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New Eastern Geographic Distribution Records of Reithrodontomys fulvescens (Fulvous Harvest Mouse)
Angela L. Larsen, Kristy L. King, Jessica A. Homyack, T. Bently Wigley, Darren A. Miller, and Matina C. Kalcounis-Rueppell

Southeastern Naturalist, Volume 17, Issue 4 (2018): N77–N83

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N77 2018 Southeastern Naturalist Notes Vol. 17, No. 4 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell New Eastern Geographic Distribution Records of Reithrodontomys fulvescens (Fulvous Harvest Mouse) Angela L. Larsen1,*, Kristy L. King1, Jessica A. Homyack2, T. Bently Wigley3, Darren A. Miller4, and Matina C. Kalcounis-Rueppell1 Abstract - The current documented Reithrodontomys fulvescens (Fulvous Harvest Mouse) distribution includes western Mississippi. Here, we report new geographic distribution records in east-central Mississippi. We conducted live-trapping in Kemper County, MS, during the summers of 2011–2015, and captured both Fulvous Harvest Mice and Reithrodontomys humulis (Eastern Harvest Mouse). We compared measurements from captured Fulvous and Eastern Harvest Mice to verify presence of both species. Our results showed 2 separate clusters of measurements, independent of our species assignment in the field, and the clusters were significantly different (P < 0.01). It is unclear whether these new records represent a range expansion or new information about the species’ historical distribution. Reithrodontomys fulvescens J.A. Allen (Fulvous Harvest Mouse) is distributed throughout most of Mexico, Louisiana, Arkansas, Oklahoma, Texas, and parts of southern New Mexico, southwest Missouri, southeast Kansas, and western Mississippi (Hooper 1952, Spencer and Cameron 1982). There have been reports of a northward expansion of the Fulvous Harvest Mouse (Baccus 1968, Findley and Pullen 1958, Long 1965). However, to our knowledge, there has been no compilation of records to suggest an eastward expansion of the Fulvous Harvest Mouse distribution. There are records of Fulvous Harvest Mouse occuring east of the documented distribution in central Mississippi, including specimens collected in central Mississippi (Scott County) by M.C. Thompson in 1959 (GBIF 2017) and in 2002–2006 in southeastern Mississippi (George, Lamar, and Perry counties; Hanberry et al. 2013). Similarly, our research reported capturing Fulvous Harvest Mice in east-central Mississippi (Kemper County) in 2011–2015 (King et al. 2014; Larsen et al. 2016; Fig. 1). Our study area is closest to where the specimens were collected in Scott County, which is ~118 km from our study area. Here, we include detailed information of Fulvous Harvest Mouse captures from 2 of our previous studies (King et al. 2014, Larsen et al. 2016) to provide evidence of a possible eastward expansion. King et al. (2014) conducted live-trapping in the summer of 2011 and 2012 in Pinus taeda L. (Loblolly Pine) stands planted in 2005 and 2010, Loblolly Pine stands intercropped with Panicum virgatum L. (Switchgrass) planted in 2011 and 2012, and monocultures of Switchgrass. Larsen et al. (2016) conducted live-trapping in the summers of 2013–2015 in Loblolly Pine stands planted in 2010, Loblolly Pine stands intercropped with Switchgrass planted in 2011 and 2012, and monocultures of Switchgrass. From these 2 live-trapping studies conducted in Kemper County, MS, we report new records of the Fulvous Harvest Mouse for east-central Mississippi. To our knowledge, these are the most eastward records of the Fulvous Harvest Mouse. Our studies used live trapping and, because we had no trap mortality associated with Fulvous Harvest Mouse captures, we did not collect voucher 1University of North Carolina at Greensboro, 312 Eberhart Building, 321 McIver Street, Greensboro, NC 27402. 2Weyerhaeuser Company, 505 North Pearl Street, Centralia, WA 98531. 