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In tropical regions, pollination is primarily carried out by animals, hawkmoths (Lepidoptera, Sphingidae), being perceived as one of the most important groups. Lepidoptera, in general, comprises approximately 146.000 species of hawkmoths, 87% of them being nocturnal. In these regions, there is a total of 7.100 species, approximately 3.100 of which are found in Brazil. Sphingidae family is one of the most representative families when it comes to pollination, being more abundant in low-altitude environments. With assistance of the wind, they migrate long distances from coastal woodlands to mountainous areas during the hot rainy season. Their abundance is even greater during cloudy nights, and during new moon phases, as they are always in search of illumination. In Brazil, most of the studies focus on the morphology of individuals for taxonomic research, being few publications on their dispersal available. Aiming mapping flight paths for feeding and reproduction, the occurrence of pollination is estimated, enabling the connectivity of forest fragments and ensuring gene exchange. To visualize this study, atmospheric models of breeze circulation such as Brazilian Regional Atmospheric Modeling System (BRAMS), GRADs, and the Three-Dimensional Kinematic Trajectory (TC3D) were employed in the Atlantic Forest of the state of Rio de Janeiro, where observations and descriptions of over 80 species are accumulated, allowing the identification of spatial patterns through the use of Geographic Information Systems. The north/south orientation proved to be dominant, potentially connecting fragments of forests with varying sizes, shapes, and conservation states, extending from coastal areas to the mountainous regions within the southeastern part of Brazil.
Department of Geography, Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
André Zaú
Department of Biosciences, Federal University of the State of Rio de Janeiro, Rio de Janeiro, Brazil
Jorge Oliveira
Department of Geography, Fluminense Federal University, NiteróI, Brazil
*Address all correspondence to: mdarrochella@gmail.com
1. Introduction
The Atlantic Rainforest is the second largest tropical rainforest of the American Continent and originally was extended along the Brazilian coast up to east Paraguay and the northeastern Argentina. Currently, the forest is recognized as one of the 25 global biodiversity hotspots1 due to its over 8.000 endangered endemic species, even though being reduced to less than 7% of its original area [1]. Other authors [2, 3] indicate that there is approximately 15% coverage of the Atlantic Forest in Brazil, including fragments in early succession stages and reforested areas, which actually do not in fact qualify as remaining forest.
The fragmentation of the Atlantic Forest has detrimental effects on biodiversity. The reduction of forested areas, whether for urban development or agricultural practices, has drastically diminished and transformed habitats, potentially leading to extinction of species, the blocking of migration routes and hindering genetic exchange [4]. Genetic exchange guarantees the maintenance of the gene pool2, on an evolutionary scale, which enables certain species to have more resistance to pests or changes in the physical environment [5].
Fragmentation occurs when an ecosystem is subdivided by human actions or even natural disturbances, such as fire, resulting in a landscape where only a few remnants of the original vegetation cover remain inserted in a matrix with completely different characteristics [6].
Depending on spatial orientation, the shape, and size, forest fragments can become less isolated, theoretically then, reducing the possibility of population decline due to mortality. For this reason, some authors [7, 8] emphasize the importance of the dispersal syndromes as the basis for connectivity, which is not necessarily, is what is used as basis to establish an ecological corridor [9, 10, 11]. In similar fashion, priority should be given to the creation of large-scale conservation units, since small fragments, even when arranged in corridors, are subject to both area and edge effects, which may hinder dispersal, migrations, and barrier crossing.
In this regard, the Guapiaçu-Macacu watershed, located in the county of Cachoeiras de Macacu (RJ), shows as a fertile ground for research on connectivity due to its spatial configuration of Atlantic Forest fragments with different shapes, sizes, and degrees of isolation in a pasture matrix. Furthermore, this biome is recognized as one of the 18 centers of lepidoptera dispersion in the neotropical region [12] and yet still exhibits an annual extinction rate of 1.8% within Lepidoptera order [13]. This fragments form a corridor that integrates themselves in private areas and diverse conservation units. Understanding how gene exchange occurs in these fragments can both make their management more efficient, as it can allow the possibility to identify problems and potentialities that assist in the creation and management of other conservation units. In a similar fashion, understanding the orientation of dispersers migratory paths3 can allow us to improve the planning and execution of ecological restoration projects for the establishment of ecological corridors.
