Moth communities correspond with plant communities in midwestern (Indiana, USA) sand prairies and oak barrens and their degradation endpoints.
Abstract: Despite immense diversity, insect conservation is typically species specific. Effective insect conservation will require efforts that capture insect species and communities at all levels of biological organization. Surrogate conservation targets, such as habitat based conservation planning tools were designed to capture poorly understood taxa such as invertebrates. We evaluated a botanically-based community filter across disturbance gradients in NW Indiana to determine if moth communities (Lepidoptera) responded similarly to vascular plant assemblages. Our 13 sample sites included high-quality ecosystem remnants (sand prairies and oak barrens) and their local degradation endpoints (exotic old fields and fire-suppressed oak woodlands). Monthly, we quantitatively sampled moths using ultraviolet light traps and inventoried vascular plant species at each sample site. Analysis of moth and plant community relationships using Bray-Curtis coefficients of dissimilarity produced statistically congruent relationships between moth and plant assemblages at the sample sites indicating that these two taxonomic groups respond to ecological gradients and disturbance similarly. Other measures of botanical community integrity used to select conservation areas such as floristic quality assessment index and diversity indices do not translate directly to measures of moth species richness or diversity. We suggest that in this system, vascular plant assemblages are a reasonable conservation surrogate for moth communities.
Article Type: Report
Subject: Oak (Environmental aspects)
Moths (Environmental aspects)
Plants (Environmental aspects)
Prairies (Environmental aspects)
Authors: Shuey, John A.
Metzler, Eric H.
Tungesvick, Kevin
Pub Date: 04/01/2012
Publication: Name: The American Midland Naturalist Publisher: University of Notre Dame, Department of Biological Sciences Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Earth sciences Copyright: COPYRIGHT 2012 University of Notre Dame, Department of Biological Sciences ISSN: 0003-0031
Issue: Date: April, 2012 Source Volume: 167 Source Issue: 2
Geographic: Geographic Scope: United States Geographic Code: 1USA United States
Accession Number: 287956850
Full Text: INTRODUCTION

Insects, by their very nature, defy comprehensive conservation planning. Their vast diversity, measured by both numbers of species as well as life history traits, preclude detailed knowledge of the status and distribution of all but a few species and at best, conservation strategies will be developed around a handful of representative taxa or imperiled species.

Almost all insect conservation efforts are focused at the level of species (Pyle, 1976, 1995; Pyle et al., 1981; Samways, 1997) or umbrella species which may serve as surrogates for other lesser known entities (Main, 1996; New, 1997); for example, New (1997) describes 22 Lepidoptera conservation case studies, all of which revolve around species-specific conservation targets. Because of this, the perception of insect conservation is species focused and biased towards charismatic taxa such as butterflies.

Insect conservation requires goals that are set at the faunal level and conservation success requires efforts that identify and conserve all species at all levels of biological organization. This seemingly daunting task could be approached using surrogate planning targets that effectively plan for and capture representative insect communities. Coarse filter, habitat based conservation planning tools were developed by conservation NGO's and governments as efficient planning tools for identifying arrays of complimentary conservation action sites (Faber-Langendoen, 2001; Groves et al., 2002; Das et al., 2006; Howard et al., 1998) and were specifically intended to avoid many of the problems associated with comprehensive species-specific planning and inventory. Defined by vascular plant assemblages (Groves et al., 2000), coarse filters are assumed to be representative of other taxa and to serve as a planning surrogate for poorly understood taxa such as invertebrates (Hunter, 1991, 2005). A strategy of conserving examples of all regional plant communities in reserves could conceivably protect the vast majority of all regional species, including regionally rare insects (Panzer and Schwartz, 1998; Shuey, 2005). Plant assemblages as coarse filter surrogates continue to be used widely despite identified limitations in intact landscapes where narrow definitions of plant assemblages would not necessarily conserve viable ecological landscapes (e.g., Noss, 1987a, b; Hunter, 1991). Despite their widespread use, coarse filter planning tools have not been widely assessed to determine if they are adequate surrogates for animal or insect communities (Su et al., 2004; Panzer & Schwartz, 1998; Shuey, 2005).

