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Article

High-Severity Wildfires Alter Ant (Hymenoptera: Formicidae) Foraging Assemblage Structure in Montane Coniferous Forests and Grasslands in the Jemez Mountains, New Mexico, USA

by
Jonathan Knudsen
1,
Robert Parmenter
2,
Theodore Sumnicht
3 and
Robin Verble
3,*
1
Department of Natural Resources Management, Texas Tech University, Lubbock, TX79401, USA
2
National Park Service, Jemez Springs, NM 87025, USA
3
Department of Biological Sciences, Missouri University of Science and Technology, Rolla, MO 65409, USA
*
Author to whom correspondence should be addressed.
Conservation 2024, 4(4), 830-846; https://doi.org/10.3390/conservation4040049
Submission received: 5 November 2024 / Revised: 2 December 2024 / Accepted: 5 December 2024 / Published: 9 December 2024
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Abstract

:
High-severity wildfires create heterogeneous patterns of vegetation across burned landscapes. While these spatial patterns are well-documented, less is known about the short- and long-term effects of large-scale high-severity wildfires on insect community assemblages and dynamics. Ants are bottom-up indicators of ecosystem health and function that are sensitive to disturbance and fill a variety of roles in their ecosystems, including altering soil chemistry, dispersing seeds, and serving as a key food resource for many species, including the federally endangered Jemez Mountain salamander (Plethodon neomexicanus). We examined the post-fire effects of the 2011 Las Conchas Wildfire on ant communities in the Valles Caldera National Preserve (Sandoval County, New Mexico, USA). We collected ants via pitfall traps in replicated burned and unburned sites across three habitats: ponderosa pine forests, mixed-conifer forests, and montane grassland. We analyzed trends in species richness, abundance, recruitment, loss, turnover, and composition over five sequential years of post-fire succession (2011–2015). Ant foraging assemblage was influenced by burn presence, season of sampling, and macrohabitat. We also found strong seasonal trends and decreases over time since fire in ant species richness and ant abundance. However, habitat and seasonal effects may be a stronger predictor of ant species richness than the presence of fire or post-fire successional patterns.

1. Introduction

Fire is an ecologically important natural disturbance [1,2]. Historically, fires burned at higher frequencies than today across much of the United States [1,2,3], and consequently, many ecosystems (e.g., lodgepole pine, tallgrass prairie, longleaf pine) are reliant on the changes and benefits associated with frequent fires and their effects [4,5]. Beginning in the late 1800s, fires were suppressed by humans across most of their historic North American range [1], resulting in altered habitat structure and increased understory growth [6]. Forest canopies became denser [7], and litter, duff, and coarse woody debris increased in many ecosystems [6,8]. Compounding the exclusion of fire in fire-dependent ecosystems are droughts and climatic unpredictability associated with a warming planet [9]. Thus, at present, when wildland fires do occur, they often occur at higher intensities and severities than historically observed [10,11].
The life history and habitat of arthropods influence their responses to wildland fire [5,12,13,14,15]. Many ground- and soil-dwelling arthropods experience limited impacts of wildfires as they burrow into heat-insulated soils [12]. Other species (e.g., surface-active predators, flying insects) may exploit burned areas and rapidly colonize these habitats [13,16]. In addition, other taxa may remain locally abundant but shift in composition or occupancy of a habitat. For example, spiders (Araneae) are more abundant in burned habitats [5], and ground beetle (Carabidae) communities changed gradually from forest species to open ground species in response to fire and clear-cutting. [15]. Fires often homogenize local habitats, and the destruction of microhabitats and changes in microclimate can reduce the numbers of specialist species and may favor more generalist groups [14].
Ants (Hymenoptera: Formicidae) are a diverse and abundant group of arthropods that form an important foundation in many ecosystems and provide valuable ecosystem services [17,18,19,20,21,22,23,24,25,26]. These include breaking down and scavenging dead organisms [26], predating upon other arthropods including pest species [18,22,27], increasing soil nutrients [20], providing protection from herbivorous arthropods [21,25], and dispersing seeds [24]. They also serve as an important food source for many threatened and endangered species [17,19,28].
Ants are affected by multiple types and severities of disturbances, and their sensitivity to wildland fire is well-documented. Ant responses to fire are often affected by changes in habitat structure and microclimate and less influenced by direct fire effects (e.g., heat, smoke; [2,3,14,29,30]). However, clear patterns in post-fire ant communities are not always evident. The overall abundance, diversity, and activity level of ant communities can decrease [23,31] or increase [2,30,32] post-fire. For example, harvester ants in the Florida sandhills increased their foraging area and numbers of ants leaving the colony in burned areas [33], and ant communities are more diverse in recently burned woodland habitats of the Gibson Desert Nature Reserve [34]. Verble and Stephen [35] found delayed decreases in occurrence of black carpenter ants (Camponotus pennsylvanicus) in Ozark forests, and Wilkinson et al. [36] found a temporary increase in overall ant abundance post fire. Additionally, effects may linger: Ant communities may take years to recover and reach their pre-fire composition [37,38]. Relatively few studies have examined the effects of mixed-severity wildfire on ground-foraging arthropods, mostly due to the logistical difficulties of obtaining pre- and post-fire data.
Our objectives were to describe the effects of wildfire occurrence, macrohabitat, and season on ant functional groups, species richness, abundance, recruitment, loss, turnover, and composition in the Valles Caldera National Preserve in the Jemez Mountains of northern New Mexico, USA. We hypothesized that macrohabitat, season, and time since fire would influence ant foraging assemblage. We predicted that burned forests would experience more pronounced drops in ant abundance and species richness than grassland habitats, and we predicted that fire would simplify functional groups [2,3,14,29,30,32,34,39]. Additionally, we predicted that vegetation type would influence ant foraging assemblage, with grasslands having a higher species richness and abundance [2,3,4,13,40] than forested areas.

