Bats (Mammalia: Chiroptera) recorded from mist-net and bridge surveys in southern Mississippi.

We surveyed communities of bats in southern Mississippi using mist nets and searches of bridges. We captured 41 individuals representing five species of bats over 28 nights of trapping. Nycticetius humeralis was the species most frequently captured in our nets, followed by Lasiurus seminolus, Myotis austroriparius, Pipistrellus subflavus, and Lasiurus borealis. These species are representative of communities of forest-roosting bats native to the southeastern United States. However, all M austroriparius were captured exclusively at the entrances of caves. Seven of 99 bridges that we searched were occupied by Corynorhinus rafinesquii as day-roosts. These seven bridges were all made of concrete and had girders or channel beams along their undersides. Our data on C. rafinesquii are consistent with findings of other studies, which suggest that the construction style of bridges plays an important role in providing day-roosts for this species.

**********

Kennedy et al. (1974) and Jones and Carter (1989) listed all species of bats (Order Chiroptera) known to inhabit Mississippi in their reviews of this state's mammals. Although a few studies (e.g., White, 1960; LaVal, 1967; Jones and Suttkus, 1975; Middleton, 1976; Cliburn and Middleton, 1983; Miller, 2000; Welch et al., 2001) have documented observations or collections of bats from distinct locations in Mississippi, no investigators have as yet compared species inventoried among different habitats in this state. Furthermore, three species of bats, Lasiurus cinereus (hoary bat), Lasiurus intermedius (northern yellow bat), and Corynorhinus rafinesquii (Rafinesque's big-eared bat), are poorly represented in collections from Mississippi (see Kennedy et al., 1974). Whether the lack of records from Mississippi for these species reflect inadequate efforts of sampling or the bats' inherent rarity is not clear. Therefore, the objectives of this study were: 1. to conduct surveys of communities of bats in southern Missi ssippi; 2. to collect specimens of Lasiurus cinereus, Lasiurus intermedius, and Corynorhinus rafinesquii.

MATERIALS AND METHODS

We defined the study area as encompassing the portion of Mississippi south of Interstate 20, which crosses the entire state in an east-west direction. Frost et al. (1986) describe much of this region.as having been historically dominated by longleaf pine (Pinus palustris) savanna, while more mesic habitats were characterized by beech (Fagus grandifolia), oaks (e.g., Quercus nigra), and magnolias (Magnolia spp.). Tupelo (Nyssa spp.) and baldcypress (Taxodium distichum) occurred along larger streams and rivers. In recent years suppression of natural fire regimes and widespread conversion of native forests to loblolly pine (Pinus taeda) plantations have dramatically altered the landscape of the region (Frost et al., 1986).

Inland habitats that we sampled included upland hardwood-pine (Pinus spp.) forest, lowland hardwood-pine forest, two limestone caves surrounded by upland, mesic hardwood forest, and cypress swamp (Table 1). The two caves, Pitts and Eucutta, are among the largest caves in Mississippi (Knight et al., 1974). At coastal sites (i.e., Davis Bayou and Mississippi Sandhill Crane National Wildlife Refuge), we typically found marshes bordered by swampy forests of wax myrtle (Myrica cerifera), sweetbay (Magnolia virginiana), and pines. We also sampled one night on Horn Island, a barrier island approximately 10 km off the coast of Mississippi. The interior of this island, managed by the National Park Service, was dominated by scrub forests of slash pine (Pinus elliotti). A few marshy ponds comprised the only sources of fresh water. Because we sought to add to the known distribution (in Mississippi) of each of our "target" bats, we chose some sampling sites based on the presence of suitable habitat (based on records in li terature) for one or more of these three species, L. cinereus, L. intermedius, and C. rafinesquii.

