Vol.: (0123456789) Aerobiologia (2026) 42:11 https://doi.org/10.1007/s10453-025-09890-w REVIEW PAPER Relationship of airborne fungal spores to epidemiological data on respiratory disease: a systematic review Dámaris A. Jiménez‑Uribe · Rosa Acevedo‑Barrios · Carolina Rubiano‑Labrador · Paloma Cariñanos Received: 28 April 2025 / Accepted: 6 November 2025 © The Author(s) 2026 Abstract  Exposure to fungal spores is associated with various types of respiratory health problems, and volumetric suction particle samplers have been used to estimate their concentrations in the atmos- phere. This systematic review analyzes the sampling of fungal spores in outdoor air worldwide and its relationship to epidemiological data on respiratory disease. Ninety-four studies were identified that met the following inclusion criteria: They were original studies published in English or Spanish between 2010 and 2024, used active volumetric impact samplers, and identified the type of fungal spores in air. Most of the studies were conducted in Europe, with a dura- tion of 1 to 2  years. The fungal taxa with the high- est records were Alternaria sp. and Cladosporium sp. Only 13% of the studies correlated fungal spore con- centrations with epidemiological variables; however, 77% of these studies concluded that there is a clear relationship between airborne fungal spore concentra- tion and the occurrence of respiratory symptoms in the sensitized population. Therefore, this study pro- vides an elaborate review of recent airborne fungal spore surveillance issues worldwide, attempting to include different perspectives of recent research on outdoor volumetric sampling, including epidemio- logical analysis. Keywords  Aerobiology Bioaerosols · Fungal spores · Health effects · Meteorological variables · Respiratory allergies · Outdoor air 1  Introduction Bioaerosols are present in all environments and have the capacity to transport toxic, allergenic, or patho- genic substances that have the potential to affect human health, agricultural productivity, and ecosys- tem stability (Bisen et al., 2012;  Woo et al., 2013). It is estimated that approximately 25% of atmospheric pollution consists of these biological particles (Cao et  al., 2014). A notable subset of bioaerosols com- prises fungal spores, which are characterized by their high reproductive capacity, year-round viability, small size, and ease of dispersion by wind (Castro, 2009; Egan et al., 2014; Pashley et al., 2012). The global diversity of fungal species is estimated to be between 2.2 and 3.8 million, of which over 112 Supplementary Information  The online version contains supplementary material available at https://​doi.​ org/​10.​1007/​s10453-​025-​09890-w. D. A. Jiménez‑Uribe (*) · R. Acevedo‑Barrios · C. Rubiano‑Labrador  Grupo de Estudios Químicos y Biológicos, Dirección de Ciencias Básicas, Universidad Tecnológica de Bolívar, Cartagena de Indias D. T. y C., POB 130001, Colombia e-mail: duribe@utb.edu.co P. Cariñanos  Department of Botany, Faculty of Pharmacy, Andalusian Institute for Earth System Research, IISTA-CEAMA, University of Granada, 18071 Granada, Spain http://crossmark.crossref.org/dialog/?doi=10.1007/s10453-025-09890-w&domain=pdf https://doi.org/10.1007/s10453-025-09890-w https://doi.org/10.1007/s10453-025-09890-w Aerobiologia (2026) 42:11 11   Page 2 of 20 Vol:. (1234567890) have been identified as potentially allergenic due to their capacity to trigger immune responses in sen- sitized individuals (Hawksworth & Lücking, 2017; Levetin et  al., 2016). A series of studies conducted in various regions worldwide have demonstrated that exposure to these fungi can elicit allergic dis- eases, including atopic dermatitis, allergic rhinitis, and asthma. These studies have found that between 19 and 45% of individuals with allergies and up to 80% of asthmatic patients are affected, contingent on the specific country and population group examined. Furthermore, the duration of symptoms has been reported to be significantly prolonged in exposed individuals, (Baxi et al., 2019; Cao et al., 2014; Mari et al., 2003; Pongracic et al., 2010; Roy et al., 2017; Tham et  al., 2017). A substantial body of research has indicated an association between elevated levels of airborne fungal spores and an increase in hospital admissions and asthma-related mortality (Lin et  al., 2016; Newson et al., 2000; Tham et al., 2014). Allergic diseases currently affect approximately 30% of the global population, with a growing preva- lence attributed to elevated concentrations of airborne fungal spores and rising global temperatures  (Lun- dbäck et  al., 2016; Pawankar et  al., 2011; Sánchez et  al., 2022). These diseases represent a significant public health burden, both economically and in terms of reduced labor productivity and quality of life (Cer- vantes-De La Torre et  al., 2018; Charalampopoulos et al., 2022). The onset of allergy-related diseases associated with bioaerosols can be predicted by identifying the fungal genera present in the atmosphere, along with their particle size, concentration, and allergenic potential. These variables can be assessed through aerobiological monitoring (Blais-Lecours et al., 2015; Uk Lee et al., 2016). To date, the majority of moni- toring efforts have been concentrated in temperate and seasonal regions, where pollen-related allergies are commonly diagnosed and reported. However, in tropical regions, characterized by high temperatures, relative humidity, solar radiation, and salinity, all of which influence fungal spore concentration and diversity, aerobiological monitoring remains limited (Huertas et al., 2018; Rodriguez-Gomez et al., 2020). Furthermore, clinical threshold values, defined as the minimum airborne concentrations required to trig- ger allergic symptoms, vary by spore type and have only been established for the most commonly studied genera: Alternaria, Cladosporium, Aspergillus, and Penicillium (O’Gorman & Fuller, 2008; Olsen et al., 2020; Pashley et  al., 2012). This limitation arises from the difficulty clinicians face in correlating spore concentrations with sensitization symptoms, primar- ily due to the lack of comprehensive data on atmos- pheric spore abundance (Anees-Hill et al., 2021; Cec- chi et  al., 2010; Sadyś et  al., 2016). Consequently, continuous bioaerosol monitoring and assessment are imperative to evaluate exposure risks and to establish reliable clinical thresholds. In this context, the present review provides a com- prehensive analysis of recent advances in volumetric sampling of fungal spores in outdoor environments, with particular emphasis on the articulation of these aerobiological data with epidemiological studies using health indicators—such as hospitalizations, skin tests, and respiratory questionnaires—for corre- lation. It has been demonstrated that a comprehensive approach, integrating spore quantification with clini- cal-population analysis, is essential for establishing reliable exposure thresholds and formulating effective preventive strategies against allergic diseases. 