Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 341 2018 NORTHEASTERN NATURALIST 25(2):341–354 Leaf Litter and Arboreal Ants (Hymenoptera: Formicidae) in a Mid-Atlantic Forest Hunter R. Mann1, Emily Rowe1, Jennifer Selfridge2, and Dana L. Price1,* Abstract - The majority of ant diversity studies have been conducted in the tropics, but the number of studies conducted in temperate regions, like the US, is on the rise. Our research measured the richness and diversity of ants (Hymenoptera: Formicidae) in a coastal, temperate forest of Maryland. We collected leaf litter along twenty-four 100-m transects during May, July, and September of 2015 and measured litter depth and mass to determine if there was a relationship with ant abundance (density). We used Berlese funnels to collect 14 ant species (4208 individuals; n = 144 samples). Neither leaf-litter depth nor mass had an impact on ant abundance or species richness. We used aspirators and hand-collection methods to examine arboreal-ant foraging preference on deciduous and coniferous trees. Total species richness for both tree types was 21; we collected 19 species from deciduous trees and 17 from coniferous trees. Four species detected on deciduous trees were not present on conifers, and 2 species from conifers were not observed on deciduous trees. We observed higher abundances on deciduous trees (P = 0.016) and detected a preference for larger trees. We provide suggestions for conservation efforts for the coastal forests of Maryland, and our study contributes to the growing species inventory of ants on Maryland’s eastern shore. Introduction Environmental-indicator taxa are sensitive to disturbance, making them useful in assessing habitat health. These taxa provide an estimate of species diversity, often in terms of the number of different species in a given area that can be monitored and used to gain an overview of changing ecosystem trends (Andersen 1997, Longino et al. 2002, McGeoch 1998). Indicator taxa can also be used to locate areas of high biodiversity (bioindicators) or estimate the impact from a specific one-time disturbance event (Caro and O’Doherty 1999). High levels of diversity in a given region act as a safeguard against the collapse of an ecosystem; thus, the loss of a single species may be filled by another, with a similar niche, or by other organisms with similar lifestyles (Ribas et al. 2003). Biodiversity surveys rely on indicator taxa for conservation planning, developing management plans, studying the impact of agricultural practices on habitat, and providing justification for the existence of protected areas (Caro and O’Doherty 1999, Spector and Forsyth 1998, Underwood and Fisher 2006). Ants (Hymenoptera: Formicidae) are often cited as excellent biodiversity and environmental indicator taxa due to their great abundance, species diversity, and ability to occupy the vast majority of terrestrial habitats (Andersen 1997, 1 University of Salisbury, 1101 Camden Avenue, Salisbury, MD 21801.2 Maryland Department of Natural Resources, 909 Wye Mills Road, Wye Mills, MD 21679. *Corresponding author - dlprice@salisbury.edu. Manuscript Editor: Daniel Pavuk Northeastern Naturalist 342 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 Vol. 25, No. 2 Bestelmeyer and Wiens 2001, Martelli et al. 2004, Underwood and Fisher 2006). Ant species often react quickly to changes in their environment, with decreased diversity levels after an ecosystem is disturbed, and increasing diversity as the habitat recovers (Longino et al. 2002, Martelli et al. 2004, McGlynn et al. 2009). Despite their global ecological importance, the majority of ant studies have taken place in the tropics (Ellison et al. 2007). In many regions of the US, basic information such as species range is incomplete even for common taxa. Partial state inventories have been conducted in California and Oregon (Ratchford et al. 2005, Sanders et al 2007a), Florida (King and Porter 2005), Oklahoma (Albrecht and Gotelli 2001), New York (Ellison et al. 2007), and Tennessee (Sanders et al. 2007b). The only state to have completed what is considered a full inventory of all ants within its borders is Ohio (Coovert 2005, Ellison et al. 2007). In Maryland, ant studies and investigation of ant ecology have mostly focused on species richness and seasonality in habitats west of the Chesapeake Bay (Lynch 1981; Lynch et al. 1980, 1988) or interactions between different ant species (Fellers 1987, 1989). Only 2 biodiversity studies have been conducted on Maryland’s eastern shore. Frye