1. Introduction
White oak (
Quercus alba L.) is an important tree species both for ecological significance and economic importance across the eastern United States. White oaks are dominating canopy that serves a vital role in forming stand structures and composition. For a wide variety of creatures, including nesting birds and arboreal animals, its wide, spreading crown provides a friendly microhabitat [
1,
2]. Acorns and fallen leaves provide a wealth of food for wildlife, supporting complex food chains and promoting biodiversity [
3,
4]. Furthermore, white oaks' vast root systems create essential symbiotic connections with mycorrhizal fungi that aid in nutrient cycling and uptake. The tree's capacity to store large amounts of carbon helps to reduce greenhouse gas emissions and mitigate the effects of climate change [
5,
6]. The white oak's vital function in the forest ecosystems of the eastern United States is further demonstrated by its patterns of succession and regeneration. Because of its effective acorn dispersal, the tree offers a plentiful supply of seeds for spontaneous growth, guaranteeing the continuation of its lineage for future generations. However, white oaks population is in danger due to the increased declining situation associated with changes in forest stand dynamics.
Oak mortality is among the most observed phenomena in oak forests in the Eastern U.S. [
7,
8,
9,
10]. The mortality starts with the oak tree browning of leaf, turning black, curl-up, and finally falling to the ground [
11,
12,
13,
14,
15]. Factors responsible for the oak decline are suspected long-term predisposing, short-term inciting, and contributing factors [
12,
13]. Predisposing factors are related to stand longevity and maturity, which are responsible for a tree's natural ability to growth inhibition and lead to injury-inducing agents. The inciting factors are related to physical or biological conditions, which are related to defoliating insects, hail, frost, and drought. Reports from inciting fac-tors showed typical crown dieback, browning, and new leaf emergence in dying trees, which eventually leads to deaths [
16]. The contributing factors are related to pathogenic fungi and boring insects that ultimately kill the trees. This decline has grappled with notable drought outbreaks, late spring frost, the emergence of saprophytic fungus due to climate change [
17], and oak borer attacks on the most vulnerable sites.
Typically, oak mortality has been targeted in red oak group species. More recently, white oak mortality (WOM) has become a prominent target across the eastern US. It seems that the spatial distribution pattern of WOM is not uniform. Poor resource sites [
18] such as droughty, poor drainage, and soil nutrient deficiency are more prone to WOM. It is because low resources have led to declining and widespread regeneration failure. Scientists also reported WOM in higher-quality mesic sites where the forest has gone through a high stand density and maturity stage [
19,
20]. The mortality is more prevalent in the self-thinning stage, as the tree species under high stand density struggle to utilize maximum resources [
21]. WOM has been observed in different topographies, from low-lying lands to valley floors [
22]. North-facing slopes where sunlight is low are more prone to high oak mortality [
23]. Besides, ecological stressors such as browsing, heavy shade, and disturbances have influenced white oaks in many parts of the eastern hardwood forests [
22].
The impacts of white oak mortality are not only limited to individual tree species but also affect entire forest ecosystems and ecological processes. For instance, white oaks were dying in greater numbers, i.e., 30% of healthy crowns, which were less than 4 meter in width across the 516-ha area of Ozark Highlands [
24]. Similarly, some 900-ha area of Baskett Wildlife Research and Education Center across the Ozark Border of Central Missouri depicted 10% of white oaks killed by drought-pathogen interactions [
25]. White oak mortality across forest ecosystems can lead to changes in nutrient cycling, water regulation, and carbon sequestration [
26,
27,
28]. White oak is a long-lived species and can sequester large amounts of carbon during its lifetime [
29,
30]. The decline of mature white oaks can disrupt carbon sinks and worsen the situation for climate change and carbon management [
31,
32]. White oaks vast canopy offers critical habitat and nutrition for a wide range of flora and fauna, promoting a complex web of biological interactions [
33,
34]. For instance, white oak is a dominant canopy species across the eastern US that shapes forest structure and composition influencing soil processes, species interactions, and wildlife habitat [
35,
36]. Its loss can lead to a change in forest dynamics such as impacting the recruitment of new individuals to alterations of the competitive relationship among plant species [
37,
38]. The decline in white oaks can impact numerous wildlife species and their microhabitats that rely upon shelter and foraging [
39]. Moreover, the white oak decline can have adverse effects on species distribution and food chain interactions as well [
40].