2. Materials and Methods

2.1. Study Site

We conducted this study in the National Park Service’s Valles Caldera National Preserve (VCNP), Sandoval County, NM, USA. Valles Caldera is a dormant volcanic caldera, 20 km wide and located in the Jemez Mountains [41] in north-central New Mexico. The site is composed of numerous forested volcanic domes with vast grassy valleys (valles) between them. Sites range in elevation from 2349 to 3439 m above sea level (asl). The VCNP is composed of two fundamental habitat components, mainly distributed by elevation gradient—forests and woodlands in the mountains, and wetlands and grasslands in the valleys. The forest habitats are further delineated by elevation; spruce–fir forests in the highest elevations (>3050 m asl), mixed-conifer forests in the mid-elevations (2740–3050 m asl), and ponderosa at the lowest elevations (<2740 m asl). The valleys are composed of montane wet meadows and montane valley grasslands. Montane valley grasslands are the dominant grassland habitat type and are composed of upland grasses [42], while montane wet meadows are primarily composed of facultative or obligate graminoids, mostly sedges and rushes [42].
We examined areas of the VCNP that burned during the Las Conchas Fire (2011). The Las Conchas Wildfire burned 63,131 hectares (631 sq km) of the VCNP from 26 June to 3 August 2011 [43]. The fire was started by a tree falling over a powerline on adjacent private land, and the initial spread was rapid, burning over 17,800 ha in the first 24 h post-ignition due to wind gusts that exceeded 65 km per hour (kph). The fire burned at a mixed severity; areas that had previously burned experienced a lower severity and higher resilience to the fire [44]. Approximately 21% of the burn area was high-severity fire, 34% moderate-severity, and 45% low-severity [45].

2.2. Field Methods

We established study sites during the first week of July 2011, immediately after the fire front had passed through various areas of the VCNP. We selected three habitat types for assessment of vegetation and ant foraging assemblages: open grasslands (valles), ponderosa pine forests, and mixed-conifer forests (Figure 1). Grassland sites within the fire perimeter were completely burned, and all forest sites sustained high-severity, stand-replacement fire. We identified six burned sites in each of the three habitats, and then we selected six unburned control sites for each habitat based on similarities in grassland community type and forest stand characteristics, elevation, and aspect, for a total of 36 study sites. Within each study site, we installed replicate permanent vegetation and ant sampling locations, which were then sampled during the remainder of 2011 through autumn 2015. In 2013, three of our control sites were burned in a separate fire (Thompson Ridge Fire) and were excluded from further analysis (N = 6 control sites from 2011 to 2012, N = 3 control sites from 2013 to 2018). Vegetation sampling methods can be found in [46].

2.3. Ant Sampling

We sampled ant assemblages using pitfall traps, which capture foraging-surface active workers. Traps consisted of three plastic cups (9 cm diameter × 12.5 cm depth) placed in triangle pattern approximately 1 m apart. Cups were inserted flush with the ground surface and covered with a ceramic tile (elevated ~3 cm on nails) to prevent rain and debris from entering the traps and filled to ~50% full with propylene glycol. Trap arrays were surrounded by metal fences to minimize disturbance by elk and bears (Figure 1B,F). Traps were left open continuously, and we sampled the traps every three weeks from mid-July to November 2011 and from May to November 2012–2015. Field samples were collected in glass jars with 70% ethanol and transported to the laboratory for analysis.

2.4. Laboratory Methods

Pitfall trap contents were sorted to family by VCNP staff. Ant specimens were further sorted to species using dissecting microscopes and identified using established taxonomic keys [47,48,49,50,51,52,53,54,55,56,57]. Voucher specimens were pointed, labeled, and pinned for archiving. Problematic specimens were verified by taxonomists. Voucher specimens are housed at Missouri University of Science and Technology, Rolla, MO, USA.