and Frye (2012) hand-sampled ants from Pinus echinata Mill. (Shortleaf Pine), P. taeda L. (Loblolly Pine) and several species of Quercus (oak) in ridge woodlands and inland dune communities of Worcester County, MD. To further expand their survey, Frye et al. (2014) used pitfall traps and leaf-litter samples to examine ant species richness and diversity of inland sand-dune communities. These studies were restricted to xeric habitats characterized by dry conditions with low moisture levels (Frye et al. 2014). We investigated species diversity and richness of ants in the E.A. Vaughn Wildlife Management Area (WMA), located in Girdletree, MD. Our primary objectives were to collect leaf-litter ants to determine if ant abundance (density) was correlated with leaf-litter depth or mass. We also collected arboreal ants from deciduous and coniferous trees to examine foraging preference for tree type. Our research provides valuable baseline ant biodiversity data for monitoring the stability and health of a coastal temperate forest of the mid-Atlantic region. Field-site Description We conducted this study in the E.A. Vaughn Wildlife Management Area (WMA), located in lower Worcester County, MD (38º4'49''N, 75º23'38''W). This WMA has been part of Maryland’s Department of Natural Resources public land system since 1943 and it has since expanded in size with subsequent acquisitions (MDNR 2016a); E.A. Vaughn WMA is currently managed by the Wildlife and Heritage Service. The E.A. Vaughn WMA consists of 1064 ha of mixed deciduous/coniferous forest (MDNR 2016a) surrounded by agriculture on all sides, a feature typical of forested areas in the mid-Atlantic region (Crist 2009). Our study site was composed of both dry uplands and low-lying wetlands that are periodically flooded by the Maryland Department of Natural Resources during the fall as part of a “green tree reservoir” (MDNR 2016b). Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 343 Methods Collections We sampled leaf litter and arboreal foraging ants from a 60,000-m2 area once a week during May, July, and September of 2015 (12 weeks total). In order to reduce the likelihood of sampling from the same colonies more than once, we sampled twenty-four, 100-m transects, set 25 m apart. To prevent sampling from nearby transects during the same week, we divided the study site into 2 sections (transects 1–12 and 13–24) and sampled 1 transect from each section each week. We followed the methods of Frye et al. (2014) and conducted research between 1000 hrs and 1600 hrs, when ants are known to be most active. We collected samples for both leaf-litter and tree-foraging ants on the same dates, at the same time of day, and from the same transects at the site location, to eliminate these factors as potential explanations for any differences detected. We sorted, curated, and identified worker ants using a variety of resources, including Coovert’s (2005) The Ants of Ohio (Hymenoptera: Formicidae), and A Field Guide to the Ants of New England (Ellison et al. 2012). We excluded queens and male ants from the analysis. We sent to Tim Foard (i2LResearch USA Inc, Baltimore, MD) all specimens that we were unable to identify. Reference specimens will be deposited in the Salisbury University Price Entomological Collection (Salisbury, MD), and the Towson University Biodiversity Center (Towson, MD). Leaf-litter ants Using a 0.25-m2 quadrat, we collected leaf litter every 20 m along each transect (6 data points per transect for a total of 12 per week). At each data point, we used a ruler to measure the depth of leaf litter in each corner and in the center, then calculated the average of the 5 depths, similar to Kaspari (1996a, 1996b). We transported all leaf-litter samples to the greenhouse for Berlese litter extraction. In the greenhouse, we used Berlese funnels made of 18.9-L (5-gallon) buckets with a 25-cm–diameter tractor funnel set inside each one. We placed a piece of 0.64 cm x 0.64 cm-mesh screen in each funnel to hold the leaf litter while allowing ants to travel through, and attached a WhirlPak bag filled with 95% ethanol to the bottom of the funnel to collect insects as they fell through the mesh. We set a 65- watt bulb above the litter for up to 96 h until the leaf litter was dry. In the lab, we weighed (g) the dry leaf litter taken from each Berlese sample after ant extraction. Arboreal foraging ants We employed aspirators and hand-collection methods to sample arboreal foraging ants every 20 m (6 points per transect) along the same 24 transect lines described