From June 1999 through July 2000 we sampled at thirteen different sites in eight counties of southern Mississippi using standard mist net procedures for insectivorous bats (Kunz and Kurta, 1988). Although we visited most localities only once during the investigation, we trapped twice at two sites. We generally selected evenings when the temperature was not expected to fall below 18[degrees]C. We collected bats using nylon mist nets 2.6 m high, ranging from 2.6 m to 9 m long and attached at each end to a metal pole approximately 3 m tall. We placed one to four mist nets at each site; we standardized our efforts by defining a net-night as one mist net opened during one night of trapping. We usually trapped just above the surface of water sources to intercept bats as they foraged or came to drink. We also placed nets over dirt roads within forested habitat and across the entrances of the two limestone caves. We chose the latter as additional sampling sites for C. rafinesquii, based on records from caves (Jones, 1977; Best et al., 1992). We opened our nets approximately 15 minutes before civil sunset and trapped for a minimum of three hours unless capture rates declined to less than one bat per hour. We determined sex and species of captured bats. We determined relative age (juvenile or adult) by noting the degree of ossification of the epiphyseal caps on the phalanges of the fingers (Anthony, 1988). Our methods followed University of Southern Mississippi IACUC protocol # 204-004.

Because several investigators (e.g., Lance and Garrett, 1997, Hurst and Lacki, 1999) have reported very little success in capturing C. rafinesquii in mist nets, we incorporated surveys of potential roosting sites (i.e., bridges) into our study. Individuals of this species have been noted to congregate along the undersides of concrete bridges by day, particularly during warmer months (Lance and Garrett, 1997; Lance et al., 2001). From May 1999 through early September 1999 we surveyed the undersides of 84 bridges located within the study area. Most of these bridges were located in and nearby the DeSoto or Bienville National Forests (NF). We checked 15 additional bridges in the Homochitto NF in June 2000. Several bridges in the DeSoto NE found to be inhabited by C. rafinesquii during previous searches by other investigators (C. Potin, pers. comm.) were included in our survey. A survey consisted of one or both of the investigators walking beneath a bridge and looking for bats along the underside of the structure . When bats were encountered, we determined the species, counted number of individuals, and noted presence or absence of young. We maintained a minimum distance of about 10 feet from clusters of bats so as not to disturb females with pups. As a result, we may have counted fewer bats than were actually present on some occasions.

RESULTS

We captured 41 bats of five different species over 28 net-nights (Table 1). Nycticeius humeralis, the evening bat, was the species most frequently caught in our mist nets, accounting for 39% of all captures (n = 16). Other species that we captured were: Lasiurus seminolus, the seminole bat (n = 11,27% of captures); Myotis austroriparius, the southeastern myotis, (n = 8, 20%); Pipistrellus subflavus, the eastern pipistrelle, (n = 3, 7%); Lasiurus borealis, the red bat, (n = 3, 7%). Nycticeius humeralis and L. seminolus occurred together at four localities. We captured two species, P. subflavus and M. austroriparius, at the entrances of both Pitts and Eucutta caves. All of the M. austroriparius that we captured in our survey were trapped at these caves. We captured two species, N. humeralis and L. seminolus, while sampling at Horn Island. We found P. subfiavus at three different sites (including Pitts and Eucutta caves) and L. borealis at three sites. We did not capture any L. cinereus or L. intermedius. Sex r atios for all species captured during the present study were: N. humeralis, 1.3 males/female; L. seminolus, 0.6 male/female; M. austroriparius,. I male/1.7 females; L. borealis, 0 males/3 females; P. subflavus, 0.5 male/female. We captured juveniles of two species, L. seminolus and L. borealis.

Six of 84 bridges that we surveyed in 1999 were inhabited by C. rafinesquii (Table 2). Three of these bridges were located in the Chickasawhay District of the DeSoto NF within a 6 km-span of a single road. The remaining three bridges were located in the DeSoto District of the DeSoto NF. The number of big-eared bats roosting beneath a bridge ranged from one to an estimated 25 individuals. We were able to identify young under two bridges on 27 May 1999, at one bridge on 10 June 1999, and under one bridge on 18 June 1999 (Table 2). One of the 15 bridges from Homochitto NF that we checked on 7 June 2000 was occupied by a solitary, male C. rafinesquii (Table 2). We found a female P. subflavus nursing a pup under another bridge in Homochitto NF.