Several research concentrated on the spatial patterns of white oaks that utilized climate, topography, and several other factors combined as well as individually [
41,
42]. There is very little research done on white oak’s spatial patterns but those are focused on local level studies. The evidence did not find clear spatial patterns for white oaks across varying scales [
43,
44], which highlights the unique approach for our study. The aim of this study is (1) to identify the spatial distribution pattern of WOM across southern, central, and northern region of the eastern U.S. and (2) and explore the potential underlying processes behind the observed spatial patterns of WOM rate. We hypothesized that the observed spatial pattern of the WOM rate is randomly distributed at local scale due to localized environmental factors and stand-scale competition, while exhibiting clustering patterns at the broad scale, likely attributed to a larger-scale environmental gradients and climatic pattern.
3. Results
3.1. White Oak Mortality Spatial Distribution Patterns
The white oak mortality (WOM) rate is spread throughout the eastern United States. However, there are differences in the magnitude of mortality rate and density distribution across different locations i.e., the southern, central, and northern regions of WOM rate. The southern region consists mostly of very low followed by low and medium WOM rates across different latitudes and longitudes (
Figure 2A). However, there are also a few plots of high as well as very high mortality rates across this region. Despite the larger area of a low and very low density of WOM rate, some locations across the southern region depicted a high-density distribution of WOM rate (e.g., parts of Arkansas and Tennessee;
Figure 2B).
Numerous plots of WOM rate were gathered that has been spread across most of the central region of the eastern US. Much of the central region showed very low and low WOM rates, however, there are also numerous plots related to medium, high, and very high WOM rates across this region (
Figure 2A). This region has the most diversified forest landscape that consists of very high-density (e.g., parts of Missouri, Arkansas, Virginia etc.) and high-density distribution (e.g., parts of Missouri, Arkansas, Tennessee, Kentucky, and Virginia) of WOM rate as compared to southern and northern regions. Most of this region has been depicted with very low density as well as low and medium density distribution of WOM rate, which is greater than very high and high-density distribution (
Figure 2B).
The majority of the northern region contains very few plots of WOM rate distributed across our study area (e.g., Illinois, Indiana, and Ohio). Though this region has very few plots of high to very high WOM rate (e.g., Missouri, west Virginia, and Virginia), the overall plot distribution to WOM rate is higher as compared to other regions (
Figure 2A). The density distribution of WOM rate is mostly depicted very low and consists of a few areas of medium, high, and very high-density distribution to certain latitudes and longitudes. Despite the larger area of land mass, this region represented a very low plot distribution of WOM rate (
Figure 2B).
The spatial distribution pattern of the WOM rate showed random patterns up to 3000 m, as the observed K value was inside the confidence envelopes, and we accept the null hypothesis (
Figure 2). Beyond this point, we reject our null hypothesis, the spatial distribution of WOM rate showed clustered until 20,000 m is statistically significant. It is due to the spatial clustering of WOM rates, as indicated by the observed K value being significantly larger than the expected K function and falling outside the confidence envelopes. This clustering pattern of the WOM rate increases as the distance increases. Most of our pattern analysis for the WOM rate depicted clustered (non-random) patterns at a larger distance across the eastern US. Thus, our spatial distribution pattern analysis of WOM showed more clustering patterns than the random pattern as distance increases.
The kernel density maps for white oaks showed the cluster spatial distribution across the declining plots of white oaks (
Figure 2). This density distribution indicated areas for higher or lower concentrations across the eastern US.
Figure 2.
Plots (A) and density distribution (B) showing WOM rate across different latitudes and longitudes.
Figure 2.
Plots (A) and density distribution (B) showing WOM rate across different latitudes and longitudes.