This research was initiated to test the applicability of botanically-based coarse filters relative to a large, species-rich group of phytophagous insects within a discrete landscape of habitat types on expansive sand deposits. Previous research focused on smaller animal groups or guilds such as butterflies, dung beetles, or birds at discrete sites (e.g., Su et al., 2004; Panzer & Schwartz, 1998; Davis et al., 2008; Barlow et al., 2007a). We choose a species rich assemblage of phytophagous insects, night flying moths (Lepidoptera) that provide a potential regional fauna exceeding 1000 species for our analysis (Metzler, unpub, data). Our sample sites were chosen to represent points across the local ecological and disturbance gradients that define natural and anthropogenic sand communities in the region. Before agricultural conversion, the region was a complex mosaic of vegetation that reflected the interplay between hydrology and fire frequency over sandy soils. Xeric dune rises supported oak barrens and woodlands while wet and mesic sites supported open sand prairie. Areas with more frequent fire had lower density of woody plant dominance than did areas with infrequent fire. Our sample sites represent points along these two continua in the present day landscape.

Specifically, we assessed vascular plant species diversity and moth species diversity across a series of sites representing high-quality sand prairie and sand barrens ecosystem remnants as well as their local degradation endpoints (old field and oak woodland/forest, respectively) to better understand the relationships between plant and moth communities in the region. Our goal was to determine if recognizable plant assemblages (sensu Faber-Langendoen, 2001) perform well as planning surrogates for a species rich assemblage of phytophagous insects as determined by their statistical congruence of alpha and beta diversity measures (alpha measures of species richness in an area while beta compares species composition between areas). If confirmed, this congruence could be further developed as a tool to assess conservation impacts of habitat management strategies, especially ecological restoration and re-creation, and could provide an independent conservation assessment tool beyond the successful establishment or manipulation of vegetation.

METHODS

The research was conducted in the Kankakee Sands Natural Region of northwestern Indiana (Fig. 1), a region originally dominated by an expansive mosaic of herbaceous wetland communities, sand prairie, and oak barrens (Homoya et al., 1985). These systems developed on glacial outwash sand deposits that undulated over a near surface water table creating a complex mosaic of wetland and upland habitats. Anthropogenic wildfire maintained the communities as open to sparsely treed. Today, the ecosystems and habitats of the region are highly fragmented from agricultural conversion and hydrologic manipulation (O'Leary & Shuey, 2003). Remnants of the natural ecosystems are still fairly common in the area. Most of these remnants are scattered black oak barrens communities (40%-60% canopy closure) and more commonly, their fire-suppressed degradation endpoints--oak woodlands (90%-100% canopy closure), which persist on well-drained sand deposits of limited agricultural value. The rarest community types, wet and mesic sand prairie, also survive as scattered and isolated fragments in the region as well. Collectively these assorted ecosystem remnants represent one of the most significant collections of sand prairie and oak barrens habitats surviving east of the Mississippi River (TNC, 2008).

We sampled a series of sites (Fig. 1) within two local, largely native landscapes exceeding 5000 ha. The sample sites were chosen to represent high-quality ecosystem remnants and regional degradation endpoints. Sites ranged in size from 15 ha to 100 ha of contiguous, similar habitat. High-quality sites included mesic sand prairie (two sites) and oak barrens (six sites, canopy closure < 50%). Sites representing degradation pathways for these communities included exotic-dominated old fields that developed from abandoned agricultural fields (two sites), one old field that succeeded to native vegetation, and two sites representing fire-suppressed, advanced succession oak woodlands (canopy closure 90100%). The sample sites were clustered within two complexes of managed conservation lands separated by approximately 40 kin. Although we were limited by habitat availability within each complex, we tried to intermingle sample site types geographically (Fig. 1). Within each complex, individual sample sites were located no closer than 0.9 km to one another.