above. For each data point, we measured the nearest coniferous and deciduous tree ranging in diameter from 50 cm to 250 cm. We observed each tree from the base of the trunk up to a height of 2 m for 15 min (Frye and Frye 2012). When an ant foraging-line was observed, we sampled only 10 ants to avoid oversampling a single species. If a tree of appropriate size or type (deciduous/conifer) was not Northeastern Naturalist 344 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 Vol. 25, No. 2 present, we recorded it as an absence. Thus, we collected samples from 144 deciduous trees and 123 coniferous trees. Deciduous trees included Quercus falcata Michx. (Southern Red Oak), Quercus bicolor Willd. (Swamp White Oak), Fraxinus americana L. (White Ash), and F. pennsylvanica Marsh. (Green Ash). Coniferous trees were Loblolly Pine and Pinus virginiana Miller (Virginia Pine). We also observed Ilex opaca Aiton (American Holly), but did not include it in this study. Limitations of hand-sampling methods include the potential for small or cryptic species to be overlooked (Underwood and Fisher 2006) and varying levels of experience among individual collectors (Gotelli et al. 2011). To avoid these biases, the same 4 people sampled each week, and we employed 15-min sample periods to allow more time to detect small or cryptic species. Data analysis We calculated 3 non-parametric species-richness estimators—Chao2, ICE (incidence based coverage estimator), and Jackknife for both leaf-litter and arboreal- foraging ant data (Chao 1984, 1987; Chao et al. 2000; Chazdon et al. 1998; Gotelli and Colwell 2010) in EstimateS 9.1 software (Colwell and Coddington 1994). We used species-accumulation curves to illustrate the rate at which new species were sampled (Gotelli and Colwell 2010). We used generalized linear models (GLM, SPSS 21.0.0) to test relationships between ant abundance and leaf-litter mass and depth. We created scatterplots to visualize the relationships between leaflitter mass (g) and ant abundances, and depth and ant abundances. We used a non-parametric Mann–Whitney U test in SPSS 21.0.0 to determine if ant abundances differed between deciduous trees versus coniferous trees. We compared differences in ant abundance by tree type, tree circumference, and the interaction of tree type and circumference with GLM (SPSS 21.0.0) log transformation. This approach allowed us to determine whether ant abundances varied between tree sizes and to test if there was an interaction between tree type and tree size. Results Leaf-litter ants We collected 14 species, including 4208 individuals of leaf-litter ants in the E.A. Vaughn WMA (Tables 1, 2). We collected Nylanderia faisonensis from all 24 transects; this species accounted for 57% of the all individuals collected. Estimates of species richness, Chao2 (14), ICE (14.57) and Jackknife (14.96) suggest 1 additional species might be found in the leaf litter (Table 2). The species-accumulation curve supports these findings, with the majority of species collected in the first 8 samples (transects; Fig. 1). We detected no correlations between ant abundance and leaf-litter depth (r2 = 0.00004, P = 0.94) or dry leaf-litter mass (r2 = 0.000009, P = 0.97). We also saw no correlation between species richness and leaf-litter depth (r2 = 0.003, P = 0.50) or dry leaf-litter mass (r2 = 0.002, P = 0.61). Our GLM analysis to examine ant density detected no effect of leaf-litter depth (F = 0.493, P = 0.484) or leaf-litter mass (F = 0.251, P = 0.618). Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 345 Table 1. Leaf-litter ants collected via Berlese extraction from 144 data points at the E.A. Vaughn WMA during May, July, and September of 2015. Species name May July September Total abundance Aphaenogaster fulva Roger 5 154 85 244 Aphaenogaster rudis Enzmann 83 13 16 112 Crematogaster cerasi (Fitch) 1 1 0 2 Formica subsericea Say 1 0 0 1 Lasius alienus (Foerster) 241 116 8 365 Lasius umbratus (Nylander) 2 203 1 206 Myrmecina americana Emery 28 55 75 158 Nylanderia faisonensis (Forel) 784 877 750 2411 Ponera pennsylvanica Buckley 185 136 101 422 Solenopsis carolinensis Forel 44 0 0 44 Solenopsis molesta (Say) 11 87 0 98 Stigmatomma pallipes (Haldeman) 3 13 5 21 Strumigenys dietrichi Smith, M.R. 1 65 12 78 Temnothorax curvispinosus (Mayr) 9 22 15 46 Total 1398 1742 1068 4208 Table 2. Species richness and abundance of leaf-litter ants collected from E.A. Vaughn WMA during May, July, and September of 2015. Statistic Leaf-litter ants Species richness 14 Total abundance 4208 Estimated