3National Council for Air and Stream Improvement, Inc., PO Box 340317, Clemson, SC 29634. 4National Council for Air and Stream Improvement, Inc., PO Box 9681, Mississippi State, MS 39762. *Corresponding author - angelallarsen@gmail.com. Manuscript Editor: David Scott Jachowski Notes of the Southeastern Naturalist, Issue 17/4, 2018 2018 Southeastern Naturalist Notes Vol. 17, No. 4 N78 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell specimens of Fulvous Harvest Mice. However, we measured hindfoot length (mm), mass (g), tail length (mm), body length (mm), and head length (mm), and we took voucher photographs of all captured individuals. We conducted our field methods under annual scientific collection permits from the Mississippi Department of Wildlife, Fisheries, and Parks and UNCG IACUC protocols 11-03 and 14-001.0. We followed the American Society of Mammalogists guidelines for use of wild mammals in research (Sikes and Gannon 2011). We captured 61 (6 captured only as juveniles, 55 adults) unique Fulvous Harvest Mice (2011: 9 [1 female, 8 males]; 2012: 25 [6 females, 19 males]; 2013: 0; 2014: 6 [1 female, 5 males]; and 2015: 21 [7 females, 14 males]) 94 times. We also captured specimens of the only other Reithrodontomys spp. in the area, Reithrodontomys humulis Audubon and Bachman (Eastern Harvest Mouse)—36 (3 captured only as juveniles, 33 adults) unique individuals (2011: 1 [0 females, 1 male]; 2012: 13 [8 females, 5 males); 2013: 2 [1 female, 1 male]; 2014: 7 [3 females, 4 males]; and 2015: 13 [4 females, 9 males]) 50 times. We captured individual Eastern Harvest Mice in each treatment type throughout our site (22 in Switchgrass monocultures, 5 in young Loblolly Pine stands, 4 in old Loblolly Pine stands, 3 in old intercropped stands, and 2 in old Loblolly Pine stands). However, Fulvous Harvest Mice were only captured in the young plots (50 in Loblolly Pine stands, 10 in intercropped stands, and 2 in Switchgrass monocultures). Furthermore, there were only 2 trap stations where both species were captured and only 1 at which both species were captured in the same year (in a Switchgrass monoculture). Our data suggest that although Eastern and Fulvous Harvest Mice can co-occur, there may be spatial partitioning and differences in Figure 1. Map of Mississippi depicting Reithrodontomys fulvescens (Fulvous Harvest Mouse) specimen records obtained from Global Biodiversity Information Facility (GBIF 2017; white circles) and our study area in Kemper County (black circle) where we captured Fulvous Harvest Mice outside of their previously known distribution. N79 2018 Southeastern Naturalist Notes Vol. 17, No. 4 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell habitat preference between the species. We report the coordinates of the trapping-grid centers where we captured Fulvous Harvest Mice in Table 1. We compared measurements (hindfoot length [mm), mass [g], tail length [mm], body length [mm], and head length [mm]) between Fulvous and Eastern Harvest Mice (Table 2). The 2 species differ in size and pelage color; the Eastern Harvest Mouse is smaller and more brown than the Fulvous Harvest Mouse, which has a more tawny, or fulvous, pelage (Fig. 2; Stalling 1997). There is evidence that Reithrodontomys megalotis Baird (Western Harvest Mouse) could be expanding its distribution due to increasing conversion of forests to agricultural lands, which suits Western Harvest Mice’s preference for grassy, weedy habitat, including fallow fields and fence rows (Choate et al. 1994, Graves 2016). However, east-central Arkansas is the far eastern edge of its known distribution. Furthermore, Western Harvest Mice have a shorter tail length (55–75 mm) than Fulvous Harvest Mice (see below; Choate et al. 1994). The only other species at our study site that has a tawny pelage is Ochrotomys nuttalli Harlan (Golden Mouse) (Fig. 2). However, Golden Mice tend to be larger overall with longer bodies, shorter tails, and greater mass (total