Moth samples were collected once monthly from May to Oct. in 1999 and 2000 from each site. Moths were sampled using 15 watt blacklight traps adapted from standard USDA (United States Department of Agriculture) design (see Thomas, 1996). A photocell was used to turn the traps on and off at sunset and sunrise. Insects are attracted to the ultraviolet light, and baffles direct the insects downward though a funnel in the trap where they are dispatched with a cyanide-based killing agent to minimize sample damage. Samples were retrieved at dawn and all moths placed in cold storage until taxonomic processing. Species were identified using wing pattern characters where possible, but a substantial number of species required genetalic dissection for determination to species. Numbers of individuals per species were tallied and recorded in a data base by sampling event.

Plant communities were characterized by simple species inventory assessments within a circle plot of 50 in radius (0.85 ha) centered on the moth trap. Species inventories were conducted within each circle plot plot in May, Jul., and Aug. 2000, capturing the full seasonal phenological progression at the sites. We did not quantitatively assess plant communities.

Data were analyzed to provide insights into patterns of community integrity, diversity, and relationships among the sites. Community integrity was assessed using the floristic quality assessment protocols developed by Swink and Wilhelm (1994) for the Chicago Region, which includes the study area. The floristic quality assessment index (FQAI) is an evaluation procedure that uses measures of ecological conservatism, expressed numerically as a coefficient of conservatism or C value, and richness of the native plant community to derive a score (I) that is an estimate of habitat quality. C values range from 0 to 10 and represent an estimated probability that a plant is likely to occur in a landscape relatively unaltered from what is believed to be a pre-settlement condition. A C of 0 is given to plants that have little fidelity to any remnant natural community and may be found almost anywhere. Similarly, a C of 10 is applied to plants that are almost always restricted to a pre-settlement ecosystem remnant. C values for all vascular plants known from the study area follow Swink and Wilhelm (1994) and the sample sites fall within the geographic scope of their research. To provide a traditional measure of diversity, we calculated Shannon H' diversity indices for the quantitative moth samples. We used regression analysis to determine if plant species numbers or these derived measures of community quality would predict moth community richness or H'.

[FIGURE 1 OMITTED]

We produced cluster analysis dendrograms to illustrate relationships among sample sites following methods in Henderson and Seaby (2004). Plant and moth data were analyzed separately using additive similarity trees based on Bray-Curtis community coefficients of dissimilarity based on species presence-absence for plants and moths as well as absolute abundance for moths to determine how sample sites related to one another. The BrayCurtis dissimilarity ranges between 0 and 1, where 0 means the two sites have the same composition, and 1 means the two sites do not share any species. We used Ward's clustering protocols to compute the distance between groups. At each iteration, all possible pairs of groups are compared and the two groups chosen for fusion are those which will produce a group with the lowest variance. The resulting groups are as homogeneous as possible.

We used the Analysis of Similarity Statistic (Clarke, 1993) to analyze the significance of the resulting relationships relative to our pre-analysis assignment to habitat type: sand prairie, old field (including dominance by both non-native and native species), oak barrens, and fire suppressed oak woodlands. To test for significance the ranked similarity within and between groups is compared with the similarity that would be generated by random chance. The samples are randomly assigned to groups 1000 times and R calculated for each permutation. The observed value of R is then compared against the random distribution to determine if it is significantly different from that which could occur at random.

We used Mantel tests to test for significance of congruence in community similarity between all pair-wise comparisons between plant and moth and communities. The Mantel test is a statistical test of the correlation between two similarity matrices. Mantel tests were performed using PopTools software (Hood, 2006) with 999 random permutations.