2 � Materials and methods This systematic review was based on the Preferred Reporting Items for Systematic Reviews and Meta- Analyses (PRISMA) methodology (Yepes-Nuñez et  al., 2021). A search of the Scopus database was conducted from May to October 2024 using a search string related to outdoor airborne fungal spores (“out- door” AND “airborne” AND “spore” OR “spores” AND “fungal” OR “fungi” EXCLUDES “Indoor Air Pollution”). Studies were initially selected based on title and abstract and subsequently by full-text evalu- ation. The exclusive use of Scopus was justified by its broad multidisciplinary coverage, which includes high-impact journals in environmental sciences, aero- biology, and mycology. Eligibility criteria were: original study published English or Spanish, or between 2010 and 2024, use of active volumetric impact samplers, and identifi- cation of airborne fungal spores. Exclusion criteria were: conference proceedings, reviews or editorials, sampling other than volumetric, identification of fun- gal taxa only by viable methods, sampling conducted indoors or conducted in the vicinity of areas that Aerobiologia (2026) 42:11 Page 3 of 20  11 Vol.: (0123456789) could produce biases in fungal spore counts (waste- water treatment plants, landfills, composting sites, crop fields, forest plantations, etc.). From the systematic review conducted, 2418 stud- ies were initially identified in the database, 193 were selected for their title and abstract, 3 were eliminated because they covered the same sampling period and location, and finally, 94 met the inclusion criteria (Fig. 1 and Online Resource 1). To extract the relevant information from each of the selected articles, a pre-designed template was used that included the following parameters: authors, year of publication, country where the sampling was carried out, continent, start and end dates of the sam- pling, duration of the sampling, methodology used, objective of the study, and fungal taxa identified. In addition, the fungal taxa most frequently identified in the articles were analyzed. To this end, specific data were extracted, such as the seasonal/annual spore integral (spores·day/m3), relative dominance (as a percentage of the total), annual total spores (spores/ m3), peak daily spore concentrations (spores/m3), days with concentrations above the clinical thresh- old (if applicable), and the number of sampling sites. Data were also collected on the statistical correla- tion between spore concentrations and various mete- orological factors, such as temperature, precipitation, relative humidity, wind speed, atmospheric pressure, and radiation (Online Resource 2). When articles presented data corresponding to dif- ferent years or monitoring locations, these were aver- aged to obtain an overall value that would facilitate analysis at the country level. Subsequently, using the data obtained, a meta-analysis was performed to eval- uate the overall behavior of the samples taken over the last 15 years, highlighting the most relevant trends and patterns in relation to meteorological variables and spore concentrations. 3 � Results and discussion 3.1 � Geographical and temporal distribution of studies Fungal spores can be found worldwide; however, in the systematic review conducted, studies were only found in 30 countries (Bisen et  al., 2012). Europe leads the research on fungal bioaerosol, with 53 pub- lished studies, followed by Asia and America. In contrast, Oceania is the least represented in this field. Spain has the highest number of studies (n = 16), fol- lowed by the UK (n = 8), Poland, and Turkey (n = 7) (Fig.  2). Despite the significant representation in Europe, many studies are concentrated in only four countries, limiting the overall understanding of bio- aerosol behavior across the continent. A similar pat- tern is observed in other continents, where only a few countries report fungal spore counts. Within the systematic review, the oldest sampling began in 1978 and, in the 1990s, there was an increase in sampling in Europe and America. In the 2000s, it reached its peak with thirty-seven studies distributed across five continents. Since 1996, non-European countries began to conduct studies; however, Africa and Oceania have not continued to publish results to date, demonstrating a large information gap on this topic in these continents (Fig. 3). Most fungal sampling lasted 1  year (n = 35) or 2  years (n = 20), and only a subset reported follow- ups of ≥ 10  years (n = 9). Although an annual cycle allows for the description of intra-annual seasonality at a site, it is insufficient to establish stable and gen- eralizable seasonality (Sánchez Espinosa et al., 2024). Furthermore, long-term analyses show that climatic conditions can modify the duration of seasons and generate upward trends in concentrations (Milling- ton & Corden, 2005; Reznik et  al., 2023). Conse- quently, multi-year series are required to understand local climatology, separate atypical years from real trends, and infer robust long-term behaviors. In this context, studies on fungal bioaerosols have pursued Fig. 1   Flowchart of the systematic review selection process Aerobiologia (2026) 42:11 11   Page 4 of 20 Vol:. (1234567890) various objectives: 78% identified and quantified spores, 42% compared counts with meteorological variables, 33% analyzed spatiotemporal variability, 13% compared sampling/analysis methodologies, and 6% implemented predictive models. This heterogene- ity of approaches reinforces the need for multi-year series and comparable protocols that integrate these objectives and allow for more robust and transferable conclusions. 