To evaluate the dispersal syndrome, it is necessary to understand the seed and pollen dispersers of the area. When it comes to a matrix dominated by rural areas with patches of Atlantic Forest nonetheless, it is understood that terrestrial dispersers are not as efficient on promoting connectivity and genetic exchange. Therefore, for long-distance dispersal, seed and pollen require aerial transportation, which can be eased by the wind or a flying agent [15].
Looking over the map of use and land cover of the watershed (Figure 1), it is possible to have a preliminary overview of the landscape conjuncture and structure, which appears as a pasture matrix with forested water dividers. In the center, numerous forest fragments of different sizes are arranged spatially forming a corridor in the SW-NE direction. Bordering this fragment, we have the presence of some agricultural areas. It is noticeable that urban occupation in the center of the watershed is incipient, with a greater concentration to the south and northeast. The area covers approximately 573, 54 Km2 and is a part of the Guanabara hydrographic watershed, located at the Atlantic slope of the Serra do Mar (locally known as Serra dos Órgaos) with rivers draining into the Guanabara bay [16].
Figure 1.
Processed image from sentinel 2ª satellite of the land cover in the Guapiaçu watershed.
The vegetation in the area is classified as dense ombrophilous forest of submontane formation (ranging from 50 to 500 m in altitude), on slopes and lowland forests on the hills (ranging from 5 to 50 m in altitude). The area displays high structural and floristic diversity; however, most of the fragments are found stripped of their original diversity [17].
The study area is under the influence of the South Atlantic Subtropical Anticyclone (ASAS) which is a system of great scale, semi-stationary, in action all over the year, but presenting variations in its position, depending on the season of the year, and the passage of cold fronts in the area.
In the summer, the strong surface heating makes it less intense over the continent and more intense over the sea. At winter, when the continent is colder than the sea, the ASAS advances on the continent, exerting a strong influence over the study area [18].
At mesoscale, the study area is under the influence of maritime, land, valley, and mountain breezes. The maritime and continental breezes occur due to the differential heating between the ocean and the continent. When the wind blows from the ocean to the continent, it is named “maritime breeze,” and when it blows from the continent to the ocean we address it as “land breeze.” The mountain breeze occurs when cold mountain air descends down the slope during the night. The valley breeze occurs during the day when warm air rises up the slope. As the study area is located in a valley and mountainous region, it is under the influence of all four breezes in different sectors.
The breezes occur within the Planetary Boundary Layer, situated between the first 2 Km of the troposphere, where the airflow (wind) is strongly influenced by the surface roughness (relief, vegetation, buildings, etc.). The friction with these obstacles is significant, thereby reducing the wind speed [19].
Lepidopterans are approximately 146.000 species of butterflies and hawkmoths, 87% being nocturnal and 13% diurnal [20]. In the neotropical regions, they add 7.100 and 7.900, respectively, in Brazil occurring between 3.100 and 3.200 species [21].
It is viable to characterize lepidopterans as holometabolic organisms, as they undergo a complete metamorphosis in four stages of their life cycle, as follows (Figure 2): the first stage is the egg; the second is the larval phase (popularly known as caterpillar), feeding as herbivores and growing; in a relatively shorter third phase, they transform into a pupa, enclosing themselves in a cocoon, where physical changes such as the development of wings occur; finally, the fourth stage is the winged one in the form of butterflies or hawkmoths, in which they will reproduce and eventually die [22, 23]. The fourth stage lasts on average 2 months, in which the lepidopterans can fly long distances to feed on nectar and potentially contribute to pollination [23, 24].