species richness Chao 2 14 ICE 14.57 Jackknife 14.96 Figure 1. Species- accumulation curve of leaf-litter ants collected from 24 transects (6 traps per transect) at the E.A. Vaughn WMA during May, July, and September of 2015. Northeastern Naturalist 346 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 Vol. 25, No. 2 Arboreal foraging ants We sampled 21 species of arboreal ants, comprising 679 individuals (Tables 3, 4). We sampled 19 species (446 individuals) from deciduous trees and 17 species (233 individuals) from coniferous trees (Table 3 ,4). Four species collected from deciduous trees were not detected on conifers, and 2 from conifers were not found on deciduous trees. Results of a paired t-test showed that overall ant abundance was higher on deciduous than coniferous trees (P = 0.016). The 2 most common species, Aphaenogaster fulva and A. rudis both preferred foraging on deciduous trees to conifers, P = 0.005 and P = 0.005, respectively. We were unable to calculate a statistical preference for other species due to a lack of data. When we ran GLM, Table 4. Species richness and abundance of arboreal ants sampled from deciduous and coniferous trees during May, July, and September of 2015 from E.A. Vaughn WMA. Statistic Deciduous Coniferous Species richness 19 17 Abundance 446 233 Estimated species richness Chao2 27.63 37.13 ICE 25.37 24.01 Jackknife 24.75 23.71 Table 3. Ants sampled from 144 deciduous and 123 coniferous trees in the E.A. Vaughn WMA, during May, July, and September of 2015. Deciduous Coniferous Species name May July Sept. Total May July Sept. Total Aphaenogaster fulva Roger 34 66 16 116 7 34 0 41 Aphaenogaster rudis Enzmann 12 28 63 103 5 7 16 28 Camponotus castaneus (Latreille) 1 5 2 8 0 3 0 3 Camponotus chromaiodes Bolton 5 4 1 10 6 2 0 8 Camponotus nearcticus Emery 8 13 2 23 3 0 2 5 Camponotus pennsylvanicus (De Geer) 2 10 2 14 5 1 4 10 Camponotus subbarbatus Emery 0 1 0 1 0 1 0 1 Crematogaster ashmeadi Mayr 1 10 0 11 12 2 10 24 Crematogaster cerasi (Fitch) 0 1 0 1 2 1 1 4 Crematogaster pilosa Emery 16 0 0 16 0 0 2 2 Formica neogagates Viereck 0 0 0 0 0 2 0 2 Formica subsericea Say 22 10 0 32 0 0 0 0 Lasius alienus (Foerster) 32 20 2 54 1 35 13 49 Lasius umbratus (Nylander) 0 2 0 2 6 17 10 33 Myrmecina americana Emery 3 3 2 8 1 0 0 1 Nylanderia faisonensis (Forel) 5 12 15 32 1 17 2 20 Stigmatomma pallipes (Haldeman) 0 0 1 1 0 0 0 0 Temnothorax curvispinosus (Mayr) 3 0 1 4 1 0 0 1 Temnothorax longispinosus (Roger) 7 0 1 8 0 0 0 0 Temnothorax morpho spp. 0 2 0 2 0 0 0 0 Temnothorax schaumii (Roger) 0 0 0 0 1 0 0 1 Total 151 187 108 446 50 122 60 233 Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 347 the interaction between tree size and tree type was removed from the model. Ant abundance was higher on deciduous trees over coniferous (F = 8.48, P = .004; Fig. 2). Tree size also had an effect on foraging behavior; ant abundance was higher on large deciduous trees over small trees (F = 5.025, P = .026; Fig. 3). We detected no difference for coniferous trees. Estimates of ant species richness are 27.63 and 37.13 (Chao2), 25.37 and 24.01 (ICE), and 24.75 and 23.71 (Jackknife) for deciduous and coniferous trees, respectively; all estimates suggest several arboreal foraging species have yet to be sampled in E.A. Vaughn WMA (Table 4). Species-accumulation curves corroborate these findings for both deciduous and coniferous trees; neither curve has neared an asymptote (Fig. 4). Figure 2. Mean abundance of ants sampled from deciduous and coniferous trees. Figure 3. Mean abundance of ants sampled from small, medium, and large deciduous and coniferous trees. Northeastern Naturalist 348 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 Vol. 25, No. 2 Discussion Of the 24 species we collected in our study, 4 species collected from leaf litter were not collected on trees—Ponera pennsylvanica, Strumigenys dietrichi, Solenopsis carolinensis, and S. molesta. Species collected only from trees include 5 species of the genus Camponotus, all of which are reported to live in forested habitats, nesting in either decaying wood, rotting logs, stumps, or in living trees (Coovert 2005, Ellison et al. 2012). We collected Crematogaster ashmeadi only from trees, and it is reported in other studies to be one of the most abundant arboreal ants sampled from pine forests (Frye and Frye 2012, Tschinkel and Hess 1999). We also observed Crematogaster pilosa, Formica neogagates, Temnothorax longispinosus, and T. schaumii only on trees. All 4 species reportedly forage on trees or on logs