length = 140–190 mm, length of tail = 67–97 mm, and mass = 18-27 g; Kays and Wilson 2009) compared to Fulvous Harvest Mice, which usually have tails twice as long as their bodies (see below and Fig. 2). The Eastern Harvest Mouse has the following characteristics reported by Kays and Wilson (2009) and Choate et al. (1994), respectively: total length = 107–128 mm and 110–130 mm; length of tail = 45–60 mm and 45–60 mm; length of hindfoot = 15–17 mm and 15–18 mm; and mass = 10–15 g and 6–10 g. The Fulvous Harvest Mouse has the following characteristics reported by Kays and Wilson Table 2. Morphological measurements of Reithrodontomys fulvescens (Fulvous Harvest Mouse, n = 21) and Reithrodontomys humulis (Fulvous Harvest Mouse, n = 13) individuals with all metrics recorded during live-trapping during the summers of 2011–2015 at our study area in Kemper County, MS. Mean SE Min Max Fulvous Harvest Mouse Hind foot length (mm) 19.33 0.22 17 21 Mass (g) 12.29 0.58 7 18 Tail length (mm) 86.95 1.25 75 96 Body length (mm) 57.38 1.90 40 79 Head length (mm) 23.10 0.59 19 27 Eastern Harvest Mouse Hind foot (mm) 13.77 0.30 12 15 Mass (g) 9.54 0.69 7 16 Tail length (mm) 52.54 0.79 49 60 Body length (mm) 56.23 1.13 52 65 Head length (mm) 20.85 0.70 16 25 Table 1. Coordinates of 15 trapping grid centers where Reithrodontomys fulvescens (Fulvous Harvest Mice) were captured in Kemper County, MS, from 2011 to 2015. Long. = longitude, Lat. = latitude. Long. (°W) Lat. (°E) Long. (°W) Lat. (°E) Long. (°W) Lat. (°E) 88.58642 32.85852 88.52557 32.87356 88.50830 32.87343 88.58745 32.85114 88.52962 32.87188 88.51455 32.87343 88.59315 32.85164 88.53002 32.80739 88.51275 32.87014 88.53453 32.86194 88.52193 32.81097 88.51277 32.87012 88.52552 32.86907 88.51880 32.80788 88.54187 32.80092 2018 Southeastern Naturalist Notes Vol. 17, No. 4 N80 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell (2009) and Choate et al. (1994), respectively: total length = 134–189 mm and 150–175 mm; length of tail = 73–116 mm and 85–95 mm; length of hindfoot = 16–22 mm and 16–20 mm; and mass = 6–25 g and 10–12 g. Even though the span of measurement values differ between the 2 references, it is still clear that total length and tail length differentiate Eastern and Fulvous Harvest Mice. To confirm that our field identifications represented 2 species, we used measurements from our captures of Fulvous and Eastern Harvest Mice during 2011–2015 in a cluster analysis by partitioning around medoids (PAM), which is a robust version of k-means cluster analysis (Kaufman and Rousseeuw 1990). Although we identified each capture to species in the field, we ensured that our field identifications were not included in our clustering analysis to keep our statistical identifications independent of our field identifications. We used a clustering analysis to provide further support for stating there are 2 species, to compare how different combinations of commonly recorded morphometric measurements separate individuals into clusters, and to assess alignment of clusters with our species identifications. We used 2 separate datasets in our clustering analysis. We first used unique captures, then we used all observations (captures and recaptures) to account for variation in measurements among observers. We began by computing a gap statistic in Program R (function fviz_nbclust in package factoextra; Kassambara and Mundt 2017, R Core Team 2017) for each dataset to determine the optimal number of clusters. For each dataset, the gap statistic resulted in 2 clusters. We conducted PAM (function pamk in package fpc; Hennig 2015, Kaufman and Rousseeuw 1990, R Core Team 2017) on records with hindfoot length and mass, as we recorded both measurements frequently. Then, we used only records with hindfoot length, mass, and tail length. Next, we used only records with hindfoot length, mass, tail length, and body length. Finally, we used records with all measurements