Figure 2.
The last three life stages of a Lepidoptera. Adapted from [22].
Hawkmoths are the least studied lepidopterans in tropical areas, and the Sphingidae is one of the most representatives when it comes to pollination. These insects are commonly found in low-altitude environments and comprise over 50 species [25]. In a study conducted in a High Mountain Forest area4 at southeastern region, it was observed that hawkmoths visited more than 80 plant species. This situation is ideal for the chosen study area. The same authors state that hawkmoths migrate from restingas to mountainous areas during the hot and rainy season.
Hawkmoths, for the most part, have nocturnal flight habits, which are closely related to temperature and precipitation5 increase, also under the influence of lunar phases [26]. At Serra do Mar in Paraná, it was perceived an increased flight activity under cloudy skies, followed by drizzle or fog, with high temperatures and moderate winds [27]. This behavior is believed to be driven by hawkmoths that need to seek light, which leads them to migrate and allows pollen dispersal. Warmer areas tend to have highest abundance of hawkmoths, although this does not necessarily reflect on the number of species [25]. Flowers pollinated by hawk moths typically exhibit nocturnal anthesis, pale coloration, nectar rich in sucrose, and sweet floral scent [22].
Recent studies investigating pollen residues and hawkmoths collected in São Paulo (Picinguaba) recorded pollination interactions between sphingid lepidoptera and the families Apocynaceae, Asteraceae, Convolvulaceae, Malvaceae, Fabaceae, Scrophulariaceae, and Rubiaceae which were already known to occur, and for the first time occurrence, Bromeliaceae, Arecaceae, Begoniaceae, Celastraceae, Combretaceae, Cyperaceae, Erythroxylaceae, Malvaceae, Melastomataceae, Piperaceae, Poaceae, Polygonaceae, Sapotaceae, Scrophulariaceae, Solanaceae, Ulmaceae, and Verbenaceae [26]. Authors emphasize that in terms of abundance, the most prevalent families were Rubiaceae (15,6%), Bromeliaceae (7,8%), and Fabaceae (6,2%).
The studies at the state park of Serra dos Órgãos, in the Picinguaba and Santa Virgínia nuclei, located in northern coast of São Paulo state, provide important contributions to the identification of families pollinated by sphingid hawkmoths. However, there is progress, as it also provides species information [25]. The identified species by the author are Apocinaceae, Caprofoliaceae, Chrysobalaneceae, Dabaceae, Malvaceae, Orquidaceae, Rubiaceae, Solanaceae, and Zingiberaceae.
The most representative families in terms of species number in this study were Orchidaceae, Rubiaceae, and Solanaceae. This is a pattern, also found by [26] in the same study area. The relationship between hawkmoth flight and increased temperature and precipitation was also evidenced by studies in the zone of Mata Mineira [28]. In their collections, the maximum number of individuals were observed in the months of January and February, while the minimum were observed in August. Regarding the flight of sphingid hawkmoths, it is known that:
These Hawkmoths are excellent flyers and have a very rapid wing beat; some fly during the day, but mostly, moths are active only at night. A large number of them feed in a manner very similar to hummingbirds, hovering in front of the flower and extending their proboscis into it [29].
The majority of hawkmoths that inhabit the Atlantic Forest, particularly in the area of Cachoeiras de Macacu and Serra dos Órgãos, show a higher number of individuals during nocturnal and crepuscular flights, in autumn months and under light rain or high cloudiness. Regarding the moon phases, the same authors describe the new moon as the ideal situation, as the absence of light prompts hawkmoths to search for this resource [30].
The chosen control point for obtaining the flight trajectories of the sphingid hawkmoths was the Guapiaçu Ecological Reserve (REGUA). This choice was based the fact that a significant number of species described in the reference catalog [31] were found in REGUA. Furthermore, one of the authors works there and regularly observes and describes the behavior and morphology of hawkmoths.
