Ecological impacts of feral pigs in the Hawaiian Islands

The foraging habits of exotic ungulate species can directly and indirectly affect native plant and animal distribution and abundance patterns. Most of the studies on feral pig interactions with other biota in the Hawaiian Islands have been published as difficult to access reports to governmental and nongovernmental organizations, graduate student theses, and a few in peer reviewed journals. In this paper we discuss the origins of pig introductions to Hawaii, their feralization process, population expansion, and interactions with native and non-native biota. We also consider the environmental effects triggered by pigs on local ecosystems and biotic communities. Feral pig activities can reduce the abundance of native plant species, enhance conditions for the establishment of invasive non-indigenous plants, and perhaps indirectly negatively impact native forest bird species. Pig foraging and traveling patterns may also lead to physical alteration of ecosystems by increasing soil erosion that may lead to watershed degradation. However, much remains to be learned about the strength and significance of aforementioned interactions and their long-term effects on Hawaiian biota and ecosystems due to some confounding events. Elucidating the dynamics and long-term ecological effects generated by pigs is a crucial step towards increasing our understanding of and more effectively managing biotic interactions.     

Vegetation Recovery 16 Years after Feral Pig Removal from a Wet Hawaiian Forest

Nonnative ungulates can alter the structure and function of forest ecosystems. Feral pigs in particular pose a substantial threat to native plant communities throughout their global range. Hawaiian forests are exceptionally vulnerable to feral pig activity because native vegetation evolved in the absence of large mammalian herbivores. A common approach for conserving and restoring forests in Hawaii is fencing and removal of feral pigs. The extent of native plant community recovery and nonnative plant invasion following pig removal, however, is largely unknown. Our objective was to quantify changes in native and nonnative understory vegetation over a 16 yr period in adjacent fenced (pig‐free) vs. unfenced (pig‐present) Hawaiian montane wet forest. Native and nonnative understory vegetation responded strongly to feral pig removal. Density of native woody plants rooted in mineral soil increased sixfold in pig‐free sites over 16 yr, whereas establishment was almost exclusively restricted to epiphytes in pig‐present sites. Stem density of young tree ferns increased significantly (51.2%) in pig‐free, but not pig‐present sites. Herbaceous cover decreased over time in pig‐present sites (67.9%). In both treatments, number of species remained constant and native woody plant establishment was limited to commonly occurring species. The nonnative invasive shrub, Psidium cattleianum, responded positively to release from pig disturbance with a fivefold increase in density in pig‐free sites. These results suggest that while common native understory plants recover within 16 yr of pig removal, control of nonnative plants and outplanting of rarer native species are necessary components of sustainable conservation and restoration efforts in these forests.     

Utilization of Sugarcane Habitat by Feral Pig (Sus scrofa) in Northern Tropical Queensland: Evidence from the Stable Isotope Composition of Hair

Feral pigs (Sus scrofa) are an invasive species that disrupt ecosystem functioning throughout their introduced range. In tropical environments, feral pigs are associated with predation and displacement of endangered species, modification of habitat, and act as a vector for the spread of exotic vegetation and disease. Across many parts of their introduced range, the diet of feral pigs is poorly known. Although the remote location and difficult terrain of far north Queensland makes observing feral pig behavior difficult, feral pigs are perceived to seek refuge in World Heritage tropical rainforests and seasonally 'crop raid' into lowland sugarcane crops. Thus, identifying how feral pigs are using different components of the landscape is important to the design of management strategies. We used the stable isotope composition of captured feral pigs to determine the extent of rainforest and sugarcane habitat usage. Recently grown hair (basal hair) from feral pigs captured in remote rainforest indicated pigs met their dietary needs solely within this habitat. Stable carbon and nitrogen isotope values of basal hair from feral pigs captured near sugarcane plantations were more variable, with some individuals estimated to consume over 85% of their diet within a sugarcane habitat, while a few consumed as much as 90% of their diet from adjacent forested environments. We estimated whether feral pigs switch habitats by sequentially sampling δ(13)C and δ(15)N values of long tail hair from a subset of seven captured animals, and demonstrate that four of these individuals moved between habitats. Our results indicate that feral pigs utilize both sugarcane and forest habitats, and can switch between these resources.     