3.2 � Taxa of airborne fungal spores In the systematic review, allergenic spores were mainly associated with the Ascomycota (82%) and Fig. 2   Worldwide sampling locations of the studies included in the review. The color of the countries indicates the number of stud- ies published. The gray rectangles represent the number of published studies per continent Fig. 3   Temporal and geographical distribution of the studies included in the review, according to the year in which their sam- pling campaigns began Aerobiologia (2026) 42:11 Page 5 of 20  11 Vol.: (0123456789) Basidiomycota (18%) phyla. The most identified spores within each phylum are presented in Fig. 4. 3.2.1 � Phylum Ascomycota The phylum Ascomycota, reported on five continents and in 82% of the articles reviewed, reproduces sexu- ally through ascospores formed in asci or asexually through conidia produced in conidiophores. Not all species generate both types of spores; their occur- rence varies between taxa (Deacon, 2005; Liu, 2024). While ascospores are usually released after wet or rainy periods, conidia are dispersed more continu- ously and include numerous clinically important allergenic taxa (Levetin et al., 2016). Among the allergenic taxa, Alternaria spp. is one of the most frequently mentioned in studies on its allergenic potential, because most exposures occur in outdoor environments  (Kasprzyk & Worek, 2006; Williams et  al., 2016). Although Alternaria concen- trations in the environment are lower than those of other allergenic spores, a significant sensitization rate of 13–17% has been observed among atopic patients, representing up to 60% of positive cases (Amado et al., 2014; Arbes et al., 2005; Lehmann et al., 2017). Patients sensitized to this genus also have a greater propensity to develop allergic reactions to other taxa (Amado et  al., 2014). Several scientific studies have established a correlation between asthma and aller- gic rhinitis with skin tests for Alternaria spp., which show sensitization generally at an early age and result in the development of childhood asthma (Moral et al., 2008; Salo et al., 2006). The studies examined show that airborne con- centrations of Alternaria exhibit marked spatial and temporal variability. Globally, the relative contribu- tion of this genus is predominantly in the range of 2% to 15% of total fungal spores, with outliers reported in Qatar (19%) and in certain contexts in Spain and Pakistan (> 10%)(Gharbi et al., 2022; Hasnain et al., 2012; Vélez-Pereira et  al., 2016). Annual concentra- tions vary considerably, ranging from a few hundred spores/m3 in regions with low occurrence, such as Cuba or certain areas of India, to more than 100,000 Fig. 4   Fungal genera reported by continent and phylum. Genus labels in dark blue = Ascomycota; genus labels in light blue = Basidi- omycota. The vertical bar at right summarizes the share of studies by phylum (Ascomycota = dark blue; Basidiomycota = light blue) Aerobiologia (2026) 42:11 11   Page 6 of 20 Vol:. (1234567890) spores/m3 in Jordan and Kuwait, indicating significant regional contrasts (Abu-Dieyeh & Barham, 2014; Al- Ahmad et  al., 2019; Almaguer-Chávez et  al., 2018; Chakrabarti et al., 2012). In terms of seasonal dynam- ics, a consistent pattern is evident in Europe, Western Asia, and the Americas, characterized by peaks dur- ing the warm, dry months, particularly between July and September, with an onset in spring (March–May) and a decline toward autumn (September–November). However, there are notable exceptions: In India, con- centrations are associated with the monsoon period, while in Mexico, peaks are concentrated during the summer (Dey et al., 2019; Ortega Rosas et al., 2020). Overall, the available evidence confirms that Alter- naria is a ubiquitous and clinically relevant genus, with a defined seasonality in temperate and subtropi- cal regions, although its abundance is highly depend- ent on local climatic and geographical factors. Cladosporium was identified in 67% of the stud- ies and is also the most frequent and the most abun- dant genus in propagule samples. This is because its spores are easily dispersed by wind (Oliveira et  al., 2009; Pyrri & Kapsanaki-Gotsi, 2015; Soleimani et  al., 2013). Its conidia are pigmented and meas- ure 5–40 × 3–13  µm, forming simple or branched chains. This genus has a strong allergenic potential that severely affects people with allergic rhinitis and asthma (Gabriel et  al., 2016; Ogórek et  al., 2012; Pomés et al., 2016; Raphoz et al., 2010). The data collected for Cladosporium show that this genus is one of the most dominant components of the atmospheric microbiota, reaching relative abundance percentages that often exceed 50% and, in some cases, reaching values above 80% of the total spores recorded (Pace et  al., 2019). The annual concentra- tions reported are remarkably high, with figures rang- ing from tens of thousands to several million spores/ m3, as observed in studies conducted in Poland, Slo- vakia, and the UK (Ščevková et al., 2023; Weryszko- Chmielewska et  al., 2018; Žilka et  al., 2024). Like- wise, daily peaks range widely, from hundreds to more than 10,000 spores/m3, evidence of high spatial and temporal variability (Ianovici, 2016; Rad et  al., 2023). In terms of seasonality, records in Europe, Asia, and America agree that peaks occur during the warm months, mainly between July and August, with a general onset in spring (March–May) and a decline in autumn (September–November) (Grinn-Gofroń et al., 2020; Kilic et al., 2020; Patel et al., 2018). In tropical regions, such as the Caribbean, a marked abundance is also reported in summer, closely related to local weather conditions (Almaguer-Chávez et al., 2018; Díaz Vázquez et  al., 2024). In summary, the findings indicate that Cladosporium has a wide geo- graphical distribution, marked summer seasonal- ity, and a predominant share of the total spore load, underscoring its epidemiological importance and its status as a leading fungal allergen on a global scale. Aspergillus spp. and Penicillium spp. are two of  the most prevalent airborne allergens, mainly associ- ated with asthma exacerbations and proliferation in high-humidity environments. A review of the exist- ing literature reveals that they are mentioned in 31% of the studies reviewed (Scott et  al., 2004; Tournas, 2005). Aspergillus and Penicillium spores exhibit a globose or subglobose morphology, with smooth or rough walls and a