017). Although fallout is considered an important source of mortality for storm-petrel populations (Rodríguez et al. 2017), its mechanism is still unknown. The main hypothesis is that light pollution may outshine natural celestial cues, interfering with storm-petrel navigation systems (Rodríguez et al. 2017). Additional hypotheses like foraging inexperience and an immature visual system have been formulated, but the information is scarce and inconclusive (Imber et al. 1975, Atchoi 2020).

Given the scarce information on the impacts of light pollution on storm-petrel populations, we conducted a scoping review to determine the main impacts of light pollution in the Hydrobatidae family. Through the scoping review, we aim to answer the following research question: What is the current state of evidence on the effects of light pollution on bird populations of the Hydrobatidae family? We expect this review will contribute to the inputs required for the development of adequate management and conservation plans for storm-petrels.

METHODS

For the scoping review, we limited our efforts to two academic platforms, Scopus and Web of Science. These platforms contain a wide array of scientific content on the selected subject.

Keyword selection and identification of relevant articles

We limited our search to keywords in two categories: (1) storm-petrel taxa, according to the Tree of Life Project (Nunn and Stanley 1998, Tree of Life Web Project 2008; Appendix 1), and (2) Light pollution (Table 1). All searches were conducted in English, considering that a first exploratory search in Spanish using the SciELO library catalog returned no results. During our search, only one full-text article (Silva et al. 2020) was found in Spanish, although its information in the library catalog was obtained in English. No restrictions were applied on geographical location.

Article selection process and eligibility criteria

We carried out the literature search on February 16, 2023. Articles published between 01/01/2000 and 31/12/2022 were compiled and uploaded to the Rayyan reference manager (Mourad et al. 2016) for duplicate detection. Articles labeled as duplicates (by the default setting in Rayyan) were individually reviewed by two authors (PS, RM) and removed from the analysis. Article titles and abstracts were screened and excluded if they did not explicitly include light pollution and its effects on Hydrobatidae species or only mentioned unspecified seabirds (Fig. 1). This first screening process was executed by two authors (PS, RM), although in case of discrepancies, all authors discussed whether to exclude or include the article in the next exclusion stage.

The full-text screening was performed using the Zotero reference manager version 6.0.4 (Zotero n.d.). Original research papers that included both light pollution and its effects on Hydrobatidae species were included. Review articles were excluded in this process, except for Silva et al. (2020), as some of their information was published for the first time. At this stage, we also incorporated relevant articles identified within the bibliographies of the originally identified articles. We selected articles from the bibliographies based on title, abstract, and full-text screenings. All studies included in this stage were used for the analysis and literature synthesis presented in the results section.

Data management and result summarizing

A Microsoft Excel version 16.0. spreadsheet (Microsoft Corporation 2021) was created with all the articles in Zotero after duplicate removal (Appendix 2). Information presented for each article includes publication year, title, journal, DOI, abstract, volume, issue, pages, library catalog, link attachment, status, exclusion stage, and exclusion criteria. Table 2 presents a summary of the relevant content of the included articles.

RESULTS

Literature profile and article characteristics

A total of 144 records were identified; 142 were from the database search, and two were added from the bibliographies of articles found in the original search (Appendix 3). After screening titles and abstracts, 20 articles were reviewed in full. Of these, 12 articles published between 2005 and 2022 were included in the final analysis, with half published between 2020 and 2022. Of the 12 articles, six presented data from the southern hemisphere and six from the northern hemisphere. All articles extracted data from locations in the Atlantic Ocean, with only four exceptions: one article from the Mediterranean Sea and three articles from the Pacific Ocean. Most articles included data recovered before the year 2013 (Table 2). In six of the included articles, data collection spanned short periods (< 4 years), while six articles had a more extended data collection period (> 4 years).