3.2. Spatial Distribution Patterns at Southern, Central, and Northern Regions WOM Rate
The spatial pattern of the WOM rate across the southern region depicted a clustering pattern until about 360 km, and we reject our null hypothesis. This is because the observed K function is above the expected K function. Also, the observed K function, statistically significant, is completely outside the envelope. This means points of WOM rate in the dataset are more closely spaced than would be expected under spatial randomness. However, the intensity of the clustering pattern, not constant, keeps increasing as the distance increases. Most of the southern region depicted aggregation to WOM rate across the local scale. We found the pattern is random at 360 km to 395 km as the K function, statistically not significant, is completely inside the envelopes. The random pattern of the WOM rate is across a broad scale but covers a low range of distance. This random pattern shows that the point of WOM rate has an equal chance of occurring anywhere in the southern region of our study area. The pattern is mostly uniform as the distance increases from 400km to 550km, and we reject our null hypothesis. It is because the observed K function across a broad scale is consistently below the envelope and is statistically significant as well. It shows that the points of the WOM rate are more regularly spaced than would be expected under spatial randomness. Also, there is a mostly uniform pattern across the larger scale which is more or less similar to the central and northern region WOM rate.
The central region of the WOM rate depicted a slightly lower intensity of observed K function at the local scale in comparison with patterns of southern and central region WOM rates. We found the clustering pattern until 200km across the central region where the intensity of clustering is much smaller, and we reject our null hypothesis. This clustering is because the observed K function, statistically significant, is outside the envelope. This indicates that there is a tendency for WOM rate points to be aggregated together in this region. Unlike other regions, the observed K function is much near to the random line at the local scale. Similarly, there is also a random pattern from 230km to 430 km as observed K function, statistically significant, is completely inside the envelope. It suggests that there is an equal chance of occurring points of WOM mostly at the medium scale than the local scale. However, broad scale is mostly facilitated with a uniform pattern starting from 430 km and beyond, and we reject our null hypothesis. And this uniform pattern is much away from the random line and contains a slightly lower intensity of observed K function in comparison to patterns of southern and northern region WOM rate. Overall, the pattern of WOM rate seems to be more random at the local scale, but at broad scale, it is less uniform, which is similar to southern region of the WOM rate as well.
The WOM rate across the higher latitude or northern region presented more clustering patterns (until 480km) than a random and uniform pattern. We found a complete clustering pattern across local, medium, and parts of broad scales in which the observed K function, statistically significant, is outside the envelope. The clustering pattern at the local scale starts with low intensity which is also much near the random line and slightly at the peak when it reaches around 20 km. This indicates that there is a tendency for WOM rate points to be aggregated or clustered together in this region. However, clustering slightly decreases as it reaches to medium scale and stops at 470km, which is at a broad scale. Unlike the southern and central WOM rates, patterns presented by the observed K function are varied across each scale. It is because random and uniform patterns are presented only across broad scales and across a few ranges of distance. Results show that the observed K function is below the expected K function at 480km to 520km, completely inside the envelope, denoting a random pattern at a broad scale. This tells us that the points are distributed in a manner consistent with complete spatial randomness (CSR). Beyond 520km, there is a uniform pattern at a broad scale but covers few distances in this region. The uniform pattern is since the observed K function is completely outside the envelope and is statistically significant as well, and we reject null hypothesis. This tells us that points are more evenly distributed across this region, and there is a tendency for points to avoid each other.
Figure 3.
Ripley’s K function showing various spatial patterns across (A) southern, (B) central, and (C) northern region WOM rates.
Figure 3.
Ripley’s K function showing various spatial patterns across (A) southern, (B) central, and (C) northern region WOM rates.