diameter ranging from 2 to7  µm. This characteristic makes it difficult to detect and dif- ferentiate spores using volumetric traps and optical microscopy, complicating the taxonomic identifica- tion process. Consequently, these spores are collec- tively classified as “Aspergillus/Penicillium type” (de Ana et  al., 2006; De Linares et  al., 2023; Piontelli, 2008) These genera have also been linked to persis- tent asthma and have been characterized as multiple specific allergens (Cramer et al., 2011; Deacon, 2005; Kousha et al., 2011; Williams et al., 2016). Aspergillus/Penicillium records indicate wide- spread presence, although in relative terms, generally minor compared to other genera: Typical dominance ranges from approximately 1–8%, with occasional increases to around 20–21%, for example in Taiwan and Cuba (Espinosa et  al., 2019; Kallawicha et  al., 2017). The reported magnitudes show marked het- erogeneity, with daily peaks ranging from tens to more than 4000 spores/m3, while annual totals fluc- tuate between hundreds and approximately 40,000 spores/m3 at various points in Spain (Vélez-Pereira et  al., 2016). The phenology is characterized by its perenniality, manifesting itself in continuous detec- tion throughout the year in various locations (Kalla- wicha et al., 2017; Naseer et al., 2024; Roy & Gupta Bhattacharya, 2020). Seasonal peaks are observed, which in the northern hemisphere are concentrated between June and October, declining toward the end of autumn; in the southern hemisphere, peaks are recorded in the southern spring (October–Novem- ber), with seasons that can extend into autumn (Nitiu Aerobiologia (2026) 42:11 Page 7 of 20  11 Vol.: (0123456789) & Mallo, 2011). Overall, the available evidence indi- cates that Aspergillus/Penicillium is ubiquitous and clinically relevant, although its contribution to the total spore load tends to be moderate and its variation is highly dependent on location and season. Based on the data provided, no additional causal mechanisms or patterns other than seasonality are observed. For Drechslera spp.,  most species of the genus are saprobes and plant pathogens. Their conidia are pig- mented, pseudoseptate and can germinate from sin- gle cells (Goh et al., 1998). Some species can cause opportunistic infections, fungal sinusitis and bron- chopulmonary mycosis (Menezes et  al., 1998). In addition, some species have been reported to be aller- gen producers and were reported in 19% of the stud- ies (Chew et al., 2000). The records available for Drechslera are scarce and heterogeneous, but they converge on a low rela- tive contribution to the total spore load of between 0.2 and 4.1%, as well as modest annual magnitudes of between 100 and 10,000 spores/m3. In Europe (Spain, Serbia, Portugal) and Asia (India), seasonality is con- centrated in the warm months of the northern hemi- sphere, with the season beginning around May–June, peaking in July–August, and ending in August–Sep- tember (Das & Gupta-Bhattacharya, 2012; Simović et  al., 2023; Sousa et  al., 2016; Vélez-Pereira et  al., 2016). Cuba shows peaks in December (Espinosa et al., 2019). Daily peaks vary widely between 4 and 175 spores/m3, suggesting marked spatial variability. Given the limited number of series and the absence of complete information in several studies, it is not pos- sible to draw additional conclusions beyond this sea- sonality and its generally minor nature. The  genus Epicoccum spp.  is frequently found in soil as a saprophyte and plant pathogen (Grinn‐ Gofroń, 2008). Epicoccum spp. were documented in 20% of the studies. Its conidia are characterized by brown color, rough walls, globular shape, and multi- cellular structure, measuring between 15 and 20 µm (Levetin et  al., 2016). Epicoccum presents a spore distribution pattern similar to that of Alternaria. In particular, it belongs to allergenic genera associated with conditions such as hypersensitivity pneumonitis and allergic fungal sinusitis (Bisht et al., 2002; Rizzi- Longo et al., 2009). Epicoccum spp. showed moderate and seasonal presence, with daily peaks that in most studies did not exceed 500 spores/m3 and annual totals generally below 3,500 spores·day/m3. Its relative dominance rarely exceeded 2%, placing it as a secondary genus in terms of atmospheric abundance. In Europe, it reached its most representative values, with summer peaks of up to 870 spores/m3 and annual totals of 200–3,000 spores·day/m3, showing well-defined sea- sons between June and September (King et al., 2011; Simović et  al., 2023). In America, the records were lower, with maximums of 100 spores/m3, present in the dry season (Espinosa et al., 2019). In Asia, values were intermediate, with peaks of around 400 spores/ m3 (Akgül et al., 2016). Epicoccum spp. behaves as a secondary genus in abundance, but with clear season- ality, whose contribution to the atmospheric mycobi- ota is particularly relevant in temperate and Mediter- ranean climates. Regarding Curvularia spp., although  rarely abun- dant,  the  conidia are frequently found in air sam- ples, characterized by thick walls, transverse septa and a curved appearance (Levetin et al., 2016). They were identified in 15% of the studies and have been associated with susceptibility to asthma and res- piratory allergies (Sio et  al., 2021). Daily peaks rarely exceeded 250 spores/m3, while annual totals were around 1,000 spores·day/m3. Its relative domi- nance was less than 2% in most of the contexts ana- lyzed. The seasonality of Curvularia was mainly related to periods of heat and humidity, reflecting its dependence on favorable conditions for sporula- tion (Almaguer-Chávez et al., 2018). In Asia, especially in Pakistan, Curvularia had the highest values, with annual totals of more than 2,000 spores·day/m3(Abbas et al., 2012). In America, maximums of 200 spores/m3 were recorded, with a relative dominance of 6%, higher than on other con- tinents (Espinosa et  al., 2019). In Europe, concen- trations were low and sporadic, with average totals of 62 spores/m3, remaining a genus of secondary presence (Fernández-Rodríguez et  al., 2014; Sousa et al., 2016). Curvularia spp. behaves as a genus that is not very abundant, but with a marked association with warm climates, being more relevant in tropical regions, where it reaches its maximum concentration values. 