1. Introduction
The high longevity of some plants in high-stress environments (e.g., nutrient-deficient environments, high competition, or extreme climates) means that they can survive under adverse conditions long enough to delay or even avoid extinction by preserving their populations throughout long hostile periods until windows of opportunity allow for their establishment [
1]. Such a strategy of the long-term maintenance of individuals in high-stress environments is known as persistence [
2]. Monitoring persistent populations is complex since they grow slowly and the environmental conditions when they were established in the past may have now changed [
3], but they can inform on the long-term environmental changes they have witnessed and the disturbances they have gone through [
4,
5,
6]. The extreme environmental conditions that persistent plants suffer determine the usually slow dynamics of these populations [
7,
8,
9]. For instance, persistent shrub populations in the Mediterranean high mountains are under intense abiotic stress and are dominated by older adults, with meager proportions of juveniles [
10], something that could also be favored by anthropogenic disturbances (e.g., deforestation and fires). This evolution as a function of environmental conditions and anthropogenic disturbances complicates their study and monitoring over long periods of time [
4,
5]. Furthermore, there is a lack of information on how the persistence of these individuals contributes to expanding the demographic strategies of the species [
7]. For these reasons, low-cost automatic monitoring of the distribution and growth of persistent shrubs with very high-resolution aerial and satellite imagery would facilitate an assessment and understanding of the effects of environmental changes and disturbances on the dynamics of long-lived plant populations.
Wildfire is the primary disturbance in the Mediterranean ecosystems of southern Europe [
11]. Fires are natural agents that have played a fundamental role in the evolution of ecosystem patterns and processes since prehistorical times [
12]. Fire action can create new ecological niches [
13,
14], be responsible for modifications in vegetation structure and distribution [
15], cause changes in erosion regimes [
16], and affect the carbon cycle and even the global climate [
17]. Fires are sometimes associated with the loss or degradation of natural areas and their associated ecosystem services (forest, livestock, and agricultural resources) [
18]. Nevertheless, there are fire-adapted ecosystems and species for which its loss is detrimental [
13]. Responses and degrees of adaptation to fire vary among plant species and some species can recover after disturbances, as is the case with resprouting [
19]. In other species, the capacity to recover after fire disturbances is minimal, and if individuals die, the only option to recover their populations is by new colonization [
10,
20]. For many years, fire regimes have been influenced by humans (e.g., [
21]). However, the cumulative effects of human actions are leading to changes in the size, intensity, and recurrence of fires, resulting in more severe impacts on biodiversity and ecosystems [
13,
18].
Aerial and satellite image classification for vegetation monitoring is one of the most widely used remote sensing tools in conservation [
22,
23,
24,
25]. Object-based image analysis (OBIA) has proved to be a successful tool for monitoring the effect of fires in the past (e.g., [
26,
27,
28,
29]), providing more accurate results with high-resolution images and without spectral information than pixel-based analysis, which presents difficulties in images of very heterogeneous environments [
30]. Pixel-based classification analyzes each pixel’s properties independently without considering spatial information or the context surrounding the pixel of interest. In addition, the small pixel size in images of very high spatial resolution means that the objects of interest are often larger and important information in the context of the image is lost through pixel-based image analysis [
31]. In contrast, OBIA considers both the spectral information of the object and its context, including its relationship with the environment [
32]. OBIA consists of two steps: segmentation, which is the division of the image into homogeneous objects (also called segments) of similar characteristics and the classification of these objects, which is their categorization into classes of interest, based on similarities in their shape, context, and spectral information [
33]. During the segmentation phase of OBIA, it is necessary to establish an appropriate scale for the division of objects according to their size in the analyzed image [
34]. Therefore, the success of the classification greatly depends on the accuracy of the segmentation [
35]. In environments where vegetation appears differentiated from the surrounding landscape, such as in high mountains, OBIA allows accurate and realistic identification by using the spectral information of the vegetation and its spatial relationship to its context. The availability of high-resolution historical photographs and their analysis with OBIA makes it possible to reconstruct the dynamics of plant populations and evaluate their changes over time.
In this study, the historical mapping of a persistent shrub (Juniperus communis) in the Sierra Nevada National Park (Spain) is constructed as a basis for evaluating its population dynamics over 40 years and its evolution after a fire that occurred in 1983. For this purpose, (i) the remote sensing identification of Juniperus communis was optimized by developing a semi-automatic work protocol that allows identifying which segmentation (separation between bush and ground) is best adapted to the characteristics of each moment in the historical images, allowing better identification of the bushes; (ii) a reconstruction of historical changes in the distribution, canopy cover, and population structure of Juniperus communis was carried out to evaluate the possible factors that have determined its post-fire recovery; and (iii) changes in the size structure of the population were compared to bibliographic references and discussed in the context of the different measures adopted to protect the area as a Natural Park in 1989 and as a National Park in 1999.