4. Discussion
Our study found that the southern region WOM rate showed a clustered pattern at the local scale, and the reason is that competition at stand level may play some dominant roles over the WOM rate. This confirms with other findings that aggregation factors were highly responsible for stand-level competition in most of the oak tree species [
64,
66]. Evidence supports that self-thinning, which is also called natural mortality, is mostly prevalent in the oak-dominated forest of eastern North America [
67,
68]. Self-thinning occurring mostly in the stem exclusion stage if deprived of sunlight, moisture, and space may lead to mortality. Also, the spatial pattern of the southern region WOM rate indicated a random pattern across a few ranges of broad scale. This random pattern is the indication of WOM occurring at regional sites including poor and good resource sites as well as various topographic and hydrological conditions [
64,
69,
70]. This random pattern of WOM rate may be due to extended periods of drought. For instance, [
44] reported that there was an extreme drought in which decline in white oaks increased from 6% (1999-2005) to 15% (2006-2010) across Ozark highlands of Arkansas and Missouri. Others found that the random pattern of WOM rate is also attributed to severe weather events such as storms, hurricanes, and droughts [
71,
72]. [
11] and [
73] documented that tropical winds followed by hurricanes killed randomly to many white oaks including other hardwood tree species and that may have resulted in a random pattern to WOM. Like the central and northern regions, our results reported that most of the southern region WOM rate had a significant uniform pattern on a broad scale. It is because the mortality in the mature stands of white oaks is not replacing themselves and would eventually generate a uniform pattern [
74]. Other reasons leading to a uniform pattern might be due to uniformity in drought or flooding and homogenous soil conditions in the region [
75,
76].
The observed clustered pattern of WOM rate in the central region raises intriguing queries concerning the underlying mechanisms affecting the dynamics of white oak mortality. Although the observed patterns can be explained by local scale causes, our investigation of site scale processes, especially self-thinning, offers a plausible explanation for the observed patterns. It is because previous studies have found that self-thinning, a natural process characterized by intraspecific competition leading to mortality, may accumulate over time and produce a clustered distribution pattern [
77]. For instance, oak forests across the central region of WOM rate (e.g., Illinois Ozark hills) had gone for massive competition with shade-tolerant species such as sugar maple, red maple, and American beech [
47,
79,
80]. In this condition, the mortality events may be driven by competition for resources within stands [
81]. The spatial clustering found in this region raises the possibility that these site-scale processes interact to affect WOM locally. However, our results also depicted a random pattern across the local scale of the central region. This random pattern may be due to environmental heterogeneity in the core area, which might result in a variety of microhabitats that have an impact on tree mortality rates [
82,
83]. For instance, disparities in soil types, moisture content, and other ecological parameters can lead to a distribution of WOM that appears random [
84,
85]. Others found that stochastic events such as disease outbreaks or drought can be the cause of random death patterns [
86,
87]. There is a greater chance for random mortality since drought followed by ice storms and other localized variables are impacting mortality in white oaks lacking a clear spatial pattern [
44]. The observed randomness at the local scale could represent temporal dynamics, in which short-term variations in environmental variables impact mortality events [
88,
89]. Over time, these variations can result in inconsistent geographic clustering. Likewise, the randomness may be facilitated by the edge effects in which the edges of white oaks stand in this region and might have faced distinct ecological conditions compared to the interior. It is because researchers documented that adjacent forest canopies had demonstrated to restrict light availability and mitigated tree growth on the outer margins of even-aged regeneration methods and gaps may exacerbate random mortality patterns [
90,
91,
92,
93,
94]. Our results also reported a uniform pattern in WOM rates at abroad scale that begs the question of what ecological factors are driving the dynamics of WOM. This unexpected uniformity could be caused by several factors. A uniform distribution of WOM rates may be facilitated by consistency in moisture content [
95], and drought-pathogen interactions [
96]. For instance, white oak stands throughout the large territory of the central region and has regular growth circumstances due to stable and consistent environmental conditions with an abundance of resources [
97]. And this stability in the environment and resources supports a uniform pattern for tree species. Similarly, the central region has a diverse ecosystem caused by human-induced variables like land use patterns such as private landownership across most of the states (e.g., Kentucky, Tennessee, Virginia, etc.). And the uniform land fragmentation resulting from landownership has contributed to a uniform distribution of WOM [
56,
98].