3.2.2 � Phylum basidiomycota Phylum Basidiomycota exhibits a vast macromor- phological producing its sexual spores, basidiospores Aerobiologia (2026) 42:11 11   Page 8 of 20 Vol:. (1234567890) externally on basidia (Reznik et  al., 2023; Rivera- Mariani et al., 2020). In non-viable light-microscopy counts from Hirst-type samplers, these basidiospores are typically small and morphologically undistinc- tive (≈3–12  µm), which precludes reliable genus- level resolution; accordingly, they are reported in aggregate as “basidiospores.” Only a few genera with pronounced diagnostic traits—such as Ganoderma or Coprinus—can occasionally be recognized under these conditions (Elbert et al., 2007; Hernández Trejo et al., 2012; Kasprzyk, 2008). Despite being frequently recorded as a pooled cat- egory, basidiospores are clinically relevant: They are known to produce allergens, and higher concentra- tions have been associated with increased emergency visits for asthma (Chane-Si-Ken et al., 2022). In this review, 18% of the studies analyzed reported Basidi- omycota, with records spanning fifteen countries across Europe, the Americas, and Asia, underscoring their broad distribution and relevance for aerobiologi- cal surveillance. Ganoderma spp., identified in 24% of the studies, is commonly known as a wood decay fungus and a prevalent genus of airborne fungal spores worldwide (Craig & Levetin, 2000; Singh & Mathur, 2021). Its spores are recognizable by their orange inner wall and spines penetrating a hyaline outer wall, ranging in size from 6.5 to 13 × 5 to 9  μm. Recent studies have identified that this genus may be the third most prevalent allergenic genus after Alternaria and Clad- osporium (Grinn-Gofroń & Strzelczak, 2011). Ganoderma spores show a well-defined sea- sonal pattern, with daily peaks between 156 and 720 spores/m3 and annual totals generally below 30,000 spores/m3, reaching their maximum in the summer and early fall months, especially in July and August. In Europe, the genus shows a regular but not domi- nant presence, with averages of 600 spores/m3 and annual totals of around 14,000 spores/m3 (Fernán- dez-Rodríguez et  al., 2018; Ščevková et  al., 2020). In America, regional differences are observed: In the Caribbean, levels reach up to 2600 spores/m3, with prolonged seasons lasting until autumn, while in South America the values are low (Almaguer- Chávez et al., 2018; Cid-Martínez et al., 2019; Díaz Vázquez et al., 2024; Emygdio et al., 2018). In Asia, in continental and semi-arid climates, Ganoderma has shorter seasons, with peaks of 300–400 spores/m3 (Abu-Dieyeh & Barham, 2014; Akgül et al., 2016). In general, Ganoderma maintains moderate abundances, but with significant peaks in tropical and Mediterra- nean areas, establishing itself as a summer genus. The genus Coprinus releases abundant basidi- ospores when humidity levels are high (Almaguer et  al., 2014; Webster & Weber, 2007). Studies have found that due to their small size, Coprinus spores can penetrate deeply into the respiratory tract, wors- ening the respiratory conditions of patients (Lev- etin et  al., 2016; Rivera-Mariani & Bolaños-Rosero, 2012). It was reported in 15% of the articles reviewed, as being associated with respiratory allergies (Cail- laud et al., 2022). Its distribution is seasonal, with peaks between spring and fall, and its concentrations vary widely, from 82 spores/m3 to peaks of 2,549 spores/m3, with a seasonal average ranging between 700 and 26,000 spores/m3. In the Americas, Coprinus showed high dominance in Cuba (15.8%), with seasons between May and October, while in Mexico and Brazil the values were much lower, with dominances below 3% (Díaz Vázquez et al., 2024; Emygdio et al., 2018; Rocha Estrada et  al., 2013). In Europe, the highest annual totals were reported in Spain and Serbia, with well-defined summer peaks, while in Asia the values were considerably lower, with annual concentrations of only 340 spores/m3 (Das & Gupta-Bhattacha- rya, 2012; Simović et al., 2023; Vélez-Pereira et al., 2016). In summary, Coprinus is not a marginal genus, as in some regions it can reach high concentration and dominance, while in others its presence is almost nil, highlighting its dependence on local factors. The genus Agrocybe was reported in 10% of the studies in the review, as an allergenic spore (Ščevková et al., 2019, 2020; Sevindik et al., 2022; Vélez-Pereira et al., 2016). It has been classified as a wet spore, like Coprinus, Ganoderma, and Leptosphaeria. These spores show higher concentrations during the night or early morning hours, and a negative correlation with air temperature has been observed. (Carlile et  al., 2001; Ščevková et  al., 2019). On the other hand, an increase in Agrocybe concentration in the air has also been reported with an increase in atmospheric NO2 levels (Damialis et al., 2015). The data available for Agrocybe shows limited dis- tribution and low dominance in atmospheric spore concentrations. In studies conducted in Turkey and Ukraine, median daily peaks range from 124 to 1,017 spores/m3, with a relatively low annual total of 1,021 Aerobiologia (2026) 42:11 Page 9 of 20  11 Vol.: (0123456789) spores/m3 in Ukraine and 1,017 spores/m3 in Turkey. The dominance of this genus is also low, with values reaching only 1.2% in Turkey and 0.2% in Ukraine, reinforcing its minor role in the total atmospheric spore load. In terms of seasonality, Agrocybe shows a defined pattern, with higher concentrations between April and August, with July and September being the peak months in the different locations(Reznik et  al., 2023; Sevindik et al., 2022). Overall, these data sug- gest that Agrocybe is a genus with moderate and sea- sonal presence in certain regions, but with a minimal contribution to the total spore load in the air. 3.3 � Sampling and quantification methodologies The seven-day volumetric spore trap based on the initial design of Hirst (1952) employs the non-viable particle impaction method, which is widely used by various organizations for pollen and spore monitor- ing. The volumetric spore sampler is usually placed on the roofs of buildings to collect a representa- tive sample of the air that has been mixed, bringing with it particles released from diverse sources (Lacey & West, 2006). In the studies analyzed, the average installation height of the volumetric sampler was 17 m above ground level. Viable and non-viable bioaerosols are identified and quantified