Invasive feral pigs impact native tree ferns and woody seedlings in Hawaiian forest

Invasive mammals can fundamentally alter native plant communities, especially on isolated islands where plants evolved without them. The globally invasive feral pig (Sus scrofa) can be particularly destructive to native plant communities. Tree ferns are an important understory component in many forests facilitating the establishment of a variety of species. However, the extent and effects of feral pig damage to tree ferns, and associated impacts on plant community regeneration, are largely unknown. We quantified the effect that feral pig damage has on tree fern growth, survival, and epiphytic woody seedling abundance over 1 year on 438 randomly selected tree ferns of three endemic species (Cibotium chamissoi, Cibotium glaucum, and Cibotium menziesii) in a Hawaiian montane wet forest with high tree fern and feral pig densities. Across all tree fern species, feral pigs damaged 13 % of individuals over 1 year. Compared with undamaged tree ferns, moderately- to heavily-damaged individuals had decreases of 4 to 27 % in trunk length increment and lost tenfold more fronds. Tree fern angle (standing, leaning, prone, or semi-prone) and woody seedling abundance co-varied with feral pig damage. Specifically, damaged tree ferns were more often prone or semi-prone and supported more seedlings, but also had annual mortality up to 34 % higher than undamaged tree ferns. Overall, feral pig damage had substantial negative effects on tree ferns by reducing growth and survival. Given the importance of tree ferns as regeneration sites for a variety of native plants, feral pig damage to tree ferns will likely alter future forest composition and structure. Specifically, feral pig damage to tree ferns reduces potential establishment sites for species that either regenerate preferentially as epiphytes or are currently restricted to epiphytic establishment due to ground rooting by feral pigs.     

Diet Overlap and Foraging Activity between Feral Pigs and Native Peccaries in the Pantanal

Inter-specific competition is considered one of the main selective pressures affecting species distribution and coexistence. Different species vary in the way they forage in order to minimize encounters with their competitors and with their predators. However, it is still poorly known whether and how native species change their foraging behavior in the presence of exotic species, particularly in South America. Here we compare diet overlap of fruits and foraging activity period of two sympatric native ungulates (the white-lipped peccary, Tayassu pecari, and the collared peccary, Pecari tajacu) with the invasive feral pig (Sus scrofa) in the Brazilian Pantanal. We found high diet overlap between white-lipped peccaries and feral pigs, but low overlap between collared peccaries and feral pigs. Furthermore, we found that feral pigs may influence the foraging period of both native peccaries, but in different ways. In the absence of feral pigs, collared peccary activity peaks in the early evening, possibly allowing them to avoid white-lipped peccary activity peaks, which occur in the morning. In the presence of feral pigs, collared peccaries forage mostly in early morning, while white-lipped peccaries forage throughout the day. Our results indicate that collared peccaries may avoid foraging at the same time as white-lipped peccaries. However, they forage during the same periods as feral pigs, with whom they have lower diet overlap. Our study highlights how an exotic species may alter interactions between native species by interfering in their foraging periods.     

Do aquatic macrophytes co-occur randomly? An analysis of null models in a tropical floodplain

One of the main issues in community ecology is the detection of structure and the identification of its related causes. In this study, co-occurrence null models were used to identify possible spatio-temporal patterns in the assemblage of aquatic macrophytes in the Upper Paraná River floodplain. The samples were obtained through the Long Term Ecological Research (LTER) Program at two different grains: (1) a coarser spatial grain in January and August 2001 (entire floodplain lagoons); (2) and a finer spatial grain in November 2006 (1 m² quadrats). The study was conducted in 36 lagoons, both connected and disconnected to the main river channel, located in the sub-basins of the Baía, Ivinheima and Paraná rivers. Two null models of species co-occurrence, the C-Score and Checkerboard indices, were used to test the null hypothesis of random structure of the aquatic macrophyte assemblages. The null models showed that the aquatic macrophyte assemblages were spatially structured in the distinct spatial grains. However, despite this general pattern, macrophyte assemblages are organized differently depending on the degree of connectivity, seasonal period and, at a finer grain, depth. Species co-occurrences were random in the disconnected lagoons during flood periods, in deep zones of the lagoons of the Baía River and in the shallow littoral zone in the lagoons of the Paraná River. Analysis of the patterns of co-occurrence indicated that competition and/or habitat preferences are probably important influences on the nonrandom structure of assemblages. However, we suppose that at least three important factors (disturbances by water level fluctuation, dispersion and facilitation) counteract potential effects of competition in specific situations, leading macrophyte assemblages to assume random structure.     