REGUA (Figure 3) covers and manages approximately 7.1000 hectares and was established and registered in 2001 as a nonprofit private association [31]. The reserve is sustained through tourism, mainly birdwatching, homing to over 500 bird species. It also boasts a great diversity of dragonflies and moths, with occasional visits from jaguarundi, capybaras, and even a jaguar (locally known as “onça pintada”) [30].
Figure 3.
Control point at the Guapiaçu ecological reserve. A—General view, B—Field visits and C—Map of the reserve. Personal archive.
From the analysis of the “Sphingidae Guide of Serra dos Órgãos” [31], it was possible to identify 110 species present in the study area and their flight months throughout the year (Table 1). It is worth to say though that some species may fly in different months around the year.
Number of species found flying in Cachoeiras de Macacu (RJ) by the month of the year.
Data in mm, taken from [32] for the period of 1968–2009.
Using Pearson’s correlation test, we obtain the index of 0.3037, indicating a weak correlation between the number of species and precipitation. It is noticeable nevertheless that the typically drier months, such as May and June, coincide with lower species numbers. On the other hand, the month with the highest occurrence was March, which denotes the transition from summer to autumn, confirming the previously mentioned findings [26].
The most efficient criterion for selecting specific species to be analyzed is abundance, which reflects the most representative ones in the area of study [31]. However, during the construction of the hawkmoth [31], there was not exactly the concern of generating abundant data, given the objective was solely to identify and describe the species.
As an alternative for selecting the most representative species in the area, we chose to rely on the expertise of Dr. Jorge Bizarro,6 a specialist zoologist with over 20 years of study on the subject. In the interview, 13 species were indicated as the most representative (Table 2), which are included in the catalog.
Species
J
F
M
A
M
J
J
A
S
O
N
D
Adhemarius daphne daphne
X
X
X
X
X
X
X
X
X
X
X
X
Manduca diffussa petuniae
X
X
X
X
X
X
X
Manduca hannibal almicar
X
X
X
X
X
Enyo lugubris lugubris
X
X
X
X
X
Enyo ocypete
X
X
X
X
X
Nyceryx coffaeae
X
X
X
Nyceryx riscus
X
X
X
X
X
Pseudosphinx tétrio
X
X
X
X
X
X
X
X
X
X
Erinnyis alope alope
X
X
X
X
X
X
X
X
X
X
X
Erinnyis ello ello
x
x
X
x
x
x
x
x
X
Xylophanes chiron nechus
x
X
x
X
x
x
x
x
x
X
Xylophanes porcus continentalis
x
x
X
x
x
x
X
Xilophanes tersa tersa
x
x
X
x
X
x
x
x
x
x
Table 2.
Hawkmoth species most observed in REGUA and their months of appearance.
From this table, it is possible to understand which species fly in which seasons of the year and thus infer about pollination by month and season. It is also possible to understand which species are more tolerant, such as Adhemários daphne daphne, that was observed every month of the year, and the less tolerant species, such as Nyceryx coffaeae, observed in only 5 months. The month of March was also the month in which the most representative species of the area were seen, with exception of Xylofanes porcus continentalis.
Recognizing the geographic distribution of these species allows us to speculate that their local behavior can be extrapolated to other areas with similar topography, climate, vegetation cover, and land use. To facilitate studies, the use of atmospheric modeling was chosen, as it presents a more feasible alternative in the absence of local baseline data, allowing approximations to be generated regarding the atmospheric behavior of these areas.
Models are simplified approximations of reality, resulting from how we perceive it, subjective representations of our interpretation as a result of rationality. Above all, it is a methodological approach, constructing “a simplified and intelligible framework of the world” [33].
Models are used according to specific objectives, which are expressed in their functioning and structure, serving as parameters for choosing which one to use. Therefore, for a research project that relies on atmospheric dynamics, it is important to remember the dependence between the atmosphere and the characteristics of the Earth’s surface due to the inherent exchange of heat, moisture, and linear momentum, with exchanges occurring between the surface and lower layers of the atmosphere. In this way, it is possible to define local forcing and local determinants for the behavior of currents, winds, and breezes [34]. This can be expressed in models.