DISCUSSION

Most of the species that we captured in our survey are representative of communities of forest-dwelling bats occurring throughout much of the southeastern United States. Nycticeius humeralis was the most common species that we captured, both in terms of number of individuals and number of sites where it was found. We collected N. humeralis in pine and bottomland-hardwood forests, and in both inland and coastal sites. Lance and Garrett (1997) captured N. humeralis more frequently than any other species in pine forests of central Louisiana, as did Krishon et al. (1997) in coastal Georgia. On Horn Island, a colony of this species apparently roosted within a boathouse at the end of a dock, utilizing the space between the tin roof and one of its concrete support columns. This structure was located approximately 300 m from the pond where we captured seven individuals of N. humeralis. Roosts for N. humeralis on Horn Island might include numerous pine snags (see Menzel et al., 2000). This species of bat had not been previously recorded from this locality.

Lasiurus seminolus was the second most-frequently captured species in our study. This species was also second to N. humeralis as the most common bat found in lowland, pine-dominated forest in Louisiana (Lance and Garrett, 1997). Menzel et al. (1998) determined that L. seminolus roosted predominantly in pine trees, while L. borealis preferred hardwoods. We captured a single L. seminolus on Horn Island, where the only trees available as roosts were pines. This species of bat had not previously been reported from Horn Island. Two of the three sites where we collected L. borealis were bottomland-hardwood forests. Miller (2000), however, captured more red bats than any other species in pine plantations of central Mississippi. Lowery (1974) described L. borealis as common and widespread in Louisiana.

Our records of M. austroriparius from Eucutta and Pitts Caves in Wayne County are especially noteworthy because this species was not recorded by Middleton (1976) in his survey of cave-dwelling vertebrates in Mississippi, although this bat had been previously captured in Pitts Cave (LaVal, 1967). Over much of its range, M. austroriparius roosts in caves (Whitaker and Hamilton, 1998), but concern exists over the decline and degradation of such habitats (Humphrey and Gore, 1992). We caught only adult males from Eucutta Cave and adult females from Pitts Cave. In this species, male and female bats tend to roost separately while pups are being born and nursed, but this segregation ends by late summer (Barbour and Davis, 1969). Lance and Garrett's (1997) survey supported Lowery's (1974) assertion that this species is widespread but uncommon in Louisiana where caves are apparently not present. However, M. austroriparius also roosts in hollow trees in bottomland forests (Hofmann et al., 1999). Pipistrellus subflavus was the only bat documented by Cliburn and Middleton (1983) from the caves that they visited. Best et al. (1992) recorded P. subflavus from limestone caves in southern Alabama, usually during cooler months. This species was the only bat that we found both at and away from caves. We obtained one specimen by netting over a stream in a pine forest interspersed with deciduous species (e.g., Magnolia virginiana, Nyssa sylvatica) and have observed this species roosting underneath concrete bridges. Findings of Menzel et al. (1999) suggest that during summer in the southern United States, P. subflavus roosts among foliage of evergreen trees located within the understory of mixed oak-pine stands.