Only four articles explicitly focused on Hydrobatidae species; the rest focused on seabirds in general. In these last articles, the percentage of Hydrobatidae species representation was below 45% of all recorded seabird groups. Only two articles used a systematic methodology for data collection (Miles et al. 2010, Wilhelm et al. 2021), while four articles used citizen science (Rodríguez and Rodríguez 2009, Murillo et al. 2013, Rodríguez et al. 2015), and three collected their data from records by vessel crew (Black 2005, Glass and Ryan 2013, Ryan et al. 2021). The remaining articles used previous reports from multiple sources (Silva et al. 2020) or from wildlife centers (Montesdeoca et al. 2017, Gjerdrum et al. 2021, Heswall et al. 2022).

Regarding light pollution, eight articles identified sources of light pollution as inshore, offshore sources, or both. Inshore sources of artificial lights were mentioned in four articles (Miles et al. 2010, Murillo et al. 2013, Wilhelm et al. 2021, Heswall et al. 2022), offshore sources were considered in three articles (Black 2005, Glass and Ryan 2013, Ryan et al. 2021), and only one article included both (Gjerdrum et al. 2021). Merely three articles quantified light pollution levels, two used VIIRS satellite data (Rodríguez et al. 2015, Gjerdrum et al. 2021), while Heswall et al. (2022) used previously measured natural sky brightness in the study area. Additional variables on light pollution effects were evaluated in 9 articles; the remaining articles recorded grounding data (location and date). Environmental variables, such as moon phase and wind, were considered in four (Miles et al. 2010, Murillo et al. 2013, Wilhelm et al. 2021, Gjerdrum et al. 2021) and one article (Wilhelm et al. 2021) respectively. Body condition was assessed in five articles (Black 2005, Murillo et al. 2013, Rodríguez et al. 2015, Wilhelm et al. 2021, Gjerdrum et al. 2021), including body mass, wing length, and plumage condition. Storm-petrel´s age class was recorded in six articles (Rodríguez and Rodríguez 2009, Miles et al. 2010, Rodríguez et al. 2015, Silva et al. 2020, Wilhelm et al. 2021, Heswall et al. 2022), while sex was only determined in 2 (Black 2005, Montesdeoca et al. 2017).

Article content

Grounding records

Grounding records were found for all Hydrobatidae genera except for Nesofregetta. Out of the 22 species of the Hydrobatidae family (Tree of Life Web Project 2008, Nunn and Stanley 1998), grounding was recorded in 12 species (Table 3). O. leucorhoa and P. marina were the most frequently mentioned species, followed by H. pelagicus.

Age class

Storm-petrel fallout occurs primarily in fledglings during their first flight leaving the colony. Of the articles that included age class in their analysis, five concluded that fledglings comprised a higher percentage of grounding records in comparison with adult storm-petrels (Miles et al. 2010, Rodríguez et al. 2015, Silva et al. 2020, Wilhelm et al. 2021, Heswall et al. 2022). According to a study (Wilhelm et al. 2021), over 99% of all grounded O. leucorhoa were fledglings. These results coincide with Silva et al. (2020), in which fledglings account for the majority of grounded individuals of O. oceanicus, O. markhami, and G. nereis. Fledgling fallout rates present seasonal variation and coincide with the storm-petrel´s reproductive season, as found by Rodríguez et al. (2015) and Heswall et al. (2022). Contrary to previously described articles, the publication by Rodríguez and Rodríguez (2009) indicated that 84% of grounded O. castro individuals and all of O. leucorhoa, P. marina, and H. pelagicus individuals were adults.

Light pollution and additional variables

Storm-petrels can be attracted to terrestrial or marine light pollution sources. Eight articles registered grounding events inshore, but only four mention the inshore light pollution source (Miles et al. 2010, Murillo et al. 2013, Wilhelm et al. 2021, Heswall et al. 2022). Of these, only Wilhelm et al. (2021) specified the type of property where stranded birds were found (e.g., residential, industry, road, highway, business), and Heswall et al. (2022) made a distinction between urban and rural areas. Three articles (Black 2005, Glass and Ryan 2013, Ryan et al. 2021) assessed bird strike events caused by offshore light pollution sources, mainly fishing vessels' deck lights. Only Gjerdrum et al. (2021) considered offshore and inshore light pollution sources, including oil and gas platforms, vessels, construction sites, and municipalities.