Our results from spatial patterns at both local and broad scale clustering in the northern region WOM rate indicate that several ecological processes and spatial factors are playing vital roles. Specific areas with comparable environmental circumstances, such as soil compaction, moisture levels, and species interactions, are examples of local scale clustering in WOM rate [
99,
100]. There is a greater chance that a specific soil type promotes the growth of a specific disease impacting white oaks' health. For instance, white oaks are impacted by exotic insects and diseases, and affect the abundance and distribution across the local scale [
101,
102]. Furthermore, given that the majority of white oaks stands were restored following destruction and the ensuing agricultural abandonment about a century ago, it is possible that site scale processes like self-thinning in white oak stands may accumulate throughout the region, which may result in a region wide WOM [
103,
104,
105]. Besides, elevation and distance from water have also an impact on the clustering observed at both local and broad scales. [
74,
106,
107] reported that environmental variability, such as elevation changes or proximity to bodies of water, had led to both local and broad-scale clustering. This is because places with comparable topographical features demonstrate clustering tendencies as a result of consistent environmental variables. There is also a greater chance that broad-scale clustering could be attributed to regional variables impacting the dynamics of WOM rates. Regional variables including climate fluctuations, historical disruptions, or landscape features that influence death rates at larger spatial scales could all be considered [
108,
109,
110]. We also found both random as well as uniform patterns across a broad scale. This could be due to various environmental stochastic processes. For instance, the northern region of the US is subjected to intermittent weather extremes, such as localized severe storms or extreme precipitation [
111,
112]. These stochastic processes result in random WOM patterns within specified broad scale ranges, reflecting the unpredictability of death events in such circumstances. A uniform WOM rate pattern at a broad scale might be the result of spatially homogeneous environmental factors. Factors include areas with constant soil types, climate, and land use methods such as private landownership, consistent logging, or land management techniques that may strongly exhibit uniform mortality rates [
56,
113,
114]. This homogeneity may result in similar WOM rates across large spatial extents. There are also edge effects and landscape characteristics that can contribute to both random and uniform patterns at broad scales. The reason may include locations near forest boundaries or regions with distinct topographical features [
115,
116]. These localized impacts may provide randomness within specific broad-scale ranges while contributing to homogeneity in neighboring places.
5. Conclusions
Our study showed various types of spatial patterns of WOM rate across the eastern US. These patterns are clustered, random, and uniform across southern, central, and northern region WOM rates with varying scales. Our research highlights mainly a clustered pattern at the local scale across each region and offers important new insights into the spatial patterns of WOM rate in these regions. The observed clustered pattern at the local scale highlights the complex interplay between the white oak population and the specific environmental circumstances across the eastern US. As stand-level competition, topographical and edaphic factors are identified as determinants that contribute to localized clustering of WOM rates. The fact that self-thinning is associated with site-scale processes highlights how complicated the mechanisms affecting white oak stands are in these regions. Such intricate ecological interactions emphasize the need to take finer-scale environmental elements into account when studying mortality events. This localized clustering highlights the necessity for site-specific solutions that consider the variability of topographical and soil conditions. However, there is also a clustered pattern at a broad scale across southern and northern region WOM rates suggesting a significant role of climate, elevation, and proximity to water bodies in these regions.
Our examination of the regional patterns of WOM rate found a distinct random pattern at broad scales both in the southern and northern regions. This random pattern suggests factors at broader scales such as climate, geology, and stochastic events may have impacted white oak stands. Similarly, climatic stressors such as water scarcity and environmental strain, have been identified as major factors in the random pattern of WOM events both at local and broad scale [
117].
Our study also unveiled a uniform pattern across all regions i.e., southern, central, and northern WOM rates. The reasons were associated with the presence of spatially homogeneous environmental conditions across a broad scale that may result in the uniform WOM pattern. Regions with consistent soil types, climate, and land use practices such as private landownerships and forest management techniques may suggest a uniform death rate across all regions. It is also evident that the land use practices, and stability of growth conditions emerge as significant factors of a uniform WOM distribution, giving valuable insights for forest management and conservation efforts in these regions exhibiting such patterns. However, further research is needed to investigate the specific mechanism underlying these relationships and to assess the long-term impacts on oak forest ecosystems. Future studies could explore the effects of multiple factors such as biotic and abiotic factors as well as land use practices, to obtain a more comprehensive understanding of the spatial pattern of WOM rate.