The lack of captures for Lasiurus cinereus and L. intermedius during our study may be further evidence of these species' rarity in Mississippi. Kennedy et al. (1974) listed only two records of L. cinereus from Mississippi (Madison and Oktibbeha Counties). Barbour and Davis (1969) categorized the species as widespread but rare in the eastern United States. Little is known of the species' preferences with regard to roosting habitat, though Lowery (1974) noted that most specimens in Louisiana came from areas dominated by pine forests. Compared to the frequency of its detection by ultrasonic equipment, L. cinereus was rarely captured by Hart et al. (1993) in Pennsylvania. Future surveys to determine presence of L. cinereus in Mississippi and elsewhere should utilize ultrasonic detectors along with mistnetting. Only two records of L. intermedius from Mississippi exist, these specimens coming from Hancock and Warren Counties (Kennedy et al., 1974). Specimens of L. intermedius are typically scarce in collections of bats from the southeastern United States (Whitaker and Hamilton, 1998). Barbour and Davis (1969) suggested that capturing L. intermedius via mist-netting is difficult, even in apparently suitable habitat. The distribution of L. intermedius closely follows that of Spanish moss (Tillandsia usneoides), a preferred roosting substrate (Barbour and Davis, 1969). At one of our coastal sites (Davis Bayou, Jackson County), we unsuccessfully attempted to locate resting individuals by probing clumps of Spanish moss using a handheld net. Other sites (e.g., wet pine savanna in Jackson County and a golf course in Forrest County) provided the open, pasture-like environments which L. intermedius favors (Barbour and Davis, 1969), but Spanish moss was scarce or not present. Efforts to identify areas in Mississippi containing both ideal foraging habitat and roosting sites for L. intermedius as described by Jennings (1958) and Krishon et al. (1997) should continue.

Another species of bat native to the study area that we did not capture was Myotis lucifugus. Richmond (1968) reported a specimen from Horn Island, but did not elaborate on the circumstances of its collection. Myotis lucifugus is largely absent from the Gulf Coast, and this record would represent a substantial extension of the species' range (see Whitaker and Hamilton, 1998). Given this bat's superficial resemblance to N. humeralis and the tendency of N. humeralis to roost in buildings (a situation where bats might come into contact with an untrained observer), we suspect that this record of M. lucifugus was actually a misidentified N. humeralis. LaVal (1967) collected a specimen of M. lucifugus from Pitts Cave, but this species was absent from Middleton's s (1976) survey and has not been observed during subsequent visits to this and nearby caves (A. W. Trousdale, personal observations).

Corynorhinus rafinesquii historically ranged across Mississippi (Jones and Carter, 1989); however, local patterns of distribution for the species are poorly known throughout its range (Jones, 1977). Lance and Garrett (1997) captured only one specimen of C. rafinesquii during their extensive mistnetting efforts in Louisiana. Bridge surveys, which we adopted for our study, provided a much better method of finding big-eared bats. Of the four colonies that we located, the two observed on 27 May 1999 (in Perry County) contained the youngest pups based on their relative size, pelage, and inability to fly. Considering that C. rafinesquii are able to fly at three weeks of age (Jones, 1977), those pups would not likely have been born before early May, which is consistent with observations from Louisiana (Lowery, 1974).

All of the bridges occupied by C. rafinesquii were made of concrete and possessed some type of structure (i.e., rectangular compartments or girders) along their undersides. Similar preferences in bridge selection by this species were noted by McDonnell and Clark (1999) in North Carolina and by Lance et al. (2001) in Louisiana. We found C. rafinesquii under two bridges in Perry County that had been used by the species during previous years (C. Potin, pers. comm). In early August 1999 we returned to a bridge in Stone County under which we had observed a C. rafinesquii colony two months earlier and found a solitary individual. Lance (pers. comm.) located the species under this same bridge in 1997. If C. rafinesquii shows fidelity to certain bridges, this behavior may have important consequences for management of the species, considering that in many areas older bridges (which typically feature compartments or girders) are increasingly being replaced by slab bridges (Lance et al., 2001).

Table 1

Locations in southern Mississippi visited by authors while conducting
mist-net surveys for bats from July 1999 through July 2000.