2.5. Data Analysis

We modeled the effects of fire presence (burned, unburned), season, year, and macrohabitat on ant abundance and ant species richness using a fully factorial analysis of variance controlled for multiple comparisons (ANOVA, alpha = 0.05, R Core Team 2013) and used Tukey HSD tests to examine differences among groups (alpha = 0.05). We examined assumptions of normality with a Shapiro–Wilk test. We categorized functional groups via AntWiki [58], examining the abundance of ants within habitat associations, nest types, diet, species associations, and social parasitism by burn treatment (X). We used linear regression to examine relationships between parasitic ants and known host ants. All data were checked for normality (Shapiro–Wilk, p < 0.05) and log-transformed when necessary to correct variance heterogeneity
We measured species turnover as the number of species disappearing and appearing between each sampling year as a percentage of the total number of species observed in the sample year [51]. We also plotted changes in abundance of select taxa over time to visualize trends with time. For this, we plotted data for all species with >5 individuals collected throughout the entire sampling period, and all genera with >10 individuals collected throughout the sampling period.
We categorized the seasons per Goforth [59] and Stahle et al. [60]. Spring was defined as 1 May–30 June, summer was defined as 1 July–31 August, and fall was defined as 1 September–30 November. This seasonal categorization follows rainfall patterns, with spring predicted to be a warm dry season, summer predicted to be a warm rainy season, and fall predicted to be cool season with intermittent precipitation. Data analyses were conducted in the SAS statistical package [61].

3. Results

We collected 66,203 ants from 2011 to 2015. Sixty species representing sixteen genera were identified (Table 1).

3.1. Ant Species Richness

Species richness varied among habitats (p < 0.0001, df = 2, Table 2, Figure 2) declined annually from 2011 to 2015 (Figure 3C). Montane valley grasslands supported the most species, while the forested ponderosa pine sites and mixed-conifer sites had fewer species and were not statistically different from one another (Figure 2B). Burned sites had a significantly higher ant species richness than unburned control sites across all three habitat types. Interactions between habitat and burn treatment influenced ant species richness (Table 2). We found strong seasonal trends in species richness within habitats (p < 0.0001, df = 26; Table 2, Figure 2A). All sites followed the same pattern of high species richness during the spring and summer seasons, with a significant drop in species richness in the fall.

3.2. Ant Abundance

Ant abundance was significantly different among the three habitats (p < 0.0001, df = 2, Table 3, Figure 2A). Montane valley grasslands had the highest mean ant abundance and were significantly different from the two forested sites. Ponderosa pine and mixed-conifer sites did not have significantly different numbers of ants. Ant abundance was not significantly different among burn severities (p = 0.1414, df = 4). Interactions between burn treatment and habitat were also not significant (p = 0.4159, df = 2). Moreover, ant abundance varied by season (Table 3, Figure 2A). In general, abundance was highest in spring and summer across most years and lowest in fall. Trends in ant abundance varied among habitats over time (Figure 3A). All three sites followed similar patterns in abundance over time, with peak abundance occurring during the spring sampling period (Figure 2A).

3.3. Turnover Rates

Rates of ant species turnover between years varied from 35.5% to 48.6% (Table 4); however, total change in species richness resulting from this turnover trended downward annually, starting from a high of 53 in 2011 and falling to 25 species in 2015. Species richness declines were greater in unburned sites than burned sites, and rates of turnover were also higher in unburned sites (Table 4).

3.4. Ant Functional Groups

Host ant and dulotic ant abundances correlated positively and weakly with one another (Figure 4). Xenobionts and host ants were influenced by burning and seasonal patterns; however, dulotic ants were only influenced by habitat (Table 5, Figure 5C).

3.5. Ant Nest Types

All ant nesting groups were influenced by burning, except wood nesting ants, which were only influenced by sites and seasons (independently, Table 6, Figure 5B). Those ants that build nests in soil with mounds near the entrance were more abundant in burned montane conifer (M = 21.71, SE = 4.930) than unburned montane conifer (M = 8.73, SE = 6.194), and were more abundant in burned ponderosa pine (M = 19.92, SE = 5.257) than unburned ponderosa pine (M = 11.20, SE = 5.937). However, in montane valleys, they were less abundant in burned sites (M = 42.11, SE = 4.648) than unburned sites (M = 69.51, SE = 4.666). Thatch nest builders were rare across all habitat types, but they were more abundant in burned sites than unburned (p = 0.0010). Ants that nested in tree holes and spaces between stones were more abundant in burned montane valleys (M = 8.65, SE = 0.092) than unburned montane valley habitats (M = 3.82, SE = 0.906), and there were no differences between burned and unburned PP and MC. Twig nesters and leaf litter nesters were less abundant in burned MC (burned: M = 9.20, SE = 4.293; unburned M = 2.70, SE = 5.394) and PP (burned PP: M = 9.08, SE = 4.570; unburned PP: M = 2.77, SE = 5.170), but they had no differences between burned and unburned MV. Finally, the ants that steal nests or nest within the nests of other species were generally rare, but they had a higher abundance in burned sites in MV (burned MV: M = 8.65, SE = 0.928; unburned MV: M = 3.83, SE = 0.931).