RESULTS

PLANT AND MOTH COMMUNITIES

A total of 295 species of native vascular plants (range = 41-91) were encountered during this study (Table 1). Prairie and barrens ecosystem remnants, including the fire suppressed barrens, supported diverse native plants communities as did the agricultural field that succeeded to native species. A total of 41 non-native vascular plants were encountered in the sites (range = 2-21) (Table 2). Non-native plants comprised between 11% and 46% of the species present in the old fields that had previously been used for agricultural production. In native ecosystem remnants not directly impacted by agricultural production, non-native plant species composed between 3% and 7% of the species present.

All native-dominated sample plots, including two degradation end points (native old field and oak woodland) had average C values near or above the threshold of 4.5 level suggested as an indicator of high natural habitat potential in the study region (Swink and Wilhelm, 1994) (Table 2). These plots have FQAI scores that range between 30 and 55. FQAI values above 35 possess sufficient conservatism and richness to be of regional significance, and areas registering above 50 are considered very high quality (Swink and Wilhelm, 1994).

In contrast the two exotic old fields were characterized by low mean C values and FQAI values indicative of highly disturbed, weed-dominated sites. Non-native species represented 32% and 46% of the species present in these sites, and although not quantified, were visually dominant. Both these fields were removed from row crop agriculture 4 y prior to sampling and were dominated by typical agricultural weeds and invasive species. Both sites are surrounded by active agricultural fields and ditches which were the primary sources for recolonization. The native old field is surrounded by oak woodland that likely served as diverse seed sources when it was removed from agricultural production approximately 20 y prior to the study. During the intervening time, a diverse assemblage of conservative native species has established on the site with an FQAI value that exceeds both of our mesic sand prairie plots (Table 1).

A total of 26,662 moths were determined to species, representing 642 species (Table 1). The number of species per site ranged between 122 and 342 (Table 2). The number of species recorded at each site is strongly correlated with the number of individuals sampled (P < 0.01, R-Sq = 89.3%).

SIMPLE PLANT-MOTH COMMUNITY RELATIONSHIPS

There is no correlation between the number moth species and native plant species (P = 0.19, R-Sq = 15.0%), invasive plant species (P = 0.20, R-Sq = 14.7%), or total plant species (P = 0.29, R-Sq = 10.0%) at the sample sites. Nor is there a relationship between the number of moths recorded and the mean C value (P = 0.21, R-Sq = 13.9%) or the FQAI score (P = 0.16, R-Sq = 16.8%) at the sample sites. Moth community Shannon H' scores were not predicted by either the mean C value (P = 0.56, R-Sq = 3.2%) or the FQAI score (P = 0.33, R-Sq = 8.5%).

COMMUNITY SIMILARITY ANALYSES AND CONGRUENCE

Dendrograms from Bray-Curtis dissimilarity analyses of plant and moth communities at the sample sites are illustrated in Figure 2. Analysis of Similarity Statistics indicates that our pre-analysis assignment to habitat types (mesic sand prairie, old field, oak barrens, oak woodland) are meaningful for both plants and moths, and samples within groups are more similar in composition than samples from different groups (plant species presence P = 0.006, moth species presence P = 0.004, moth species abundance P = 0.008).

[FIGURE 2 OMITTED]

The dendrograms for plants and moths produced two contrasting sets of community relationships. For plant species, there is no intuitive relationship between our pre-assigned habitat types and the resulting dendrogram. Other than the close similarity of two adjacent prairie sites and the two exotic old fields, there are no easily interpreted relationships.

In contrast, both analyses of moths (presence only and abundance by species) produced well ordered dendrograms with a basic dichotomy between relatively treeless habitats (prairie and old fields) and treed habitats (barrens and oak woodlands). Within the relatively treeless group, the native old field clustered strongly with native prairie, while the exotic old fields formed a separate cluster. It is worth noting that woody trees and shrubs that visually dominate savanna and woodlands (species of Quercus, Salix, Sassafras, Rhus, Populus, and Prunus) were reported in all habitat types. For treed habitats, there was no segregation between open oak barrens and closed canopy oak woodlands.