Currently, Numerical Weather Prediction (MNT) allows us to forecast events with a 15-day lead time, and 5-day accuracy, which is possible thanks to the computers at the Centre for Weather Forecasting and Climate Studies (CPTEC) of the National Institute for Space Research (INPE) and other institutions that combine regional- and global-scale data [34]. For climate models to yield reliable results, it is necessary to dispose of a solid database as long as a thorough understanding of the variables influencing the process, as well as a long period of data collection.
CPTEC employs various models such as the Global Circulation Model of the Atmosphere (GCM) for global weather prediction and ETA for regional prediction. The Regional Atmospheric Modeling System (RAMS) and the Brazilian Regional Atmospheric Modeling System (BRAMS) are used for studying spatially limited areas with high spatial resolution. BRAMS allows the interpolation of data from nearby meteorological stations to create wind field analysis (Figure 4), while GRADS provides analysis at three different scales as grids (Figure 5) [36].
Figure 4.
Example of wind and temperature field generation in model BRAMS model with graphical interface of the TC3D model. BRAMS does not work under a graphical interface. It is a mathematical model, being necessary then the use of the program “GRADS” to further visualization of the generated atmospheric data. Source: [34].
Figure 5.
Examples of grids that can be analyzed with the GRADs model. BRAMS does not operate under a graphical interface. BRAMS is a mathematical model, requiring the use of the GRADS program for the visualization of generated atmospheric data. Source: [35].
One of the major advantages of numerical modeling of the atmosphere is the possibility of obtaining spatial and temporal information on a much larger scale than available in conventional synoptic networks. However though, it is important to validate the models for tropical atmospheric conditions, as most of them were developed for mid-latitude atmospheres on a global scale [35].
BRAMS, along with the GRADS (Figure 5) and Three-Dimensional Kinematic Trajectory (TC3D) programs, offer an advantage in this regard, as they have been adapted for validation in the tropical atmosphere. Using these models, it is possible to analyze potential connectivities and dispersal trajectories of hawkmoths at three different scales, with the third grid covering the entire watershed, the second grid connecting it to the upstream mountainous region, and the first grid covering the entire Serra do Mar.
BRAMS is a regional-scale numerical model, adapted from RAMS, developed by the University of Colorado, which allows us to predict variables such as temperature, wind, humidity, and precipitation. It solves the equations of atmospheric dynamics and includes numerous submodels that relate soil-vegetation-atmosphere system, turbulent flux exchanges, radiative transfers, cloud microphysics, and more [37].
The BRAMS was adapted to represent the state of the tropical atmosphere [35]. The equations used include the equations of motion, thermodynamics, continuity for the mixing ratio, and mass continuity. With parameterizations, it is possible to generate information on solar radiation, humidity processes (clouds, liquid, and ice precipitation), sensible and latent heat, soil layers, water surfaces, vegetation, among others [38].
As the nights of new moon are favorable for the increased occurrence of sphingid hawkmoths, all new moon nights [39] from the years 2015, 2016, 2017, and 2018 were investigated, each month, based on observations obtained from the website of the Department of Astronomy at the University of São Paulo (USP), in order to run the BRAMS model for each of these nights, simulating their atmospheric conditions. This way, it was possible to understand under which atmospheric conditions the moths could have flown.
With this, it is possible to simulate the atmospheric conditions for each month, for the species of hawkmoths found during this month. Each new moon event presents the ideal conditions for hawkmoth flight, allowing the calculation of the flight trajectory for each species, since they could theoretically take advantage of the winds to minimize energy expenditure. Similarly, the data can be divided by season, to understand certain wind patterns and flight behavior. From the flow field (generated by the TC3D model), trajectories can be calculated forward (departure from a specific point) and backward (arrival at specific location). At this research, the team chose to use a departure point, the REGUA, which is believed to be the origin of dispersion of the described and observed hawkmoths.