3.6. Ant Diet

All ant diet types were influenced by burning; however, omnivorous and actively predaceous ants showed site-specific relationships with burning (Table 7, Figure 5B). The way in which fire impacted a functional group varied widely (Figure 5B). For example, active predators were less abundant on burned montane valley sites (M = 13.54, SE = 3.180) than unburned montane valley sites (M = 27.79, SE = 3.197), but they were more abundant on burned ponderosa pine sites (M = 13.17, SE = 3.602) than unburned ponderosa pine sites (M = 5.43, SE = 4.06). Burned montane valley sites had more non-predaceous omnivorous ants (M = 38.23, SE = 2.960) than unburned montane valley sites (M = 22.58, SE = 2.970), and there were no differences in non-predaceous omnivores between burned and unburned ponderosa pine and montane conifer sites. Honeydew-tending ants showed no differences between burned and unburned MV and MC sites, but they had a higher abundance on burned ponderosa pine sites (burned M = 18.21, SE = 3.303; unburned M = 8.22, SE = 3.729). Finally, dulotic ants showed no differences between burned and unburned MC and PP, but they had a higher abundance on burned MV (M = 13.18, SE = 1.640) than unburned MV sites (M = 1.82, SE = 1.65).

4. Discussion

The results of this study indicate that ant assemblages in the Valles Caldera National Preserve are diverse and resilient to the immediate effects of wildfire. We found a decreasing ant abundance and ant species richness with the time since burning. The effects of wildland fire on species richness and ant abundance in the VCNP are related to the time since the fire, site, and fire occurrence. Ant species composition is driven in part by the burn effect and habitat variables.
Our study and most previous studies examined the ant community structure utilizing burned versus unburned sites. These studies have occasionally found no difference in species richness or ant abundance between burned and unburned sites [62,63]; however, most studies have found decreases [12,31,38,39] or increases [2,30,32] in ant species richness and abundance. Our study aligns with the studies that have found changes in the leaf litter arthropod community structure as a result of wildland fire, and it extends these results further to show that the time since a fire also results in differences in ant foraging and nesting assemblages. Interestingly, wood nesting ants increased in abundance in burned areas, suggesting that downed deadwood availability may have increased post-burn, even after the consumption of existing wood resources during the fire.
Studies of fire intensity (e.g., [39,64]) show markedly different patterns of short-term ant species richness and abundance to in our study. These studies occurred in two distinct fire-adapted ecosystems, so geography, fire history, and local factors may drive these differences. Nonetheless, further examinations of how fire intensity and burn severity influence insect community structure are warranted [65,66].
Ant species richness and abundance peaked immediately post-fire for all (including unburned) sites. While all habitats then experienced declines in ant species richness and abundance, these declines were less precipitous in unburned habitats, suggesting that inhospitable conditions associated with the year (rainfall, high temperatures, increased predator abundance, or some combination of these factors) may be differentially impacting burned habitats, with the burned habitats yielding more favorable conditions. This result was unexpected and may be a result of local climatic factors (e.g., precipitation, temperature) also differentially impacting burned habitats via increased solar radiation [67] and water-induced erosion [68]. Changes in ant abundance over time corresponded with changes in herbaceous vegetation from 2011 to 2015 and may have been a major factor in the observed reduction in ant species and abundance over time. The increase in plant density (grasses, forbs) would enhance the “architectural complexity” of the microhabitat on the ground, which would slow down ant movements and reduce the likelihood of ants being collected in pitfall traps, resulting in fewer ants and fewer species (Figure 3). The composition and structure of ant assemblages was not homogenous among habitats or the time since the fire, suggesting that the observed assemblages are mostly driven by shifts in habitat and environment and not by short-term behavioral changes. This could also be a function of the elimination of litter (needles, branches, etc.), which in years 2–5 would have simplified the ground “architecture” and allowed ants to move more freely and rapidly, thereby increasing trap captures. Immediately after the fire, in 2011, there was still a deep layer of ash, so ant movements should have been somewhat constrained. But after the first monsoon storms in late July, most of the ash was washed away or matted down to form a relatively hard surface, making ant movement easier. Thus, the observed increased numbers of ants and species in the burned habitats may be an artifact of increasing trap capture probabilities.
Differences in abundance of ant functional groups among burned and unburned habitats suggest that the effects of fire may impact functional diversity more than taxonomic diversity. In other studies of ants and wildfire, this has also been observed [69]. Those groups that benefited from simplification of the vegetation architecture (i.e., soil nesters with mounds) tended to increase in areas where the vegetation architecture was significantly impacted by fire (i.e., montane conifer and ponderosa pine forests), but they were relatively unimpacted or negatively impacted in areas where the vegetation architecture was already less complex pre-fire (montane grasslands). Interestingly, the abundance of ants that steal the nests of or nest within the nests of other species was higher in burned sites than unburned sites. This may also be a result of a simplified vegetation structure and increased ease of detecting possible hosts, or conversely, this could be a result of decreased defensive abilities of host ants.
Seasonal differences in insect composition, including ant species richness, abundance, and composition, are well-established in the literature [70,71], as are differences by habitat type [3,14]. Our study found that montane valley habitats supported more ants than other habitat types, possibly due to the resilience of grasslands to fire [72], and species composition differed between forested and grassland habitats. We also demonstrate the interactive effects of fire, macrohabitat, time since fire, and season on arthropod communities, a result supported by a recent meta-analysis by Vasconcelos et al. [73], which also found that wildland fire decreases the alpha diversity of ants, while vegetation type and complexity play a strong role in determining ant species richness. At the VCNP, after the Las Conchas wildfire, similar interactive effects of fire and habitat were observed for another ground-dwelling arthropod group (Araneae, [74]), but not for a more mobile group (Lepidoptera, [75]).
In summary, we have demonstrated that ant communities at the VCNP are resilient to high-severity wildland fire. We have also shown that wildfire effects on ant communities persist in this system for at least five years. Finally, we have supported previous work that found that the macrohabitat interacts with wildland fire to drive local ant species richness. This and concurrent work on other taxa at the VCNP will provide a biological framework for appropriately managing fire in montane southwestern ecosystems as changing climates promote a longer fire season, more intense fires, and more frequent fires [10].