Response of floodplain grassland plant communities to altered water regimes

Floodplain grasslands are often composed of a mosaic of plant communities controlled by hydrological regime. This article examines the sensitivity of floodplain grassland plant communities to water regime using reciprocal transplantation of an inundation grassland and a flood-meadow within an English floodplain. Experimental treatments comprised control, transplanted and lifted plots; the last treatment, in order to elucidate any disturbance effects of transplantation. Plant community response was analysed using species abundance and their ecological traits. Results from both communities showed substantial annual variations related to hydrology, including significant species changes, but generally, vegetation seemed to be responding to drier conditions following a major flood event. This ‘drying’ trend was characterised by increased species diversity, a greater abundance of competitive species and fewer typical wetland plants. Transplanted community composition increasingly resembled receptor sites and transplant effects were most pronounced the first year after treatment for both vegetation types. Differential responses to water regime were detected for the two plant communities. The inundation grassland community was particularly dynamic with a composition that rapidly reflected drying conditions following the major flood, but transplantation into a drier flood-meadow site prompted little additional change. The flood-meadow community appeared more resistant to post-inundation drying, but was sensitive to increased wetness caused by transplantation into inundation grassland, which significantly reduced six species while none were significantly favoured. The effects of disturbance caused by lifting the transplants were limited in both communities, although five species showed significant annual fluctuations. The study shows that small alterations in water regime can prompt rapid vegetation changes and significant plant species responses in floodplain grasslands, with effects probably magnified through competitive interactions. The dynamic properties of floodplain vegetation demonstrated by this study suggest that its classification, management and monitoring are challenging and ideally should be based on long-term studies.     

Waterbird herbivory on a newly created wetland complex: potential implications for site management and habitat creation

Estimations of waterbird herbivory at a newly created wetland, indicate that overwintering waterbirds have the potential to remove a large proportion of the above-ground macrophyte standing crop in the main lagoons. Observed lake state switching and changes in macrophyte species composition may have been linked, in part, to waterbird herbivory. Significant differences were observed in the distribution of certain species of waterbird between the lagoons, which may reflect food preferences. Between 30% and 50% of the herbivorous waterbird community utilised alternative on-site feeding areas over the winter period thus reducing the impact of grazing intensity in the larger open water areas. These results emphasise the advantages of incorporation of site heterogeneity, specifically distinct hydrological units, within the design of wetland creation schemes. Hypereutrophic conditions, with turbid water and dense filamentous algal blooms were experienced on the Reservoir Lagoon yet the lagoon supported high densities of overwintering birds (> 100 individuals ha^–1. This led to an alleviation of grazing pressure on other water bodies thereby promoting the development of macrophyte communities elsewhere on site. The need for work to provide a better understanding of critical thresholds of herbivory in determining community composition and abundance, and particularly the role of eutrophication in these processes is highlighted.     

Vegetation response to removal of non-native feral pigs from Hawaiian tropical montane wet forest

Globally, non-native ungulates threaten native biodiversity, alter biotic and abiotic factors regulating ecological processes, and incur significant economic costs via herbivory, rooting, and trampling. Removal of non-native ungulates is an increasingly common and crucial first step in conserving and restoring native forests. However, removal is often controversial and there is currently little information on plant community responses to this management action. Here, we examine the response of native and non-native understory vegetation in paired sites inside and outside of exclosures across a 6.5–18.5 year chronosequence of feral pig (Sus scrofa) removal from canopy-intact Hawaiian tropical montane wet forest. Stem density and cover of native plants, species richness of ground-rooted native woody plants, and abundance of native plants of conservation interest were all significantly higher where feral pigs had been removed. Similarly, the area of exposed soil was substantially lower and cover of litter and bryophytes was greater with feral pig removal. Spatial patterns of recruitment were also strongly affected. Whereas epiphytic establishment was similar between treatments, the density of ground-rooted woody plants was four times higher with feral pig removal. Abundance of invasive non-native plants also increased at sites where they had established prior to feral pig removal. We found no patterns in any of the measured variables with time, suggesting that commonly occurring species recover within 6.5 years of feral pig removal. Recovery of species of conservation interest, however, was highly site specific and limited to areas that possessed remnant populations at the time of removal, indicating that some species take much longer (>18.5 years) to recover. Feral pig removal is the first and most crucial step for conservation of native forests in this area, but subsequent management should also include control of non-native invasive plants and outplanting native species of conservation interest that fail to recruit naturally.     