by optical microscopy, which also allows detailed observation of their structures. Their efficiency is improved by using stains such as meth- ylene blue, which allow them to be classified mor- phologically. This method, called direct counting, is nowadays one of the most widespread analytical techniques. To obtain better visibility of the parti- cles, phase contrast microscopes are used to observe nearly invisible particles, fluorescence microscopy, which uses ultraviolet or near-ultraviolet illumina- tion that causes fluorescent particulate compounds to emit light, or scanning electron microscopy (SEM), in which the surface of the bioaerosol is scanned with an electron beam (Després et al., 2012; Pöhlker et al., 2012). This systematic review found that 17% of the stud- ies used light microscopy for taxon identification and counting, while 6% used culture techniques, fluores- cence microscopy or DNA analysis to compare results between analytical methods. Despite uniformity in sampling methodologies and the units in which con- centrations are expressed (spores/m3) (Galán et  al., 2007), studies vary in the way they are reported. Some provide daily averages, others monthly aver- ages, others the annual integral, and some present data in total spore counts while others provide them by fungal taxon. This makes quantitative comparisons between the studies analyzed very difficult (Janssen et al., 2021; Symon et al., 2025). 3.4 � Relationship with meteorological factors Meteorological parameters such as temperature, rela- tive humidity, solar radiation, and atmospheric pres- sure influence the production, release, dispersion, deposition, diversity, and concentration of bioaero- sols (Bertolini et  al., 2013; Polymenakou & Manda- lakis, 2013; Şakiyan & Inceoǧlu, 2003). In 49% of the reviewed studies, the relationship between fungal spore concentrations and meteorological variables was analyzed by statistical correlation coefficients. 3.4.1 � Temperature Temperature is one of the most influential factors that, depending on the season and time of day, affects the metabolism, reproduction, and viability of fungal spores, correlating directly with atmospheric concen- trations (Mantoani et al., 2025; Millington & Corden, 2005; Pasanen et al., 2000). In the case of dry spores, Alternaria and Cladosporium showed consistent pos- itive correlations with minimum, average, and maxi- mum temperatures in Mediterranean regions such as Morocco and Spain, reaching peaks of more than 600 spores/m3 on the warmest days and increasing the number of days above the clinical threshold dur- ing the summer (Bardei et al., 2017; Sabariego et al., 2012; Sánchez Espinosa et al., 2024). Less abundant genera such as Drechslera and Curvularia also fol- lowed this pattern, with summer increases in coun- tries such as India and Mexico, reflecting their close link to the growth and aging cycles of grasses in hot conditions (Das & Gupta-Bhattacharya, 2012; Rocha Estrada et  al., 2013). Similarly, Aspergillus/Penicil- lium was more prevalent in warm climates, with posi- tive associations with average temperature and daily maximums above 400 spores/m3 in Asian studies (Kallawicha et al., 2017; Naseer et al., 2024). Other less abundant genera, such as Drechslera and Curvularia, recorded their highest concentrations in the summer months in Cuba and Spain, coinciding Aerobiologia (2026) 42:11 11   Page 10 of 20 Vol:. (1234567890) with rising temperatures, which reinforces their ther- mophilic nature (Almaguer-Chávez et  al., 2018; Fernández-Rodríguez et al., 2014). Coprinus showed variable behavior: In Cuba and Ukraine, it reached high annual totals with peaks during the warm months of July and August, while in Mexico and Brazil, its presence was marginal, probably limited by local conditions (Almaguer-Chávez et  al., 2018; Emygdio et  al., 2018; Reznik et  al., 2023; Rocha Estrada et  al., 2013). Overall, the evidence confirms that temperature acts as a facilitating factor for dry spores (Alternaria, Cladosporium, Aspergillus/Peni- cillium, Drechslera, Curvularia), while in genera such as Coprinus its effect depends more on the inter- action with substrate availability and local conditions. 3.4.2 � Relative humidity Relative humidity showed different effects depend- ing on the type of spore. In the group of dry spores, such as Alternaria and Cladosporium, negative cor- relations with humidity were observed in several studies in Europe and North Africa (Filali Ben Sidel et  al., 2015; Sarda-Estève et  al., 2019). In Granada and Seville, Cladosporium concentrations decreased significantly on days with humidity above 70%, while Alternaria reduced its presence in saturated envi- ronments, maintaining its maximum levels under dry summer conditions (Aira et al., 2013). This pat- tern also extended to less abundant genera such as Drechslera and Curvularia, whose summer peaks were associated with dry periods following grass senescence. Aspergillus/Penicillium showed simi- lar behavior: Their concentrations decreased in very humid environments, confirming their preference for relatively dry and hot periods (Espinosa et al., 2019). In contrast, moist spores responded in the opposite manner. In Spain and Greece, Ganoderma recorded marked increases on days with humidity above 75%, reflecting its dependence on moisture for the matu- ration of fruiting bodies and active spore release (Charalampopoulos et  al., 2022; Elvira-Rendueles et  al., 2013). Coprinus showed a comparable pat- tern: In Cuba and Serbia, its highest loads coincided with humid periods in summer and fall, while in drier regions such as Mexico, its presence was marginal (Almaguer-Chávez et al., 2018; Rocha Estrada et al., 2013; Simović et al., 2023). In summary, relative humidity is a critical deter- minant of the composition of atmospheric mycobiota: Dry spores (Alternaria, Cladosporium, Aspergil- lus/Penicillium, Drechslera, Curvularia) proliferate under dry conditions and decline under high humid- ity, while moist spores (Ganoderma, Coprinus) reach their maximum levels in moisture-saturated environ- ments, defining opposite patterns that explain much of the regional variability observed. 