To determine the possible flight trajectories from the wind field (Figure 6), The Three-Dimensional Kinematic Trajectory (TC3D) model was adopted. This is a nonconvective three-dimensional model applied over the surface, which allows estimation of directions and altitudes of particles suspended in the atmosphere based on wind circulation data7.
Figure 6.
Example from application of the TC3D model.
The model uses a color scale that makes possible to estimate the altitudes reached by suspended particles. The author emphasizes that the application of this model enables the estimation of both outgoing and incoming trajectories, defining their vertical and horizontal paths within the atmospheric mixing layer [40].
Having REGUA as a reference point, it is possible to present flight trajectories of moths for each new moon night throughout the year, indicating which species would potentially be involved in pollination and the distances they could reach.
After conducting all the simulations for the fur dates in each season of the year, over 4 years, the trajectories were georeferenced using the ArcGIS software, creating a layer overlaying the satellite image of the area at the GRADS grid level 2 scale. All trajectories were vectorized to create a single synthesis map containing the 16 simulations, providing an overview of the spatial and biogeographic arrangement of the pollination potential and connectivity of the Southeast Brazilian Atlantic Forest.
The preprocessing, processing, and postprocessing steps were performed using the BRAMS model, and the calculation of the trajectories was done using the Three-Dimensional Kinematic Trajectory (TC3D). Using the BRAMS model, it was simulated the wind field, while the TC3D model calculated the potential trajectories of a particle dispersed by the wind from a specific point, on a predefined date and time.
For this purpose, four nights of the new moon were chosen throughout the year, with intervals of 2 to 3 months, once each season. The models generated results for each season from 2015 to 2018. For each year starting from 2015, an additional day was added forward, as new moon events occur within a weekly period.
The interpretation of trajectories is based on state delimitation on a map without the geographical characteristics of the area. Thus, it is possible to identify only the direction, speed, and altitude at which an inert particle moves in the atmosphere. This way, the hypothesis was made that hawkmoths can take advantage of the sea and land breezes systems and potentially can follow the trajectories, given that they have full flight autonomy.
Breezes are atmospheric phenomena resulting from the differential heating between the land and the ocean. They are mesoescale phenomena whose reach depends on the topography. They occur in the planetary boundary layer which is in the first few kilometers above the surface.
It is important to note that even though there are no geographical elements in the graphic representation of GRADS, the evaluation is not arbitrary but requires some level of abstraction and knowledge of the landscape characteristics of the area and the political division of municipalities. This allows to make assumptions about the landscapes where a particle disperses, as well as its final destination during the chosen 24-hour period.
5.1 Summer trajectories
The trajectories were simulated starting at 8 PM, considering daylight saving (Figure 7) during the twilight period, for 4 years starting in 20158. The species observed in January and following these trajectories are Adhemarius daphne daphne, Manduca diffussa petuniae, Manduca hannibal almicar, Enyo lugubris lugubris, Erinnyis alope alope, Erinnyis ello ello, Xylophanes chiron nechus, Xylophanes porcus continentalis, and Xylophanes tersa tersa. That means that potentially 10 out of the 13 most representative species could follow these trajectories.
Figure 7.
Potential summer trajectories.
Regarding the predominant direction of the sea breeze system, it is evident that during summer, the behavior did not change drastically over the four simulated years. There was a predominance of the land breeze, blowing toward the south and southwest.
5.2 Autumn trajectories
The new moon experiment in March, as a sampling for autumn, presents the species that would follow such trajectory, including among them: Adhemarius daphne daphne, Manduca diffussa petuniae, Manduca hannibal almica, Enyo lugubris lugubris, Enyo ocypete, Nyceryx coffaeae, Nyceryx riscus, Pseudosphinx tétrio, Erinnyis alope alope, Erinnyis ello ello, Xylophanes chiron nechus, and Xylophanes tersa tersa. That shows that potentially 12 out of the 13 most representative species can fulfill these trajectories.