Evidence suggests a potential relationship between moon phases and grounding registers (Miles et al. 2010, Murillo et al. 2013, Gjerdrum et al. 2021, Wilhelm et al. 2021). As established by Wilhelm et al. (2021), during the new moon (lowest moon visibility), O. leucorhoa had the highest grounding rates. On the other hand, there are few stranding records during the full moon. This lack of records is supported by findings made by Gjerdrum et al. (2021), Miles et al. (2010), and Murillo et al. (2013) in which the largest stranding events of O. leucorhoa (Miles et al. 2010, Gjerdrum et al. 2021) and O. oceanicus (Gjerdrum et al. 2021) occurred during the moon’s low illumination phase and most of O. hornbyi fallouts (Murillo et al. 2013) occurred during the first quarter of the moon or new moon. In addition, Miles et al. (2010) considered the percentage of the night that the moon was above the horizon but found no interactions with the moon phase in the analysis of O. leucorhoa grounding rates. Although these articles suggest a potential association between moon phase and groundings, the possibility of a causal link requires further evaluation and is beyond the scope of this study.

Population-level impacts and species’ threat status

According to IUCN (2022), among the 12 storm-petrel species mentioned in the articles included in this study, nine have decreasing population trends, two have unknown statuses, and only one is considered stable. Additionally, O. hornbyi and O. markhami are assessed in the Red List as Near Threatened, O. leucorhoa as Vulnerable (O. leucorhoa), and O. gracilis as Data Deficient. However, the effect of light-induced groundings as a threat to storm-petrel populations is rarely assessed.

Out of the 12 articles included, only five considered the population impact of light pollution (Appendix 3). The articles by Rodríguez and Rodríguez (2009), Ryan et al. (2021), and Rodríguez et al. (2015) found that the percentage of individuals killed annually by fallout is less than 1% of the respective species' population. According to Miles et al. (2010), the number of grounded O. leucorhoa and H. pelagicus is low compared to their breeding population, and Glass and Ryan (2013) stated that it is unlikely that grounding records of P. marina and Fregetta spp. have a significant effect at a population level.

DISCUSSION

This scoping review provides an overview of the state of evidence on the effects of light pollution on storm-petrel (Hydrobatidae) populations worldwide. We discuss the most relevant findings recovered from the analyzed scientific literature, which include (1) the spatial and temporal distribution patterns of light-induced grounding, (2) the implications of these results for future research on light pollution, (3) the knowledge gaps in the reviewed literature, and (4) the limitations encountered during this review.

Relevance of findings for scientific research and literature

This article is the first global review to focus on light pollution effects on the members of the Hydrobatidae family, aside from broader reviews that include other seabird taxa (Rodríguez et al. 2017, Rodríguez et al. 2019, Dias et al. 2019, Silva et al. 2020). Data for storm-petrels on this subject is scarce, with only twelve scientific articles included in the final analysis of this study. Although there is a trend of increasing research on the effects of light pollution on storm-petrels, most articles used data recovered before 2013, which may be outdated information on light pollution and storm-petrel populations. Furthermore, few articles focused specifically on studying the effects of light pollution on storm-petrels. Most articles treated seabirds as a single group, probably due to methodological limitations or difficulty related to the taxonomic identification of closely related storm-petrel species. Given the specific focus of this scoping review on Hydrobatidae, research articles on light pollution effects on other Procellariiformes (i.e., Procellariidae, Diomedeidae, Pelecanoididae) were not included.

Storm-petrel records are limited mainly to the North American and European coastlines (Fig. 2), even though the eastern Pacific Ocean has the highest storm-petrel diversity due to phylogenetic radiation in the late Miocene (Warheit 1992). This pattern is also noticeable in the number of studies assessing light pollution effects on genera and species. Oceanodroma, the most diverse genus within Hydrobatidae, with 13 species out of 22 (Tree of Life Web Project 2008), was evaluated in seven articles. Within this genus, O. tethys and O. markhami, present in South America, were the least studied. The limited research on the effects of light pollution on storm-petrels found primarily in the southern hemisphere, and the distribution pattern record centered in the northern hemisphere may be another case of the identified trends of research limitations in the Global South (Dados and Connell 2012, Collyer 2018). This pattern, however, could also be influenced by the elevated levels of light pollution in regions of the northern hemisphere, such as the coastlines of Europe and North America (Falchi et al. 2016).