3.4.3 � Other meteorological factors Among secondary meteorological variables, wind played a significant role in the dispersion and trans- port of spores. Studies conducted in Europe observed that moderate wind speeds favored the upward and horizontal movement of light genera such as Clad- osporium and Aspergillus/Penicillium, while intense gusts reduced local concentrations through dilution (Antón et al., 2019). Atmospheric pressure, although less studied, showed significant associations in cer- tain regions: Pressure drops preceded increases in Leptosphaeria concentrations, possibly due to mete- orological instability favoring spore release and sus- pension (Sadyś et al., 2015). Solar radiation has a dual effect: High levels tend to increase concentrations of Cladosporium and Gan- oderma, while in genera such as Leptosphaeria they decrease on very sunny days (Grinn-Gofroń et  al., 2020; Ščevková et  al., 2019). Beyond climate, cer- tain spore traits favor their persistence in the air: A resistant cell wall gives them tolerance to dryness, and dark pigments such as melanin protect them from ultraviolet radiation and oxidative damage (Hopke et al., 2018; Suthar et al., 2023). Together, these fac- tors help explain why pigmented dry spores tend to predominate in warm, dry, and windy environments, in addition to the effect of weather conditions. In summary, meteorological factors such as tem- perature, humidity, wind, pressure, and radiation interact synergistically, modulating the daily and sea- sonal dynamics of the atmospheric mycobiota, favor- ing dominant genera and limiting those most sensitive to extreme conditions (Zhai et al., 2018). 3.5 � Clinical thresholds In order to diagnose allergic diseases, it is essen- tial to identify the concentration of allergens in the Aerobiologia (2026) 42:11 Page 11 of 20  11 Vol.: (0123456789) atmosphere and to establish the relationship between exposure to these concentrations and the onset of symptoms. This enables the definition of appropriate clinical threshold values (Ghosh et al., 2022). Various governmental agencies and private organizations have proposed exposure limits for fungal spores; however, many of these thresholds are expressed in colony- forming units (CFU), which are based on viable sam- pling techniques that present significant limitations. This is primarily because, although some bioaerosols are viable, they are not always cultivable and may fail to form colonies under standard laboratory con- ditions, resulting in underestimated concentrations ( Becher et al., 2000; Chi & Li, 2007; de Aquino Neto & de Góes Siqueira., 2000; Jo & Seo, 2005; Kim et  al., 2018; Pegas et  al., 2010; Wainwright et  al., 2004; Yang & Heinsohn, 2007). Other studies have established threshold values based on non-viable sampling methods; however, these efforts are generally limited to the most exten- sively studied fungal genera and show significant var- iability across countries. For instance, the proposed threshold for Alternaria is 50 spores/m3 in the United Kingdom, 80 spores/m3 in Poland, and 100 spores/ m3 in Finland (Frankland & Davies, 1965; Ranta & Pessi, 2006; Rapiejko et  al., 2007). For Clad- osporium, the clinical threshold is 3,000 spores/m3 in the United Kingdom and Croatia, 2800 in Poland, and 4000 in Finland (Frankland & Davies, 1965; Peter- nel et al., 2004; Ranta & Pessi, 2006; Rapiejko et al., 2007). Conversely, studies conducted in Germany on Aspergillus spp. suggest that allergic symptoms may emerge at concentrations as low as 50 spores/m3 (Holmberg, 1987). Given the heterogeneity of clinical thresholds reported across countries, one of the most useful epi- demiological indicators identified in the reviewed lit- erature is the number of days per season that exceed the clinical threshold. This parameter not only reflects the duration of exposure risk for sensitized indi- viduals but also provides a more direct link between atmospheric concentrations and potential health outcomes (Ortega-Rosas et  al., 2025; Vélez-Pereira et al., 2023). Based on the articles analyzed in this review, consistent data on days above threshold were found mainly for Alternaria and Cladosporium. For Alter- naria (threshold ≥ 100 spores/m3; 25 site-seasons ana- lyzed), the median number of days above threshold was 37 (IQR 28–58), with a range of 4–125 and a mean of 47. Notably, 80% of sites exceeded 15 days, 72% exceeded 30 days, while 20 and 16% surpassed 60 and 90 days, respectively. At the continental level, the median exposure was highest in Africa (89 days), followed by Europe (42.5  days) and Asia (28  days), indicating that populations in semi-arid and Mediter- ranean climates face the longest risk windows. For Cladosporium (threshold ≥ 3000 spores/ m3; 17 site-seasons), the median was 65  days (IQR 40–90), with a range of 12–147 and a mean of 64.4. Here, 88.2% of sites had ≥ 15 days, 76.5% ≥ 30 days, and 64.7% ≥ 60  days, while nearly 30% experi- enced ≥ 90 days above threshold. The highest values were reported in Asia (median 95 days) and Europe (median 64  days), with more moderate exposure in Africa (22  days). For Aspergillus/Penicillium, data availability was limited (n = 1, 2  days above thresh- old), underscoring the need for further epidemiologi- cal studies before meaningful comparisons can be made. Taken together, these findings highlight that Alternaria and Cladosporium can sustain extended seasonal exposures above clinical thresholds, often for several consecutive weeks, with significant regional variability. 3.6 � Relationship with epidemiological studies Numerous researchers have postulated that the health implications associated with airborne fungal spores are contingent on three primary factors: (a) the fun- gal genera present in the atmosphere, (b) their aller- genic potential, and (c) their environmental concen- tration (Ghosh et  al., 2022; Roponen et  al., 2002). While these three factors are critical for estimating health risks, most studies have addressed only one or, at most, a combination of two, without integrating all three factors simultaneously. This tendency is exemplified in the present sys- tematic review, which revealed that a mere 13% of the analyzed studies correlated fungal genera and airborne spore concentrations with epidemiologi- cal data. The evaluated populations in these stud- ies were primarily composed of patients treated at healthcare facilities for respiratory conditions, including coughing, recurrent colds, dyspnea, sneezing, wheezing, asthma, and allergic rhinitis. A notable aspect of the analysis is the absence