It is noticeable that there is no standard behavior (Figure 8). In the years 2015 and 2017, there was a predominance of the sea breeze in the north direction, with changes in direction after crossing the limits of Serra dos Órgãos. In 2016 and 2018, the initial direction of the land breeze was not the same but later followed the southwest pattern.
Figure 8.
Potential autumn trajectories.
5.3 Winter trajectories
Regarding the species that fly in June, as a sampling for the winter period, we have then: Adhemarius daphne daphne, Enyo lugubris lugubris, Pseudosphinx tétrio, and Erinnyis alope alope. This demonstrates that out of the 13 most significant hawkmoth species in the area, only 4 of them would potentially follow that trajectory. The winter season breezes (Figure 9) behaved similarly throughout the years 2015, 2016, and 2017, originating from the sea with an initial direction tending toward the northwest. Among these 3 years, 2016 showed a tendency toward the northeast at the end. The year 2018 presented a very different result, with the occurrence of the land breeze, following a westward direction, with a tendency toward the west and later south.
Figure 9.
Potential winter trajectories.
5.4 Spring trajectories
September represents the beginning of spring in the Southern Hemisphere and includes the species here mentioned that would potentially follow this trajectory: Adhemarius daphne daphne, Manduca hannibal almicar, Enyo lugubris lugubris, Enyo ocypete, Nyceryx coffaeae, Nyceryx riscus, Pseudosphinx tétrio, Erinnyis alope alope, Xylophanes chiron nechus, Xylophanes porcus continentalis, and Xylophanes tersa tersa. This trajectory is potentially followed by 11 out of the 13 most representative species in the area.
Analyzing the behavior of spring breezes (Figure 10), it is noticeable that in the years 2015, 2017, and 2018, there is a tendency toward the south-southwest direction, indicating a continental origin. Only in the year 2016, the dominance of the sea breeze occurred, with a trajectory toward the northwest and later toward the northeast.
Making an observation of all the trajectories, it becomes evident that the majority occurred in the north/south directions, which is a different pattern than expected—southwest/northeast, following the orientation of the river drainage. Only 3 out of the 16 trajectories followed this pattern. It also became evident that in the autumn and winter trajectories, there is a predominance of the sea breeze, oriented toward the north. Similarly, in spring and summer, there is a predominance of the land breeze toward the south.
Observing the synthesis map (Figure 11), it is possible to notice that in the northern direction of REGUA, there is an abundance of small to rounded fragments and a higher number of deforested areas or abandoned pastures. This indicates that under the influence of this potential orientation of the hawkmoth pollination syndrome, there is a pathway that can serve as a basis for planning new ecological restoration projects or the implementation of new conservation units.
Figure 11.
Potential flight paths of hawkmoths. Legend: Circle in orange—REGUA, black lines—Winter trajectories, red lines—Summer trajectories, pink lines—Spring trajectories, yellow lines—Autumn trajectories.
They can reach the state of Minas Gerais, and along their path, are able to potentially promote connectivity with the high-montane ombrophilous forests in the Serrana region of Rio de Janeiro state, where important conservation units are located. Subsequently, they can cross the Paraíba do Sul river valley, connecting the fragments of semi-deciduous stationary forests.
The trajectories in south direction extend beyond the coastal limit, reaching the ocean along their path, and they traverse landscapes of rolling hills, submontane ombrophilous forests, abandoned pastures, as well as coastal sandy areas or restingas.
The three trajectories at southwest direction would allow connectivity with mangrove ecosystems and submontane ombrophilous forests, as well as important conservation units with low- and mid-montane ombrophilous forest in the metropolitan area of Rio de Janeiro city.