Species richness in tropical fresh waters of Australia

General ecological expectations about the relationship between latitude and species richness are that at low latitudes (the tropics) species richness is greater than at higher latitudes (temperate and polar regions). Recent work suggests that this may not be the case for several habitat types and biological groups in Australia. Results are conflicting: on present evidence (admittedly sparse) it appears that in Australian tropical fresh waters species richness is generally depressed in zooplankton and littoral microfaunal communities, but not in macroinvertebrate communities in typical streams and in fish communities (and perhaps also in amphibian and reptile communities). The situation is indeterminate for tropical phytoplankton and macrophyte communities.     

No sitting on the fence: protecting wetlands from feral pig damage by exclusion fences requires effective fence maintenance

Feral pigs damage the significant ecological and cultural values of tropical Australian wetlands. Control measures such as culling, baiting, and trapping can reduce overall pig populations, but do not eliminate the substantial physical damage to wetlands that can occur from just a few individuals. Exclusion fences have been adopted as a potential technique to prevent damage to selected wetlands. To test the effectiveness of exclusion fences we measured the physical damage caused by pigs to multiple wetlands in the Archer River catchment of tropical northern Australia. Wetlands were fenced using a typical cattle exclusion fence, a specific pig exclusion fence or had no fence. Initial analyses of these fence treatments showed no significant difference in the intensity of physical pig damage to exposed wetland sediments and fringing vegetation. However, several of the pig exclusion fences were found to have been compromised. Reanalysis indicated wetlands with functioning pig exclusion fences had no physical pig damage and this was significantly less damage than in all other treatments. In contrast, wetlands with compromised pig exclusion fences had damage that was statistically equivalent to sites without fences or with cattle exclusion fences, but in individual cases had the worst damage recorded in any of the treatments. Compromised pig exclusion fencing of wetlands can thus be worse than having no fencing at all. This suggests that the successful prevention of pig damage to wetlands by exclusion fences requires ongoing and effective fence monitoring and maintenance regimes.

     

Prevalence of Tree Regeneration by Sprouting and Seeding Along a Rainfall Gradient in Hawai'i

Drought stress in tropical dry forests is thought to result in greater asexual regeneration via vegetative sprouting ( e.g ., basal, root, and branch layering) than occurs in moister tropical forests. We tested this hypothesis by examining the prevalence of tree sprouting and seeding in tropical forests located along a rainfall gradient on the island of Hawai'i. Additionally, we examined the potential for novel disturbance, feral pig Sus scrofa rooting and trampling, to alter patterns in tree regeneration mode. We found greater sprouting (in terms of relative density and basal area) in dry forests than in mesic and wet forests, supporting the hypothesis. We also found that feral pig disturbance is negatively correlated with the relative density and basal area of seedlings in wet forests, but is positively correlated with the relative importance of sprouting, and the richness and diversity of sprouting species. Our results suggest rainfall regimes may be an important factor controlling broad-scale patterns in tree regeneration mode, and that exotic ungulates can significantly modify such patterns with potential consequences for the structure and dynamics of tree populations and communities.     

Impact of nonnative feral pig removal on soil structure and nutrient availability in Hawaiian tropical montane wet forests

Conservation and restoration of ecosystems impacted by nonnative ungulates increasingly involves their removal and exclusion. While the influence of nonnative ungulate removal on plant communities is commonly monitored, impacts on underlying ecological processes are seldom quantified. Here we examined how nonnative feral pig (Sus scrofa) removal from Hawaiian tropical montane wet forests affects soil physical and chemical properties. Unique to this study, measurements were taken in paired sites inside and outside of five feral pig removal units representing a ~20 year, highly constrained chronosequence where other potentially confounding variables are held constant. Additional targeted measurements were taken inside and outside of a single exclosure in areas characterized by ‘low’ versus ‘high’ feral pig activity. Overall, nonnative feral pig removal increased stable soil aggregates and porosity, and decreased bulk density, water-filled pore space, and soil moisture content. Further, feral pig removal increased soil nutrient regeneration as evidenced by increased extractable cations, increased resin available NO₃ ^? and total inorganic N, and enriched foliar δ¹⁵N. Increasing time since feral pig removal was positively related to net nitrification and total net inorganic N mineralization, and negatively related to pH and net ammonification. Results from both the chronosequence and targeted sampling were consistent in direction and support a central role of feral pig removal in modifying soil physical and chemical properties. Changes in soil properties following ungulate removal coincided with large increases in understory vegetation cover, highlighting the need to better understand aboveground-belowground linkages following nonnative ungulate removal.     