Author Contributions

Conceptualization, R.V., J.K. and R.P. methodology, R.P., R.V. and J.K.; software, J.K. and R.V.; validation, R.V., T.S. and R.P.; formal analysis, R.V., J.K. and T.S.; investigation, J.K., R.V. and R.P.; resources, R.P. and R.V.; data curation, R.V. and T.S.; writing—original draft preparation, J.K.; writing—review and editing, R.V., T.S. and R.P.; visualization, R.V.; supervision, R.V. and R.P.; project administration, R.V. and R.P.; funding acquisition, R.V. and R.P. All authors have read and agreed to the published version of the manuscript.

Funding

This work was funded in part by a grant from the National Park Service, Southern Colorado Plateau Cooperative Ecosystem Studies Unit (CESU, Task Agreement P15AC01085) to Texas Tech University. Texas Tech University’s Department of Natural Resources Management also provided funding.

Data Availability Statement

All data are available on the U.S. National Park Service Integrated Resource Management Applications (IRMA) DataStore website.

Acknowledgments

Valles Caldera National Preserve staff and volunteers assisted with sample collection and sorting. C. Mitchell, C. Owens, and M. Hernandez Del Mar assisted with ant preparation and identification. Missouri University of Science & Technology provided student support.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Photos of study sites within each habitat. (A) Control mixed-conifer forest. (B) Burned mixed-conifer forests. (C) Ponderosa pine forest. (D) Burned ponderosa pine forest. (E) Control grassland. (F) Burned grassland. Photos B and F show pitfall traps inside elk enclosure fences. Photo credit: Valles Caldera National Preserve staff.
Figure 1. Photos of study sites within each habitat. (A) Control mixed-conifer forest. (B) Burned mixed-conifer forests. (C) Ponderosa pine forest. (D) Burned ponderosa pine forest. (E) Control grassland. (F) Burned grassland. Photos B and F show pitfall traps inside elk enclosure fences. Photo credit: Valles Caldera National Preserve staff.
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Figure 2. (A) Relationship between site (X), fire occurrence, season, and mean ant abundance (Y) from 2011 to 2015. (B) Relationship between site (X), fire occurrence, season, and mean ant abundance (Y) from 2011 to 2015. Bars indicate standard error. Orange color code indicates burned sites, and green color code indicates unburned sites. Black bars = MC. Grays bars = MV. White bars = PP.
Figure 2. (A) Relationship between site (X), fire occurrence, season, and mean ant abundance (Y) from 2011 to 2015. (B) Relationship between site (X), fire occurrence, season, and mean ant abundance (Y) from 2011 to 2015. Bars indicate standard error. Orange color code indicates burned sites, and green color code indicates unburned sites. Black bars = MC. Grays bars = MV. White bars = PP.
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Figure 3. (A) Seasonal ant abundance by fire treatment. (B) Seasonal abundance of eight most frequently collected genera. (C) Seasonal species richness by fire treatment.
Figure 3. (A) Seasonal ant abundance by fire treatment. (B) Seasonal abundance of eight most frequently collected genera. (C) Seasonal species richness by fire treatment.
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Figure 4. Host ant (X) and dulotic ant (Y) abundance show weak relationships that change with fire treatment. Red line and points are from burned sites (R2 = 0.0027), and green line and points are from unburned sites (R2 = 0.00614). Each point is a site at a given sampling time.
Figure 4. Host ant (X) and dulotic ant (Y) abundance show weak relationships that change with fire treatment. Red line and points are from burned sites (R2 = 0.0027), and green line and points are from unburned sites (R2 = 0.00614). Each point is a site at a given sampling time.
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Figure 5. The ways in which fires impacted ant functional groups varied among associations (A), nest types (B), and feeding habits (C). Dulotic = a social parasite. Host = a species enslaved by a social parasite species. Xeno = a xenobiont that shares a nest (mutualistically or commensally with other species), diet (P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids), and nest code (W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests).
Figure 5. The ways in which fires impacted ant functional groups varied among associations (A), nest types (B), and feeding habits (C). Dulotic = a social parasite. Host = a species enslaved by a social parasite species. Xeno = a xenobiont that shares a nest (mutualistically or commensally with other species), diet (P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids), and nest code (W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests).