At the end of the trajectories, it is noticeable the reach of the inland stationary forests, where forest cover is more sparse, with thousands of small fragments, possibly disturbed and under the influence of the edge effect. It is interesting to note that such pollination syndrome via the effect of breezes is not present in the East-West directions, with basically no connectivity between REGUA and the Costa Verde and the lakes region of Rio de Janeiro (locally addressed as “Região dos Lagos”).
In this sense, it is believed that the Atlantic Forest fragments in the county of Cachoeiras de Macacu, as well as their corresponding watersheds and aerial areas, potentially participate in a large dispersal pathway through the breeze system, extending from coastal ecosystems to high-altitude ecosystems in the mountainous inland areas.
The spatial arrangement of connectivity presented should still be seen as a potential, and its validation will only be possible with observations and markings of the studied fauna, which is methodologically possible, but was financially unviable for this research. At least, it is noteworthy that the species Adhenarius daphne daphne and the order Manduca can be seen as important bioindicators of connectivity, given their body size and occurrence throughout all of the months of the year.
The study area, as a representative sampling of the Atlantic Forest conjecture, in the context of habitat fragmentation and ecological restoration action plans, served much more than a locus of individual analyses, revealing connections between fauna and flora with distant areas. It is encompassed mountainous areas covered by fragments of the Atlantic Forest with ecosystems of semi-deciduous seasonal forests, open ombrophilous forests, mixed ombrophilous forests, advanced successional ombrophilous forests, as well as submontane coastal lowlands with ombrophilous forests, restinga ecosystems, and mangroves. This study covered a vast mosaic of landscapes through the application of integrated methods from ecology and meteorology.
Little is still known about flight trajectories of lepidopterans in Brazil, with at most the knowledge of where they may occur. Therefore, we present a possible spatial arrangement, as vast Brazilian literature demonstrates the occurrence of the species discussed in this chapter in the areas where the TC3D model simulated the trajectories.
We sincerely hope that the modeling will serve as an incentive for new researchers to seek methodologies to understand the connectivity of the forest fragments and landscapes based on pollination syndromes.
This research was made possible thanks to numerous partnerships between different individuals and institutions. First and foremost, we would like to thank Mr. Nicholas Locke, manager of the Guapiaçu Ecological Reserve (REGUA), for his support in the fieldwork and accommodation, as well as Dr. Jorge Bizarro, research director of the reserve.
We would like to express our gratitude to the Engineer José Maria de Castro Junior, for his assistance in using the BRAMS, GRADs, and the TC3D models.
A special thanks to the biologist and chemist Nathália Pombo Gil for reviewing and translating the original text.
Lastly, we extend our gratitude to the institutional support provided by the Department of Geography at Federal Fluminense University, the institute of Biosciences at Federal University of the State of Rio de Janeiro, the Graduate program in Geography at Federal University of Rio de Janeiro, and the Carlos Chagas Filho Foundation for Research Support of the State of Rio de Janeiro.
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Notes
The term refers to prime concern areas for biodiversity conservation worldwide [14].
The term can be understood as genetic elasticity.
Pollen and seeds.
Forests with altitude greater than 1600 m [25].
The low luminosity of the sky on rainy days causes more hawkmoths to fly in search of luminosity [23].
Jorge Bizarro is a PhD in Zoology, co-author of the hawkmoth catalog, and REGUA director (Ecologic Reservation of Guapiaçu), used as a reference on a great number of scientific papers.
It is worth mentioning that TC3D model does not display detailed land cover and topography in its graphical interface. The program only uses political boundaries on a flat white surface. The vectors, however, represented by the particle in ascent trajectories are colored to express their altitude, and points with time stamps, providing information on the speed of dispersion.
The research considers that each event simulated by the model represents a flight path for feeding and reproduction.
Written By
Marcio D’Arrochella, André Zaú and Jorge Oliveira
Submitted: 13 July 2023Reviewed: 14 July 2023Published: 19 January 2024
Open access peer-reviewed chapter