[FIGURE 3 OMITTED]

Mantel tests indicated that community similarity between plants and moths (with both species abundance and presence only) was significantly positively correlated (P < 0.05; Fig. 3). Sites that were more similar based on plant species composition, also supported similar moth communities.

DISCUSSION

Although intuitively apparent, the basic assumption that underlies the "coarse filter" conservation strategy has not been widely tested to determine if speciose animal assemblages respond to environmental gradients as expressed via plant communities. Those studies that have examined this have focused primarily on species richness measures or groups that might serve as indicator taxa and have generated inconsistent results (e.g., Kati, 2004; Barlow et al., 2007b). For example, Howard et al. (1998) reported little congruence in species richness between woody plants, butterflies, large moths, birds, and mammals in Ugandan forests remnants. Oertli et al. (2005) report no relationship between bees, grasshoppers, and aculeate wasps, in a mosaic landscape in the Swiss Alps. Similarly, our assessments of alpha diversity show no relationships between simple measures of plant and moth communities. In contrast to most findings, Panzer and Schwartz (1998) found that for conservative insect species (a subset of the total fauna representing regionally rare species only associated with high-quality prairie and savanna habitats), 80-85% of the variance in richness among sites could be explained by native plant species richness when coupled with area (Panzer et al., 1995).

We concur with Su et al. (2004) who argue convincingly that conservation planning tools and assessments focused on [alpha]-diversity miss the point. Divergent taxa respond differentially to environmental gradients and relationships between measures of species richness in divergent taxonomic groups may be spurious. Simple measures of [alpha]-diversity likely provide few if any useful insights for conservation planning across divergent taxonomic communities in our system.

Measures of community integrity (coefficient of conservatism and floristic quality index) have also been used as surrogates to assess the conservation value of sites in the study region (Swink and Wilhelm, 1994). These assessment tools are increasingly being adapted for use across much of the Midwest to prioritize conservation actions and to evaluate the conservation value of management and restoration (Andreas and Lichvar, 1995; Bernthal, 2003; Herman et al., 1997; Lopez and Fennessy, 2002; Rooney and Rogers, 2002). Although these measures provide insight into plant assemblages and the presence of regionally rare species, they do not necessarily predict [alpha]-diversity of insects associated with sites. In an explicit test of this relationship, Panzer and Schwartz (1998) found no relationship between botanical measures of community integrity and s-diversity of conservative insects across the Chicago region. Our results support this, and across our ecological mosaic, and we find no correlation between measures of community integrity and the moth species richness or Shannon H'. Increased botanical integrity does not translate directly to measures of moth species richness or diversity.

Su et al. (2004) argue that [beta]-diversity is relevant to conservation planning and suggest that patterns of community similarity are more appropriate assessments of congruence between communities. Using data for plants, birds and butterflies in montane meadow habitats, they found that sites that supported similar communities of plants also supported similar communities of birds and butterflies. In the context of coarse-filter conservation, congruence between community similarity measures of divergent taxa indicate that at the community level, they are indeed responding to the same environmental gradients (but not necessarily responding in the same way). Our study was designed explicitly to assess cross-taxon congruence of community similarity relationships ([beta]-diversity) relative to other more common [alpha]-diversity measures of surrogate relevance in closely related botanical communities across subtle ecological gradients. Our results indicate that moth communities are highly congruent with plant communities within the study region--sites with similar plant assemblages also support moth communities that are similar as well. Given that moth species generally have very specific hostplant requirements this relationship makes sense. As hostplant species turn over based on community type, the individual moth species they support turn over as well.