under bark (Coovert 2005). Leaf-litter ants We collected 14 species of leaf-litter ants (4208 individuals) from a 60,000-m2 area during May, July, and September of 2015, in the E.A. Vaughn WMA. Species richness and biodiversity at our survey site within E.A. Vaughn WMA are similar to those reported for other temperate leaf-litter ant studies conducted in the mid- Atlantic (Ellison et al. 2007, Lynch 1981, Lynch et al. 1988). In Edgewater, MD, on the Coastal Plain west of the Chesapeake Bay, Lynch (1981) reported a maximum of 14 species collected from litter samples in 3 different habitats. Lynch et al. (1988) collected 4124 ant specimens representing 22 species in a mature deciduous forests of the Maryland coastal plains, and Ellison et al. (2007) reported 21 species in Black Rock Forest (Cornwall, NY) when using sieved litter samples. Altered or fragmented habitats with low diversity tend to be dominated by a small number of extremely abundant generalist species (Resende et al. 2013). In Figure 4. Species- accumulation curves for arboreal foraging ants sampled from 24 transects of paired deciduous (black squares; 144 trees total) and c o n i f e r o u s trees (black circles; 123 total) at the E.A. Vaughn WMA during May, July, and September of 2015. Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 349 the E.A. Vaughn WMA, the most notable of these species was N. faisonensis, supporting the findings of Lynch et al. (1988), who reported N. faisonensis (reported as Paratrechina faisonensis) to be the most abundant species, making up 61.2% of the total individuals collected. This species is distributed throughout the eastern US, from New Jersey to Florida, west to Ohio and throughout the southeast (Coovert 2005, Trager 1984). It has also been reported as one of the most common ant species in the Mid-Atlantic (Coovert 2005, Kjar 2009), especially within deciduous forests or mixed deciduous–pine forests (Trager 1984). Similar to the findings of Lynch et al. (1988), Ponera pennsylvanica was the second most abundant species found in our study site. This species was present in small numbers at over half of our collection points (78 out of 144), and we collected it from 23 of 24 transects. This species is widely distributed across much of the eastern and central US (MacKay and Anderson 1991) and reportedly forms small colonies under rocks, within rotting wood, and is rarely seen on the surface (Coovert 2005). Additional species sampled in high abundance (>100 specimens) include, Aphaenogaster fulva, and A. rudis, Lasius alienus, and L. umbratus, where the latter was only abundant at 1 collection point (196 individuals of the 206 collected). Kjar (2009) reports A. fulva as one of the most common ant species in the region along with N. faisonensis. Leaf-litter mass and depth The role of leaf-litter mass in predicting ant abundance has been the focus of numerous studies, with some authors reporting a correlation between ant abundance and increased amounts of leaf litter. Research in a Costa Rican forest suggests that leaf-litter mass plays a key role in determining ant abundance and species richness (Lopes and Vasconcelos 2008, McGlynn et al. 2009). These authors reported increases in ant abundance with increasing litter mass. Studies of this type, however, are not unanimous in their findings. Lynch et al. (1988) found a significant negative correlation with ant abundance and leaf-litter mass indicating that ant abundance increased as leaf-litter mass decreased. Wilkie et al. (2010) found no association between ant species richness and leaf-litter mass, while Kaspari (1996a) found only a very weak, positive connection (in the form of nest-site abundance). Our findings suggest that neither leaf-litter depth nor mass had an effect on species richness or abundance (density) of leaf-litter ants. This finding was not unexpected, considering that the composition of trees in terms of density, size, and type varied only slightly throughout the study site, with little variation in leaf-litter mass and depth throughout (134.7–377.1 g and 1.1–5.0 cm, respectively). Arboreal foraging ants Ants are one of the most common arthropod groups utilizing tree trunk surfaces (Hanula and Franzreb 1998); thus, these habitats are important for the examination of ant ecology and foraging behavior. The most abundant arboreal species sampled include Aphaenogaster fulva, A. rudis, and Lasius alienus. Both A. fulva, and A. rudis have wide ranges that encompasses most of the eastern continental US, and are frequently observed