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Table 1. Species observed during this study from 2011 to 2015. Habitats (MC = mountain conifer, MV = montane valley, PP = ponderosa pine), associations (F. Dulotic = facultative social parasite, engages in raiding behavior; O. Dulotic = obligate social parasite, depends on raiding host species for workforce; T. Dulotic = temporary social parasite, takes over host colonies for colony foundation; Host = a species enslaved by a social parasite species; Xeno = a xenobiont that shares a nest site mutualistically or commensally with other species), diet (P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids), and nest code (W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests). ? = taxonomic grouping is too broad for description or data are unknown. In the associations column, numbers represent known species associations, corresponding to the “No.” column. Associations with blank columns are those for which no data were available. An ‘x’ in a column denotes the presence of a species in a habitat type.
Table 1. Species observed during this study from 2011 to 2015. Habitats (MC = mountain conifer, MV = montane valley, PP = ponderosa pine), associations (F. Dulotic = facultative social parasite, engages in raiding behavior; O. Dulotic = obligate social parasite, depends on raiding host species for workforce; T. Dulotic = temporary social parasite, takes over host colonies for colony foundation; Host = a species enslaved by a social parasite species; Xeno = a xenobiont that shares a nest site mutualistically or commensally with other species), diet (P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids), and nest code (W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests). ? = taxonomic grouping is too broad for description or data are unknown. In the associations column, numbers represent known species associations, corresponding to the “No.” column. Associations with blank columns are those for which no data were available. An ‘x’ in a column denotes the presence of a species in a habitat type.
No.SpeciesMCMV PPAssociationsDietNesting Type
1Camponotus herculeanus x OHW
2Camponotus modocx xXeno 20PHW
3Camponotus ocreatus xx OU
4Dorymyrmex bicolor x PHS
5Dorymyrmex flavusxxx O?
6Dorymyrmex insanus x PHS
7Forelius mccooki x PSU
8Formica accretaxx Host 55O?
9Formica altipetensxxxHost 29, 30OHSU
10Formica argenteaxxxHost 55; Xeno 39OHWU
11Formica aservaxxxF. Dulotic 22, 28BHWUT
12Formica biophilica x Host 27OSU
13Formica bradleyi x Host 30OH?
14Formica subaenescensxx Host 55PHWSU
15Formica gnavaxxxXeno 46OH?
16Formica laevicepsx x OHSUT
17Formica lasioidesxx Host 30; Xeno 24, 38, 42, 50OSU
18Formica limataxxx OSU
19Formica montana xHost 29OST
20Formica neoclaraxx Host 29, 55; Xeno 2OHWSU
21Formica neogagatesxxxHost 27, 30, 55; Xeno 32OHSU
22Formica neorufibarbisx Host 11, 29, 30, 55OHWU
23Formica obscuriventris xHost 27OUT
24Formica occultaxx Host 55; Xeno 17, 39, 50OSU
25Formica opaciventrisx x OST
26Formica pallidefulvaxx Host 27, 29OWS
27Formica pergandeixxxF. Dulotic 12, 21, 23, 26, 29BSU
28Formica podzolicaxx Host 11, 27, 55OSUT
29Formica puberula xxF. Dulotic 9, 13, 17, 21, 22BSU
30Formica wheeleri xxF. Dulotic 9, 13, 17, 21, 22BST
31Formica xerophilaxx OSU
32Lasius americanus xxXeno 21PHU
33Lasius aphidicolaxxxT. Dulotic 34, 38, 39OHWSU
34Lasius crypticusxx Xeno 45OHWU
35Lasius humilis X OHU
36Lasius neonigerxx Host 33, 40OHSU
37Lasius ponderosaexxxHost 33OHSU
38Lasius pallitarsisx xHost 40; Xeno 10, 17, 50OHWU
39Lasius sitiensxxxXeno 10, 24, 45, 50OHU
40Lasius subumbratusxxxT. Dulotic 36, 38HWU
41Lasius xerophilusxxx OHS
42Leptothorax crassipilis xXeno 17, 24, 50OHSU
43Leptothorax canadensisxx OHWUL
44Liometopum luctuosumx OHU
45Monomorium minimumx Xeno 14, 34, 39OHWSUD
46Myrmecina americana x Xeno 15PSL
47Myrmica americanaxx PSL
48Myrmica brevispinosa x PSL
49Myrmica discontinuaxx PSL
50Myrmica hamulataxx Xeno 17, 24, 38, 39, 42PSL
51Myrmica incompletaxxx PHSL
52Myrmica latifronsxxx PSL
53Myrmica monticolaxxx PSL
54Pheidole sp.xxx ??
55Polyergus mexicanusx O. Dulotic 8, 10, 14, 20, 21, 22, 24, 28BD
56Solenopsis invicta x PH?
57Solenopsis pilosula x PSD
58Stenamma diecki x PSCL
59Tapinoma sessilexxx PHWSUCLD
60Temnothorax rugatulus x OHWSUCLD
Table 2. Multivariate analysis of variance comparing mean ant species richness (Y) among sites, fire treatment, seasons, and interactions (X). DF = degree of freedom. SS = sum of squares. * = significance (alpha = 0.05). F35,1227 = 8.14. p < 0.0001.
Table 2. Multivariate analysis of variance comparing mean ant species richness (Y) among sites, fire treatment, seasons, and interactions (X). DF = degree of freedom. SS = sum of squares. * = significance (alpha = 0.05). F35,1227 = 8.14. p < 0.0001.
FactorDFSSFp
Year*Season*Site*Burned432.612.000.0920
Year*Season*Site413.110.800.5220
Year*Season*Burned221.972.700.0678
Season*Site*Burned43.170.190.9413