recorded: hindfoot length, mass, tail length, body length, and head length. For each dataset, we visualized our measurements from the 2 species using an ordination plot from our PAM clustering Figure 2. Photos of (a) 2 Reithrodontomys fulvescens (Fulvous Harvest Mice; tags K1244 and K1659); (b) a Reithrodontomys humulis (Eastern Harvest Mouse); and (c) an Ochrotomys nuttalli (Golden Mouse) captured at our study area in Kemper County, MS. In each case, 2 frames of each individual are shown. Eastern Harvest and Golden Mice have similar physical characteristics to Fulvous Harvest Mice, but we ensured proper identification via measurements and pelage color. Fulvous Harvest Mice have distinctly longer tails relative to their body length, and longer tails compared to Eastern Harvest and Golden Mice. Eastern Harvest Mice have gray pelage, Fulvous Harvest Mice have orange pelage, and Golden Mice have yellow pelage. All photos © K. King and have not been altered. N81 2018 Southeastern Naturalist Notes Vol. 17, No. 4 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell analyses (factoextra package; Kassambara and Mundt 2017). We used analysis of similarity (ANOSIM, vegan package; Oksanen et al. 2013) to test if the 2 clusters statistically differed. We accepted differences as statistically significant at the α = 0.05 level. We found the same overall results for both of our unique individuals and observations datasets. When we used the 2 most common measurements, hind foot length and mass, PAM misidentified 10 of 96 individuals and 10 of 136 observations. When we included tail length measurements, PAM misidentified 1 out of 77 individuals and 1 of 106 observations. The addition of body length resulted in 0 misidentified of 62 individuals and 84 observations. Finally, when we included all measurements, PAM again resulted in 0 misidentified out of 34 individuals and 49 observations. When we included sex in the analysis, there were no major differences. Female Eastern Harvest Mice (n = 3) clustered closer together than female Fulvous Harvest Mice (n = 7), but it is important to note our low sample size. For all datasets, clusters were significantly different from each other based on ANOSIM (P < 0.01 for all comparisons; Fig. 3). Specifically, for our dataset with all recorded measurements of individuals (n = 34), Fulvous and Eastern Harvest Mice groups were significantly different from each other (R = 0.97, P < 0.01). Thus, morphometric measurements differed between the 2 species we identified in the field and were 100% effective at classifying Eastern and Fulvous Harvest Mice. We conclude that the Fulvous Harvest Mouse occurs in east-central Mississippi. There are 2 possible explanations of why this new eastern distribution has not been previously reported. One scenario is that the Fulvous Harvest Mouse has historically occurred in east-central Mississippi, but may not have been sampled or may have been misidentified. For example, Clark and Durden (2002) captured small mammals throughout Mississippi, but only reported capturing Eastern Harvest Mice. Clark and Durden (2002) may not have Figure 3. Ordination plot of Reithrodontomys fulvescens (Fulvous Harvest Mouse; cluster 1, n = 21) and Reithrodontomys humulis (Eastern Harvest Mouse; cluster 2, n = 13) measurements taken at time of capture from unique individuals captured in Kemper County, MS, from 2011 to 2015. Hindfoot length, mass, tail length, body length, and head length were included, and each species clustered separately. 