Implications for studying light pollution

To determine the extent of light pollution as a threat to the survival of storm-petrel populations, we must first understand which individuals are more susceptible to being disturbed by artificial light sources. One of the variables proven to be fundamental is the age group, as fledglings frequently get disoriented by artificial lights on their first flight toward the sea (Aubrecht et al. 2010, Rodríguez et al. 2017). Six of the evaluated articles considered this variable, and almost all found that storm-petrel fledglings were more frequently grounded by light pollution than adults. These results support evidence from previous studies (Le Corre et al. 2002, Rodríguez and Rodríguez 2009), suggesting that fledgling fallout and mortality may influence population decline (Troy et al. 2011). Even though the reason for fledgling disorientation and attraction toward artificial lights has not been fully established, four main hypotheses have been proposed: (i) inexperienced fledglings may confuse artificial lights with bioluminescent prey (Imber 1975), (ii) storm-petrel fledglings may associate light with a food source since the only light they encounter as nesting chicks is emitted from the burrow entrance used by parents to deliver food (Ainley, personal communication as cited in Rodríguez et al. 2017), (iii) the interplay between an undeveloped visual system and innate behavioral inexperience as fledglings (Atchoi et al. 2020), and (iv) artificial lights may limit their ability to use the moon and stars as navigation cues (Telfer et al. 1987). Understanding the mechanisms of fledgling attraction will allow the proposal of mitigation measures to reduce light pollution effects (Atchoi et al. 2020), particularly during fledgling periods in high-risk areas (Dias et al. 2019).

Among the environmental factors identified in numerous studies to affect storm-petrel susceptibility to light pollution (e.g., wind direction and speed, moon phase, length of time the moon was above the horizon), the moon phase is the most frequently mentioned. An association between grounded storm-petrels’ body condition and subsequent mortality has been established by Wilhelm et al. (2021). However, standardization of data collection may reveal patterns that have remained undetected. Studies performed by Wilhelm et al. (2021), Gjerdrum et al. (2021), Miles et al. (2010), and Murillo et al. (2013) suggest that stranding records focus on nights with the lowest moon visibility, while there are fewer storm-petrel strandings during the full moon (i.e., high moon visibility). In a recent study, Rodríguez et al. (2023) argued that the reduction in strandings during full moon nights may be associated with a lower attraction effect of artificial light when overlapped with natural moonlight. Future studies focusing on storm-petrels’ use of the moon as a navigational cue may reveal the mechanism of disorientation and attraction to artificial lights.

Knowledge gaps in the reviewed literature

One of the limitations of the current state of evidence on the effects of light pollution on the Hydrobatidae family is that only two of the articles from our study used a systematic methodology for data collection. The accuracy of storm-petrel grounding records depends on the search effort, so systematic searches by qualified personnel are needed to increase the proportion of dead birds found (Rodríguez et al. 2017). However, accurate information is difficult to obtain. Effective monitoring programs need to consider a wide array of factors such as the area of impact, the search area and intensity (Rodríguez et al. 2014, Rodríguez et al. 2017), the probability of a carcass to be found (e.g., due to size, carcass accessibility or time of day; Rodríguez et al. 2017, Malinarich et al. 2018 in Silva et al. 2020), light-induced grounding hotspots (Crymble et al. 2020), proximity to the colony, and how quickly a grounded storm-petrel may disappear due to predation or decay (Rodríguez et al. 2017).