Year*Site*Burned229.583.630.0268 *
Year*Site2157.3319.32<0.0001 *
Year*Burned1126.5431.07<0.0001 *
Season*Burned218.022.210.1099
Season*Site435.562.180.0688
Season*Year2158.1419.42<0.0001 *
Site*Burned212.751.570.2093
Year1457.87112.44<0.0001 *
Season2996.99122.41<0.0001 *
Site2150.3018.45<0.0001 *
Burned1126.5431.07<0.0001 *
Table 3. Multivariate analysis of variance comparing mean ant abundance (Y) among sites, years, fire treatment, seasons, and interactions (X). DF = degree of freedom. SS = sum of squares. * = significance (alpha = 0.05). F35,1227 = 9.04. p < 0.0001.
Table 3. Multivariate analysis of variance comparing mean ant abundance (Y) among sites, years, fire treatment, seasons, and interactions (X). DF = degree of freedom. SS = sum of squares. * = significance (alpha = 0.05). F35,1227 = 9.04. p < 0.0001.
FactorDFSSFp
Season*Site*Burned*Year442,194.701.090.3581
Season*Burned*Year224,901.901.290.2753
Season*Site*Year41161.000.300.8765
Site*Burned*Year2125,303.606.500.0016 *
Season*Site*Burned45624.601.460.2127
Season*Burned228,718.901.490.2260
Season*Site4306,994.107.96<0.0001 *
Site*Burned216,934.700.880.4159
Season*Year266,358.003.440.0324 *
Site*Year259,479.203.080.0461 *
Burned*Year1133,598.4013.850.0002 *
Season2604,899.8031.36<0.0001 *
Burned119,106.901.980.1595
Year154,294.105.630.0178 *
Table 4. Number of species appearing and disappearing between years. Species turnover is calculated as the (number of appearing species + number of disappearing species divided by the total number of species observed) × 100.
Table 4. Number of species appearing and disappearing between years. Species turnover is calculated as the (number of appearing species + number of disappearing species divided by the total number of species observed) × 100.
Year and TreatmentTotal SpeciesSpecies AppearingSpecies DisappearingSpecies TurnoverPercent Change
201153n/an/an/an/a
Burned47n/an/an/an/a
Unburned46n/an/an/an/a
20124913944.8−7.6
Burned4010332.5−14.9
Unburned3710745.9−19.6
20134512435.5−8.2
Burned363727.7−10
Unburned3111654.8−16.3
20143715348.6−17.8
Burned2611146.2−27.8
Unburned9181211−70.1
20152510144.0−32.4
Burned1401285.7−46.1
Unburned151220+40.0
Table 5. Multivariate analysis of variance comparing mean abundance of xenobionts, host species, and dulotic species (Y) among sites, years, fire treatment, seasons, and interactions (X). p-values are reported (alpha = 0.05).
Table 5. Multivariate analysis of variance comparing mean abundance of xenobionts, host species, and dulotic species (Y) among sites, years, fire treatment, seasons, and interactions (X). p-values are reported (alpha = 0.05).
XenoHostDulotic
Burned0.00900.00030.7277
Season<0.0001<0.00010.1467
Site<0.00010.00120.0058
Season*Burned0.51780.09870.6736
Site*Season0.00010.00260.1079
Site*Burned0.17260.00060.6646
Table 6. Multivariate analysis of variance comparing ant nesting types (Y) among sites, years, fire treatment, seasons, and interactions (X). W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests). p-values are reported (alpha = 0.05).
Table 6. Multivariate analysis of variance comparing ant nesting types (Y) among sites, years, fire treatment, seasons, and interactions (X). W = in rotten wood or under bark; S = in soil with crate or soil mound at nest entrance; U = under stones or logs, specifically at the interface between earth and object, and can also be at the bases of plants; T = thatch mound; C = cavities such as tree holes and spaces between stones; L = in twigs or pockets of leaf litter or in acorns; D = steals nests or nests inside other species’ nests). p-values are reported (alpha = 0.05).
WSUTCLD
Burned0.96000.71810.04010.00110.07110.01040.0415
Season<0.0001<0.0001<0.00010.07190.00020.0043<0.0001
Site0.00440.6347<0.00010.2745<0.0001<0.0001<0.0001
Season*Burned0.1122<0.00010.26040.23120.12310.98700.1781
Site*Season0.07720.0131<0.00010.30000.00360.00450.0084
Site*Burned0.98400.00050.22140.37040.0241<0.00010.0413
Site*Burned*Season0.94790.54300.61010.27940.21190.88000.1717
Table 7. Multivariate analysis of variance comparing ant diet types (Y) among sites, years fire treatment, seasons, and interactions (X). P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids.
Table 7. Multivariate analysis of variance comparing ant diet types (Y) among sites, years fire treatment, seasons, and interactions (X). P = omnivorous and actively predaceous; O = omnivorous and not actively predaceous, i.e., primarily a scavenger; B = omnivorous and dulotic; H = honeydew, i.e., tending aphids, mealybugs, or coccids.
POBH
Burned0.46220.00120.00460.0180
Season<0.0001<0.00010.0147<0.0001
Site<0.0001<0.00010.0019<0.0001
Season*Burned0.79260.007850.23370.5669
Site*Season0.0012<0.00010.25440.0018
Site*Burned0.00260.0242<0.00010.6983
Site*Burned*Season0.51610.09020.13160.0156
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MDPI and ACS Style