4. Discussion
In this work, we developed a protocol for automatically monitoring persistent shrub populations (i.e.,
Juniperus communis L.) over several decades by a semi-automatic optimization of OBIA on very high-resolution aerial or satellite images. The semi-automatic extraction of the combination of parameters that provides the best segmentation of the images by OBIA is the main contribution to this article, being normally a very time-consuming process [
48]. According to [
46], determining the best segmentation parameters is a laborious process. Nevertheless, this work shows a method to determine the effect of the variation of the segmentation parameters (scale, shape, and compactness) highlighting the scale in this technique [
44,
58], as well as the effect of the variation of the other parameters on the result. The increasing use of OBIA in different fields of biodiversity conservation highlights its success in vegetation monitoring, having been used for multiple purposes, such as forest inventories [
59,
60], changes in vegetation cover [
61], or studies of vegetation structure [
22]. OBIA has been used for monitoring long-lived animal species [
62] but not persistent plants after a fire, as we have performed in this work.
The positive effect on the population of
J. communis of the designation of Sierra Nevada as a Natural Park in 1989 and as a National Park in 1999 can be observed. After Sierra Nevada acquired both protection statuses, the area occupied by
J. communis increased, perhaps due to the increased protection of the individuals that were recovering and colonizing after the 1983 wildfire. Nevertheless, an increase in the number of individuals is only observed after the creation of the National Park in 1999, which may be due to this increased protection or to the slow growth of junipers, which may delay the moment of detection by aerial images due to the small size of the juveniles [
10]. In the period of 1977–1984, there was a great decrease in the number of individuals due to the fire that occurred in 1983.
J. communis is a species whose seeds largely die after fires and has no ability to resprout [
10,
20]. The largest junipers were less affected by fire but smaller individuals were greatly affected (
Figure 6), as occurs in other species [
63]. Such homeostasis of the largest individuals may be due to the typical strategy observed in persistent species. This, together with the low success in the recruitment of the species in the study area and, therefore, the loss of colonization capacity of the burned areas [
10,
20,
36], has led to a lack of juniper regeneration, so only the largest individuals remained, not reaching levels of canopy cover similar to those detected in 1977. In this area, there has also been strong pressure from livestock and irrigation channels, which have now been largely abandoned [
40,
41]. In this context, a decrease in human populations in the Sierra Nevada has been observed in recent decades, which may imply an improvement in
Juniperus communis populations due to a decrease in human land use [
41,
64]. Nevertheless, the almost null recovery of
J. communis after anthropic disturbances is remarkable [
10,
20]. The improvement of this population does not seem to be affected by the decrease in habitat quality [
65], despite the fact that an increase in temperatures and a decrease in the protective snow cover has been observed in recent decades, with an increase in dryness in the area and an extreme drought in 2005 [
66]. Although the populations of
J. communis of Sierra Nevada have been reported to be highly resistant to snowfall, summer drought, browsing, and trampling by livestock [
42,
66], it is necessary to avoid further losses in these populations given the complexity of their restoration [
42].
Although the method described in this work can provide high accuracy, the results should be analyzed with caution. Shadows from tall specimens can influence segmentation, delimiting areas of uncertainty [
67,
68]. In addition, the spatial resolution and contrast of the images are slightly variable, which can lead to an overestimation of individuals in some cases, especially in panchromatic or low-resolution images [
46,
69]. To minimize the effect of errors in image acquisition, which can affect the identification and classification of individuals [
70], it is necessary to take into account the time of year, day, and time of image captures. Likewise, the large size observed in some shrubs may be due to the union of several individuals in the same patch of dense vegetation. This makes it practically impossible to differentiate them by OBIA due to their similar characteristics, or even to identify them in the field, which can lead to an underestimation of the number of individuals in some cases [
61]. For better identification of individuals in future studies, it may be useful to use hyperspectral or multispectral images [
71] together with deep learning [
72] to facilitate the identification of individuals with similar characteristics.
The selection of an appropriate range of canopy sizes and shapes when applying examples for both segmentation and classification validation can affect image analysis [
56]. The detection of small individuals may be conditioned by the spatial resolution of the images, which will generate more or less accurate segments with respect to reality depending on this resolution. Likewise, a classification may appear to be conditioned either by an excessive heterogeneity of the images, which may mean that, in some cases, an individual has been wrongly classified, or by an excessive homogeneity of the images, with insufficient differentiation between objects [
73]. The use of a larger number of images, with shorter time intervals between them can lead to more accurate monitoring of populations [
61].