in forest and forest-edge habitats. Lasius alienus is Northeastern Naturalist 350 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 Vol. 25, No. 2 extremely common throughout most of North America and Europe (Coovert 2005). Aphaenogaster fulva is noted as being more common in old-growth forests (Kjar 2009), so its relative abundance in the fractured E.A. Vaughn WMA with nearby agriculture and roads is surprising. Conversely, the presence of generalist species such as L. alienus, which is more likely to be found near human habitation, is less surprising (Kjar 2009). We found a significant preference by the arboreal ants of the E.A. Vaughn WMA for foraging on deciduous trees rather than coniferous trees. Reasons for a preference are likely multifaceted and cover a number of factors relating to various evolutionary history traits. Interactions with other insect species that use bark as a conduit from the forest floor to the canopy (Majer et al. 2003), differences in bark structure (Majer et al. 2003; Menzel et al. 2004; Nicolai 1993, 1995), and tree size/ diameter (Tschinkel and Hess 1999; Verble and Stephen 2009a, b) have all been reported to have an effect on arboreal-ant–foraging behaviors. Our estimates of species richness on both deciduous and coniferous trees are high. Although we recognize that more ant species are likely present at this site, the low number of ants sampled from conifers suggests the Chao2 prediction of 37.13 species is unlikely. Tree size We considered tree size to be a potential variable in the foraging behavior of arboreal ants. In our study, ants foraging on deciduous trees preferred larger trees to medium or small ones; we found a significant difference between foraging on large and small trees (P = 0.039). Several authors have suggested the increased structuring of the bark seen on older, larger trees contributes to a greater abundance of arthropods (Nicolai 1993, Ulyshen 2011). Tschinkel and Hess (1999) found no difference in species richness within different tree-size classes, but observed a change in species composition favoring arboreal species as tree size increased. The authors also noted an increase in the proportion of trees supporting ants among larger size categories. Conclusions Our findings regarding ants’ preferences for tree size and type for foraging in a fractured temperate forest reinforces the idea that ants generally forage more often on deciduous trees than conifers. Thus, if many ant species prefer deciduous trees, the increased planting and spread of coniferous trees may have implications for ant species richness in the region. Research on ground beetles (Coleoptera: Carabidae) suggested that mixed forests with both deciduous and coniferous species were important for the well being of some species, and monocultures of conifers had a negative impact on diversity (Koivula et al. 1999). Studies in Europe found that replacing conifer monocultures with a polyculture of different tree species increased diversity for species that were not already rare (Felton et al. 2010a, b). Thus, maintenance of mixed forests, or at the very least, prevention of conifer monocultures is likely important for sustaining healthy ant populations. Due to the habitat variation at this site, and the overall mission of conservation of sensitive species (Maryland Northeastern Naturalist Vol. 25, No. 2 H.R. Mann, E. Rowe, J. Selfridge, and D.L. Price 2018 351 Department of Natural Resources 2012), we recommend further assessment of ant ecology in this region. This research will provide valuable information regarding long-term management practices of the coastal forests of Maryland. Acknowledgments We thank a number of people who were critical to the completion of this project, including Mallory Hagadorn, for help with the collection of ants; Roman Jesien and John LaPolla, for their initial help with project design; and John also for his comments regarding this manuscript. We are grateful to John Moulis for providing us with permits for ant collection in the E.A. Vaughn WMA, Tim Foard for his help with ant identifications, and Eric Leibgold for his suggestions and comments regarding the final report. This research would not have been possible without support and funding provided by Salisbury University, including a Guerrieri undergraduate summer fellowship for Emily Rowe, graduate research and presentation awards, and a Henson undergraduate research grant. Literature Cited Albrecht, M., and N.J. Gotelli. 2001. 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