Knudsen, J.; Parmenter, R.; Sumnicht, T.; Verble, R. High-Severity Wildfires Alter Ant (Hymenoptera: Formicidae) Foraging Assemblage Structure in Montane Coniferous Forests and Grasslands in the Jemez Mountains, New Mexico, USA. Conservation 2024, 4, 830-846. https://doi.org/10.3390/conservation4040049

AMA Style

Knudsen J, Parmenter R, Sumnicht T, Verble R. High-Severity Wildfires Alter Ant (Hymenoptera: Formicidae) Foraging Assemblage Structure in Montane Coniferous Forests and Grasslands in the Jemez Mountains, New Mexico, USA. Conservation. 2024; 4(4):830-846. https://doi.org/10.3390/conservation4040049

Chicago/Turabian Style

Knudsen, Jonathan, Robert Parmenter, Theodore Sumnicht, and Robin Verble. 2024. "High-Severity Wildfires Alter Ant (Hymenoptera: Formicidae) Foraging Assemblage Structure in Montane Coniferous Forests and Grasslands in the Jemez Mountains, New Mexico, USA" Conservation 4, no. 4: 830-846. https://doi.org/10.3390/conservation4040049

APA Style

Knudsen, J., Parmenter, R., Sumnicht, T., & Verble, R. (2024). High-Severity Wildfires Alter Ant (Hymenoptera: Formicidae) Foraging Assemblage Structure in Montane Coniferous Forests and Grasslands in the Jemez Mountains, New Mexico, USA. Conservation, 4(4), 830-846. https://doi.org/10.3390/conservation4040049

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