Most data about the magnitude of fallout comes from rescue programs carried out by communities, not necessarily as part of systematic monitoring programs (Rodríguez et al. 2017). The success of rescue campaigns, usually conducted by volunteers, relies on public awareness and successful media campaigns (Crymble et al. 2020). These rescue programs allow the generation of conservation knowledge for rare or poorly-known species, which otherwise would be too expensive or intractable to acquire (Rodríguez et al. 2017). Yet, data obtained through rescue campaigns or volunteering programs generally go underreported and should be interpreted as minimum numbers since (1) some grounded birds could be found by people unaware of the light pollution-seabird problem and hence not be reported (Rodríguez et al. 2015), (2) grounded fledglings tend to hide in dense vegetation, holes or crevices, so an unknown number of grounded birds are never found (Reed et al. 1985), (3) the general public is biased toward the collection of live birds and usually do not report or collect dead birds (Rodríguez et al. 2014, Rodríguez et al. 2015, Crymble et al. 2020), and (4) the effort is inherently greater in highly trafficked areas, leading to a possible sampling bias (Burt et al. 2023). Improved data collection and analysis will arise from combining rescue program information with rigorous scientific methods (Rodríguez et al. 2017). These volunteer rescue campaigns, as part of a citizen science approach, provide the opportunity to gather information that, due to resource and time constraints, would not be possible through traditional scientific methods (Kobori et al. 2016), as it may help to monitor stranding events in a broader geographical scale (Burt et al. 2023). Despite the great potential that citizen science has to advance marine and coastal conservation (Cigliano et al. 2015), awareness and community rescue campaigns still need to be encouraged so that the general public understands the threat of light pollution to seabirds, becomes more involved with conservation efforts, and demands solutions from the local and national governments (Rodríguez and Rodríguez 2009, Burt et al. 2023).

There is a considerable gap in the knowledge of light pollution effects on the Hydrobatidae family, which needs to be addressed to propose appropriate management conservation measures. Although some articles identify light pollution sources, the intensity, and spectrum are generally not quantified. This aspect of light pollution is relevant as the spectrum (wavelengths) and intensity of city lights at night are highly diverse, resulting in a mosaic-like distribution that can affect birds' responses in different magnitudes (Dominoni et al. 2013). Other essential factors to consider on light pollution effects are the temporality of light pollution sources (e.g., time of the day, duration, seasonality), the relationship between gender and incidence of light pollution attraction, and the distance between light pollution sources and the storm-petrel breeding populations or their flight paths (Troy et al. 2011, Silva et al. 2016; Rodríguez et al. 2017, Norambuena et al. 2021, Wilhelm et al. 2021). In fact, a recent study on Leach’s storm-petrels found that even a partial reduction in ALAN resulted in a decrease in strandings of up to 57% (Burt et al. 2024). Understanding the potential relationship between the attributes of light pollution sources such as intensity and distance attracting storm-petrels, may allow to quantify the exposure of dark areas and colonies to light pollution (Rodríguez et al. 2017, Norambuena et al. 2021, Wilhelm et al. 2021, Ryan et al. 2021). These and other previously mentioned gaps in our understanding of the effects of light pollution on storm-petrels, particularly at a population level, need to be addressed in future research, as they hold significant implications for conservation management.

Limitations of this study

This scoping review has two main limitations. First, our literature search was conducted exclusively in English, with a prior search in Spanish that yielded no results. However, it is possible that relevant articles written in other languages may exist but were not accessible for inclusion in our analysis. Second, we focused exclusively on scientific articles indexed in Scopus and Web of Science platforms. Information from other web search tools, such as Google Scholar, ProQuest, ScienceDirect, and SciELO was not included. Since we also limited our analyses to scientific research papers published between 2000 and 2022, we acknowledge that important information from earlier research or found in the gray literature, such as government reports and environmental impact assessments, was not considered. Yet, focusing solely on academic scientific articles allowed us to identify relevant information gaps and biases in recent light pollution and storm-petrel research.

CONCLUSIONS AND FUTURE PERSPECTIVES

We identified a series of knowledge gaps that must be further studied to propose effective mitigation measures for light pollution effects on members of the Hydrobatidae family. Future research would benefit from focusing on (1) standardizing data collection of storm-petrel groundings and associated environmental variables, (2) prioritizing systematic collection of grounding data or increasing available information through citizen science programs, (3) assessing the impact of light pollution at a population level to comprehend the effect of light-induced mortality in the species population trends, and (4) characterizing light pollution sources.

As light pollution rapidly increases globally with the expansion of cities, we need to start prioritizing research to determine effects on wildlife and adequately propose management plans and mitigation measures to ensure the well-being of other-than-human beings.

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