2018 Southeastern Naturalist Notes Vol. 17, No. 4 N82 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell sampled areas with Fulvous Harvest Mice, or they may have misidentified Fulvous as Eastern Harvest Mice if unaware of Fulvous Harvest Mouse presence in parts of Mississippi. In this case, our study provides records to extend the known Fulvous Harvest Mouse distribution range to include east-central Mississippi. Alternatively, the Fulvous Harvest Mouse may not have historically occurred in east-central Mississippi. In this case, our records show a Fulvous Harvest Mouse eastward range-expansion that may be concomitant with the northward expansion (Baccus 1968, Findley and Pullen 1958, Long 1965). Eastern Harvest Mice have been captured throughout Mississippi, including previous studies in the same county as our work (Hood et al. 2002). However, beyond the studies we mentioned above, we are not aware of other studies in eastern Mississippi that report capturing Fulvous Harvest Mice. Future research should determine whether or not the Fulvous Harvest Mouse has historically been in east-central Mississippi because range expansions are important in studies of community structure (Le Roux and McGeoch 2008), genetic changes (Excoffier et al. 2009), and hybridization and speciation (Bocxlaer et al. 2010) which can affect biodiversity in the expansion area. Acknowledgments. Our research was supported by the following funding sources: Weyerhaeuser Company, Catchlight Energy LLC, National Council for Air and Stream Improvement, Inc., US Environmental Protection Agency (EPA), American Society of Mammalogists, North Carolina Academy of Science, and University of North Carolina at Greensboro. Weyerhaeuser Company employees operated under a Statement of Work with CLE during this study. This publication was developed under STAR Fellowship Assistance Agreement no. FP-91763501-0 awarded by the US EPA. It has not been formally reviewed by EPA. The views expressed in this publication are solely those of the authors, and EPA does not endorse any products or commercial services mentioned in this publication. We truly appreciate all our field technicians; R. Kelble, K. Paolini, E. VanNatta, A. Andrews, D. Bradley, J. Franklin, M. Martin, R. Meyer, A. Matteson, K. Corson, E. Smerdon, A. Wagoner, A. Culpepper, and G. Mikel. We thank Ray Iglay for discussions on small mammals in this region. We appreciate the people who helped us with logistics at our field site: Mississippi State University, Steve Emerson, and the employees of the Scooba Weyerhaeuser office. We also thank the East Mississippi Sportsman Association and Charlie Studdard with the Kemper County Hunting Club for lodging. Literature Cited Baccus, J.T. 1968. Two noteworthy records of rodents from Baylor County. The Southwestern Naturalist 13:362–362. Bocxlaer, I.V., S.P. Loader, K. Roelants, S.D. Biju, M. Menegon, and F. Bossuyt. 2010. Gradual adaptation toward a range-expansion phenotype initiated the global radiation of toads. Science 327:679–82. Clark, K., and L. Durden. 2002. Parasitic arthropods of small mammals in Mississippi. Journal of Mammalogy 83:1039–1048. Choate, J.R., Jr, J.K. Jones, and C. Jones. 1994. Handbook of Mammals of the South-Central States. Louisiana State University Press, Baton Rogue, LA. 320 pp. Excoffier, L., M. Foll, and R.J. Petit. 2009. Genetic consequences of range expansions. Annual Review of Ecology, Evolution, and Systematics 40:481–501. Findley, J.S., and B.E. Pullen. 1958. The Fulvous Harvest Mouse in New Mexico. Journal of Mammalogy 39:306–307. Global Biodiversity Information Facility (GBIF). 2017. GBIF occurrence download. Availoable online at http//www.GBIF.org. Accessed November 2017. Graves, G.R., 2016. Geographic range of the Western Harvest Mouse (Reithrodontomys megalotis) in Arkansas. Journal of the Arkansas Academy of Science 70:268–270. Hanberry, B.B., S.L. Edwards, S. Demarais, and J. Jones. 2013. Small-mammal responses to intensively established pine plantations in coastal plain Mississippi. Southern Journal of Applied Forestry 37:53–58. N83 2018 Southeastern Naturalist Notes Vol. 17, No. 4 A.L. Larsen, K.L. King, J.A. Homyack, T.B. Wigley, D.A. Miller, and M.C. Kalcounis-Rueppell Hennig, C. 2015. Fpc: Flexible procedures for clustering (version 2.1-10). 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