of age Aerobiologia (2026) 42:11 11   Page 12 of 20 Vol:. (1234567890) stratification in 8 studies, with 5 of them focusing exclusively on pediatric and adolescent populations. To assess the relationship between respiratory diseases and airborne spore concentrations, six studies applied skin prick tests using fungal aller- gen extracts and compared positive results with bio- aerosol counts(Abbas et al., 2012; Al-Ahmad et al., 2019; Chakraborty et  al., 2014; Dey et  al., 2019; Katotomichelakis et  al., 2016; Kilic et  al., 2020). Three studies employed statistical correlation meth- ods to evaluate the association between the number of respiratory disease cases and allergenic spore concentrations (Batra et  al., 2022; Chakrabarti et  al., 2012; Ortega Rosas et  al., 2020). Two addi- tional studies employed respiratory health question- naires to assess clinical conditions and subsequently compared those results with aerobiological sam- pling data (Al-Nesf et al., 2022; Chen et al., 2011). However, only one study directly compared daily spore concentrations with clinical threshold values reported in the literature (Weryszko-Chmielewska et al., 2018). The collective findings of these studies indicate a clear relationship between the concentra- tion of airborne fungal spores and the onset of res- piratory symptoms in sensitized individuals. Despite the pervasive recognition of the detri- mental health implications associated with fun- gal spores, a comprehensive, universally accepted dose–response relationship that delineates accept- able exposure limits remains elusive. This dearth of knowledge stems from three interrelated fac- tors. Firstly, there is an absence of consistent dose–response data. Secondly, the availability of real-time bioaerosol identification and quantifica- tion is insufficient. Thirdly, methodological vari- ability among existing studies hinders cross-com- parability. Consequently, there is an imperative for the development of robust longitudinal research conducted across multiple environments and geo- graphic regions, employing standardized and vali- dated sampling methodologies. Such studies should not only identify and quantify airborne fungal taxa but also correlate these findings with clinical indi- cators of respiratory health. It is only through this comprehensive approach that we can generate appli- cable scientific evidence that will directly benefit sensitized populations and contribute to improving their quality of life. 4 � Conclusion Despite the ubiquity of fungal bioaerosols, the extant evidence is concentrated in a limited number of con- texts. Specifically, outdoor volumetric sampling was carried out in 30 countries, mostly with 1–2  year campaigns. The geographical and temporal coverage of the data is restricted, which limits global infer- ence and the capture of interannual variability. This results in notable gaps in regions such as Africa and Oceania, as well as in tropical and semi-arid climates. In order to enhance existing knowledge and mitigate regional biases, there is a necessity for the establish- ment of coordinated observation networks and the implementation of extended time series. These meas- ures are instrumental in characterizing regional pat- terns and trends with greater resolution. The clinical link is the primary factor hindering the attainment of accurate results in the studies con- ducted. A mere 13% of the research has established a correlation between concentrations and genders with epidemiological indicators, such as skin tests, medi- cal consultations, or hospitalizations, as well as ques- tionnaires. The issue of scarcity, when considered in conjunction with the heterogeneity of clinical thresh- olds, poses a significant challenge in determining applicable dose–response relationships and compar- ing exposure thresholds between regions. The results of the study identify prolonged expo- sure intervals for genera of high health relevance. Alternaria (≥ 100 spores/m3) demonstrated a median of 37  days across 25 site-seasons, while Clad- osporium (≥ 3000 spores/m3) exhibited a median of 65  days in 17 site-seasons, exhibiting significant regional variability. This evidence thus provides a scientific basis for the implementation of specific seasonal alerts and the prioritization of both genera in surveillance. Conversely, the observed taxonomic landscape (Ascomycota 82% and Basidiomycota 18%) provides a framework for the standardization of allergen panels and the allocation of monitoring resources. In summary, the current evidence base facilitates progress in surveillance; however, further research with an epidemiological focus is necessary to trans- late this into clinical and public health decisions. This should include longitudinal and multicenter studies that integrate clinical and environmental variables Aerobiologia (2026) 42:11 Page 13 of 20  11 Vol.: (0123456789) from their design, the adoption of comparable met- rics explicitly linked to health outcomes, geographi- cal expansion to underrepresented regions, and more extensive campaigns that capture interannual vari- ability. It is only in this manner that aerobiological monitoring will be capable of providing support for the formulation of preventive recommendations, the generation of seasonal alerts, and the establishment of a prioritization of allergens based on the actual risk to the population. Author contributions  All authors contributed to the concep- tion and design of the study. The literature search, data analy- sis, and writing were carried out by Dámaris A. Jiménez-Uribe. Rosa Acevedo-Barrios, Carolina Rubiano-Labrador, and Pal- oma  Cariñanos critically reviewed the paper and commented on earlier versions of the manuscript. All authors read and approved the final manuscript. Funding  Open Access funding provided by Colombia Con- sortium. This work was supported by Department of Science, Technology and Innovation-COLCIENCIAS (Colombia) through the doctoral fellowship to D.A. J.-U. Data Availability  No datasets were generated or analyzed during the current study. Declarations  Conflicts of interest  The authors declare no competing inter- ests. Open Access  This article is licensed under a Creative Com- mons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Crea- tive Commons licence, and indicate if changes were made. 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