Location (county)      Habitat (4)  Date(s) visited

Van Hook Golf               A       July 1999
  Course (Forrest)
Rails-to-Trails             A       July 1999
  (Lamar)
Bluff Creek at Bluff        B       3 August 1999
  Creek Rd (Stone)


Pitts' Cave (Wayne)         C       4 August 1999

Horn Island, GINS (1)       D       12 August 1999
  (Jackson)
Davis Bayou, GINS (1)       B       14 August 1999,
  (Jackson)                         13 October 1999
Beaver pond adjacent        E       17 August 1999,
  Cabin Rd,                         19 August 1999
  SCCNWR (2)
  (Adams)
Gillirad Lake, N            F       18 August 1999
  edge, SCCNWR (2)
  (Wilkinson)
Tiger Creek at Hwy          B       2 September 1999
  15 (Jones)
Eucutta Cave                C       16 September 1999
  (Wayne)
MSSCNWR (3)                 G       September 1999
  (Jackson)
Holliman Property           B       6 April 2000
  (Lamar)
Spector Farm (Pearl         B       1 July 2000
  River)

                       Species (# individuals)
Location (county)      collected

Van Hook Golf          None
  Course (Forrest)
Rails-to-Trails        None
  (Lamar)
Bluff Creek at Bluff   Lasiurus borealis * (1),
  Creek Rd (Stone)     L. seminolus * (7),
                       Nycticeius humeralis * (2),
                       Pipistrel subflavus * (1)
Pitts' Cave (Wayne)    Myotis austroriparius (5),
                       P. subflavus (1)
Horn Island, GINS (1)  L. seminolus * (1), N.
  (Jackson)            humeralis * (7)
Davis Bayou, GINS (1)  L. seminolus * (2),
  (Jackson)            N. humeralis * (6)
Beaver pond adjacent   L. borealis (1),
  Cabin Rd,            N. humeralis (1)
  SCCNWR (2)
  (Adams)
Gillirad Lake, N       L. borealis (1),
  edge, SCCNWR (2)     L. seminolus (1)
  (Wilkinson)
Tiger Creek at Hwy     None
  15 (Jones)
Eucutta Cave           M. austroriparius (3),
  (Wayne)              P. subflavus (1)
MSSCNWR (3)            None
  (Jackson)
Holliman Property      None
  (Lamar)
Spector Farm (Pearl    None
  River)

(1)Gulf Islands National Seashore;

(2)St. Catherine's Creek National Wildlife Refuge;

(3)Mississippi Sandhill Crane National Wildlife Refuge

(4)Key to categories of habitat: A = upland mixed hardwood-pine forest;
B=lowland mesic hardwood-pine forest; C=upland mesic hardwood forest;
D=coastal marsh; E=bottomland hardwood forest; F=cypress swamp; G=wet
pine savanna

* New county record for a species
Table 2

List of counties in southern Mississippi in which the authors observed
bridges occupied by Corynorhinus rafinesquii. (1)

                             # bridges used  # bats seen per
Location   Date(s) Surveyed  by C. RAF. (2)      bridge

Jones      10 June 1999,           2               12
           2 September 1999                         3
Perry      27 May 1999             2             11, 25
Stone      18 June 1999            1                9
Wayne      2 September 1999        1                1
Wilkinson  7 June 2000             1                1

           Pups observed
Location   under bridge?

Jones           Yes
                No
Perry        Yes, Yes
Stone           Yes
Wayne           No
Wilkinson       No

(1)Exact locations of bridges may be obtained from the authors.

(2)Corynorhinus rafinesquii

ACKNOWLEDGMENTS

The authors wish to acknowledge the Mississippi Wildlife Heritage Program of the Mississippi Museum of Natural Science for providing funding for this project. The U.S. Fish and Wildlife Service, the U.S. Forest Service, and the National Park Service provided access to their lands, as well as technical support. J.G. Himes and V. Shervette assisted in data collection.

D.A. Miller provided valuable technical instruction. R.F. Lance and C. Potin shared their data from bridge surveys. M.L. Kennedy assisted with identification of Myotis specimens. S.W Rosso gave advice, encouragement, and logistical assistance.

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Austin W. Trousdale (1)

(1.) Author for correspondence. Department of Biological Sciences, Post Office Box 5018