The effect of seasonal variation in abiotic factors on fish community structure in temporary and permanent pools in a tropical wetland

1. Fish community structure depends on biotic interactions and abiotic variables. Abiotic variables appear to gain importance in highly variable freshwater systems, such as tropical wetlands where a marked seasonal hydroperiod (dry and wet seasons) modifies water quality and quantity, differentially affecting fish survival and, consequently, modifying species richness and abundance. 2. We evaluated the relationship between abiotic variables and fish community structure in variable (temporary) and stable (permanent) pools that were interconnected in a tropical wetland with marked annual dry and wet seasons. 3. All fish species were able to occupy any of the studied pools, but our results showed distinctive fish community structures in permanent and temporary pools. Community structure was related to temperature, depth, pH and macrophyte coverage. Total fish abundance in the wetland was negatively related to water depth and positively related to macrophyte coverage. 4. Null models of co‐occurrence indicated a non‐random pattern at the wetland scale and a random pattern within groups of pools with similar characteristics, suggesting that fish communities are structured according to habitat features. We conclude that seasonal abiotic variation and habitat characteristics in this highly variable pristine wetland play major roles in structuring fish communities.     

Carbon sequestration in freshwater wetlands in Costa Rica and Botswana

Tropical wetlands are typically productive ecosystems that can introduce large amounts of carbon into the soil. However, high temperatures and seasonal water availability can hinder the ability of wetland soils to sequester carbon efficiently. We determined the carbon sequestration rate of 12 wetland communities in four different tropical wetlands—an isolated depressional wetland in a rainforest, and a slow flowing rainforest swamp, a riverine flow-through wetland with a marked wet and dry season, a seasonal floodplain of an inland delta—with the intention of finding conditions that favor soil carbon accumulation in tropical wetlands. Triplicate soil cores were extracted in these communities and analyzed for total carbon content to determine the wetland soil carbon pool. We found that the humid tropic wetlands had greater carbon content (P ≤ 0.05) than the tropical dry ones (96.5 and 34.8 g C kg^?1, respectively). While the dry tropic wetlands had similar sequestration rates (63 ± 10 g Cm^?2 y^?1 on average), the humid tropic ones differed significantly (P < 0.001), with high rates in a slow-flowing slough (306 ± 77 g Cm^?2 y^?1) and low rates in a tropical rain forest depressional wetland (84 ± 23 g Cm^?2 y^?1). The carbon accumulating in all of these wetlands was mostly organic (92–100%). These results suggest the importance of differentiating between types of wetland communities and their hydrology when estimating overall rates at which tropical wetlands sequester carbon, and the need to include tropical wetland carbon sequestration in global carbon budgets.     

ILLEGAL TRANSLOCATION AND GENETIC STRUCTURE OF FERAL PIGS IN WESTERN AUSTRALIA

Abstract: Unlike many regions in the world where wild pigs (Sus scrofa) are threatened, in Australia they are a significant invasive species. As such, the molecular ecology of feral pigs was investigated to understand their social and population genetic structure. Samples from 269 adult animals were collected over their distribution in southwestern Australia. Using 14 highly polymorphic microsatellite markers, we identified 7 inferred feral pig populations that had moderate heterozygosity (mean = 0.580) and displayed a high level of differentiation (mean R_ST = 0.180). In revealing the genetic structure of feral pigs, we detected anomalies in the putative origin of some individuals. Samples from these animals were collected from 2 main areas: recently colonized regions that were previously uninfested, and established feral pig populations, where animals from geographically isolated areas had been introduced. In the latter, these corresponded to areas that were in close proximity to public road access and towns. Given the large distances immigrants were found from their population of origin (from 50 to >400 km), the generally low levels of dispersal of southwest feral pigs, and the grouping and sex of these pigs, we suggest that these individuals have been deliberately and illegally translocated to supplement recreational hunting stocks. Additionally, we could not detect any genetic contribution in these feral pigs from domestic pig herds, suggesting that the deliberate release of domestic pigs to restock feral populations is relatively uncommon. Our molecular data allowed some inferences regarding the success or lack thereof of current management practices, and offered considerable insights into the dynamics of the feral pig populations and identification of “new” approaches that may allow for better control of this highly destructive species.     