In order to successfully implement successful insect conservation programs that reach beyond a handful of representative taxa and or imperiled species, we clearly need planning surrogates that can be used in lieu of assessments of entire insect communities--not more surrogates (sensu Kremen et al., 1993; Ricketts et al., 2002). We suggest that defined vascular plant assemblages (communities) (Faber-Langendoen, 2001) are a strong candidate for an insect-inclusive surrogate. Plant communities and ecological systems are generally defined and used across most of the Western Hemisphere (Josse et al., 2003, 2007; Grossman et al., 1998; Comer et al., 2003). Our work and that of others (Oertli et al., 2005; Summerville and Crist, 2004, 2008) demonstrates that some insect communities respond to the same ecological gradients as those that shape plant communities. What is lacking, are the assessments such as Fraser et al. (2009) that will determine if other insect taxa, especially non-phytophagous insects, respond similarly. No surrogate planning tool will be representative of all biodiversity in a region. But understanding both the strengths and shortcomings of surrogate tools that are used widely will be critical if we, as a society are serious about conserving the broadest sampling of biodiversity as possible.

Acknowledgments.--We thank the Indiana Department of Natural Resources for access to lands they manage. This research grew out of long conversations with Ron Panzer, Northeastern Illinois University, on the challenges facing insect conservation. Diane Debinski, Iowa State University, provided useful comments on an earlier draft of this manuscript and we are appreciative. Funding for this research was provided by TNC's Hoosier Science Fund, supported by private donors.

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SUBMITTED 8 MARCH 2011

ACCEPTED 16 NOVEMBER 2011

JOHN A. SHUEY

The Nature Conservancy, 620 E Ohio Street, Indianapolis, Indiana 46202

ERIC H. METZLER

Adjunct Curator of Lepidoptera, Michigan State University, East Lansing, Michigan 48824 and National Museum of Natural History Research Collaborator, P.O. Box 45, Alamogordo, New Mexico 88311

AND

KEVIN TUNGESVICK

Spence Restoration Nursery, 2220 East Fuson Road, Muncie, Indiana 47302

Corresponding author: e-mail: jshuey@tnc.org
TABLE 1.-Total species encountered and the range of site-
specific variation for plant and moth  species sampled in
sand prairies and barrens habitats

          Total     Maximum   Minimum   Mean    SD
          species

Plants      295        97        41       62   17.3
Moths       642       342       122    212.3   65.8

TABLE 2.--Breakdown by sampling site for plant and moth
communities. See text for explanation of mean C value and
Floristic Quality Index

                                             Oak       Oak       Oak
                         Praire   Praire   Barrens   Barrens   Barrens
                           1        2         1         2         3
Vascular Plants
# of native species        52       51       66        67        91
# of exotic species        3        3         2         3         7
mean C value              4.3      4.4       5.1       4.7       4.9
Floristic Quality         30.7     31.6     41.1      38.1      46.4
 Index (FQI)
Moths
# of species              221      136       314       22S       205
* moths collected         1360     807      4721      1702      2123
Shannon H (1)             4.43     3.67     4.40      4.42      4.43

                           Oak       Oak       Oak       Oak
                         Barrens   Barrens   Barrens   Woodland
                            4         5         6         1
Vascular Plants
# of native species        89        63        59         50
# of exotic species         6         5         2         3
mean C value               5.9       5.2       5.0       4.4
Floristic Quality         55.3      41.1       380       31.2
 Index (FQI)
Moths
# of species               237       219       246       342
* moths collected         2481      1896      2613       4433
Shannon H (1)             4.32      4.20      4.05       4.37

                                    Exotic   Exotic
                           Oak       Old      Old     Native
                         Woodland   Field    Field     Old
                            2         1        2      Field
Vascular Plants
# of native species         54        28       25       41
# of exotic species         4         13       21       5
mean C value               4.4       2.0      1.1      5.0
Floristic Quality          32.7      10.7     5.3      31.9
 Index (FQI)
Moths
# of species               165       122      187      137
* moths collected          1482      660      1698     686
Shannon H (1)              3.55      3.97     4.17     4.03
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