Fish diet composition in floodplain lagoons of an Australian dryland river in relation to an extended dry period following flooding

Floodplain rivers worldwide are threatened by loss of connectivity to their floodplains and hence reduced benefits from floodplain energy subsidies. Dryland rivers with ‘boom and bust’ ecological responses to flooding and extended dry periods may be particularly vulnerable. This paper describes variations in dietary composition of three fish species of contrasting trophic position in dryland floodplain lagoons with variable flood inundation and drying histories. The study species were Ambassis agassizii - a microphagic carnivore, Leiopotherapon unicolor - a carnivore/omnivore, and Nematalosa erebi - an algivore/detritivor. Despite the range of food items recorded in fish guts, each species fed mostly on relatively few food categories and few food items within each category. Most of the spatial (i.e. among lagoons) and temporal dietary variation was associated with different proportional contributions of these food items. Given the absence or low magnitude of flooding during the study period, temporal changes in diets of the three species are probably the result of successional changes in composition of invertebrate prey as the dry season progressed. The focus of each fish species on relatively few food categories and a few reliable food items within each category may be the most profitable foraging strategy when food resources are limiting in progressively drying floodplain lagoons.     

Eradication of Feral Pigs From Pinnacles National Monument

Abstract: Feral pigs (Sus scrofa) have caused considerable damage where they have been introduced around the world. At Pinnacles National Monument, California, USA, managers were concerned that feral pigs were damaging wetland habitats, reducing oak regeneration, competing with native wildlife, and dispersing nonnative plant species through soil disturbance. To address these threats the National Park Service constructed an exclosure around 57 km² of monument land and through cooperation with the Institute for Wildlife Studies eradicated all feral pigs within the area. Trapping, ground-hunting, hunting dogs, and Judas techniques were used to remove feral pigs. Trapping techniques removed most pigs, but a combination of techniques was required to cause eradication. A series of bait sites and transects across the monument helped focus removal efforts and facilitated detection of the last remaining feral pigs in the exclosure. Consistent funding and cooperation from the National Park Service allowed for a seamless and comprehensive program that provided intensive removal of feral pigs. The successful eradication of feral pigs at Pinnacles National Monument should encourage managers in other areas to implement future control or eradication programs.     

Will fencing floodplain and riverine wetlands from feral pig damage conserve fish community values?

Installation of feral pig (Sus scrofa) exclusion fences to conserve and rehabilitate coastal floodplain habitat for fish production and water quality services remains untested. Twenty‐one floodplain and riverine wetlands in the Archer River catchment (north Queensland) were surveyed during postwet (June–August) and late‐dry season (November–December) in 2016, 2017, and 2018, using a fyke net soaked overnight (~14–15 hr) to test: (a) whether the fish assemblage are similar in wetlands with and without fences; and (b) whether specific environmental conditions influence fish composition between fenced and unfenced wetlands. A total of 6,353 fish representing twenty‐six species from 15 families were captured. There were no wetland differences in fish assemblages across seasons, years and for fenced and unfenced (PERMANOVA, Pseudo‐F < 0.589, p < .84). Interestingly, the late‐dry season fish were far smaller compared to postwet season fish: a strategy presumably in place to maximize rapid disposal following rain and floodplain connectivity. In each wetland, a calibrated Hydrolab was deployed (between 2 and4 days, with 20 min logging) in the epilimnion (0.2 m) and revealed distinct diel water quality cycling of temperature, dissolved oxygen and pH (conductivity represented freshwater wetlands), which was more obvious in the late‐dry season survey because of extreme summer conditions. Water quality varied among wetlands in terms of the daily amplitude and extent of daily photosynthesis recovery, which highlights the need to consider local conditions and that applying general assumptions around water quality conditions for these types of wetlands is problematic for managers. Though many fish access wetlands during wet season connection, the seasonal effect of reduced water level conditions seems more overimprovised when compared to whether fences are installed, as all wetlands supported few, juvenile, or no fish species because they had dried completely regardless of the presence of fences.