Linking Transportation and Conservation

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The significance of roads has expanded beyond pavement

A 'bats in bridges' report by RD Wildlife Management

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Linking Conservation and Transportation: A 'Bats in Bridges' Report Task No. 5372-33

A report for New Mexico Department of Transportation

April 2013

Prepared by RD Wildlife Management Smith, Holly J. Stevenson, Justin S.

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PREFACE

Bats are fascinating members of New Mexico’s natural environment; diminutive, captivating, and the only mammals capable of true flight. Bats are intrinsic to healthy ecosystems, community integrity and vital ecological processes. They provide valuable ecosystem services (insect suppression, pollination, seed dispersal), products and provisions (tequila, sisal, cactus fruits), cultural benefits (educational, recreational, spiritual) and contribute considerably to New Mexico’s mammalian diversity. Without these marvelous flying creatures, “benefits that humans inadvertently and unsuspectingly derive from bats will be forever lost or severely diminished, causing both known and unknown consequences to the ecosystems in which they have evolved” (Kunz et al. 2011). This report presents the first worldwide documentation of bat occupancy in manmade transportation infrastructures. In recent decades, environmental responsibility and stewardship practices have become intrinsic to the project development process of state DOTs. Consideration to wildlife movements (mammal collision mitigation measures), ecosystem impacts and timetables (stream crossings and fish spawning, bird incubation periods) are now at the forefront of transportation planning. Numerous states have become environmental champions, actively engineering and retrofitting bridges to accommodate bat colonies. Sixty percent of North American bat species exploit bridges and occupancy rates can surpass 90%. Yet, definitive guidelines to address bat occupation (including identification and mitigation procedures) are nonexistent. Both personally and professionally, we have become incredibly invested and interested in this issue. Within the last three years, we have realized not only the impacts of operational activities, but the opportunity for conservation triumphs. While this request was put forth by the NMDOT Environmental Bureau, we nonetheless sought to provide holistic standards because every phase of the project development process potentially impacts bats. Operational activities that adversely affect bats primarily include roost destruction, modification of habitats and direct disturbance during critical life phases (maternity and weaning periods,


hibernation). Even those projects with uncomplicated scopes (e.g., pavement rehabilitation and reconstruction, bridge deck replacement, guardrail and fencing) and minimal environmental impacts may be distressful to resident colonies. We, therefore, include recommendations for new construction, demolition (reconstruction/rehabilitation) and maintenance activities. We strove to consolidate the extensive amount of information we’ve amassed into complete, but practicable guidelines for NMDOT personnel, officials and subcontractors. These procedures are recommended to ensure bat welfare, consistency of approach and provide a sound, scientific foundation for effective planning and implementation of policies. The procedural advice we provide illustrates exemplar measures; thus, they may necessitate modification for feasibility, site specific differences, and project constraints (e.g., phase of development, available budget, conservation status of respective bat species). The first three sections elaborate on roost ecology, ethology (behavior) and biology; which provide an overall framework for understanding the interplay between bat physiology and roost selection and appreciating the influence of location and environment. These considerations facilitate the identification of current and potential roosts, their relative significance, potential interactions and appropriate measures by which to minimize or eliminate impacts. The succeeding sections address the ‘who, what, when, where and why’ of bat/bridge occupancy and tenable complications relative to swallow nest occupation and winter activity. We also present concerns and implications with respect to the deficiencies of current survey techniques and propose specific directives and requirements. While we incorporate recommendations as they pertain to individual sections within the report, Appendix A details the measures by which to abate negative impacts on resident bat communities or prevent avertible outcomes resulting from significant alterations to the immediate landscape. Three of four threatened and endangered species endemic to New Mexico have been either documented in bridges or is likely to do so. However, comprehensive information on the populations and distributions of the state’s 29 species is immensely deficient. The Western Bat Working Group, who assigns conservation priority statuses to western bat species, recognizes the “need to gain a regional perspective and more complete distributional information” and that “population status and trend data are lacking and seriously needed for most species.” This deficiency is one reason we include substantial references to international species, particularly European bats; which parallel those endemic to New Mexico - in appearance, life history requirements and roost selection. The other reason is the continent’s national implementation of the Habitats Directive (92/43/EEC); which protects all bat species, their habitats and roosts. The indiscriminate decline of Europe’s bat populations and the respective legal responsibility of its constituent countries elicit widespread scientific interest and conservation initiatives. As a result, these countries provide the most progressive and comprehensive volume of research and mitigation measures relative to the planning, design and implementation of national road schemes. Clearly, a close association exists between bridges and roads and thus, their standards provide an incredible foundation for this report.


Two of the three publications that reference New Mexican bridge roosts are limited in scope; Chung-MacCoubrey (1999) identifies only 1 colony/1 bridge and Geluso et al. (2005) note 1 individual/1 bridge. These studies provide the rationale for an NMDOT statewide bridge evaluation - Chung-MacCoubrey and Geluso et al. demonstrate the paucity of available information for New Mexico, and Geluso and Mink (2009) illustrate the potential rate of occupancy. Because references to bridge habitation in New Mexico are limited, and to help apply statements/thoughts to New Mexican bats, we have included the following superscript designations throughout the main text (1 indicates New Mexico-specific species, 2 indicates North American species). We hope not only to communicate the importance of establishing standards for NMDOT activities, but to illuminate the value and awesomeness of bats and impart the desire to conserve and protect these mammals. With the tragedy of white-nose syndrome (ca. 7 million deaths) and the unpredictability of its westward expansion, any measures in consideration of bats is of enormous conservation value. Needless to say, it is at NMDOT’s discretion whether to elaborate on, develop protocols from, or provide basic principles due to these recommendations. However; an instituted framework, with this report as its foundation, can establish the standard by which all transportation authorities could orchestrate bat mitigation, management and conservation initiatives.



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ighway infrastructures are omnipresent features of human activity and provide productivity benefits (e.g., transport, market expansion, production and distribution, accessibility and tourism) and connect regional development. However, linear infrastructures (e.g., roads, highways) represent one of the most consequential anthropogenic impacts on natural ecosystems. Although adverse effects from highway infrastructure exist (mortalities; barrier effects; isolation, restriction of animal movement, habitat patchiness, population fragmentation), these networks can establish otherwise esoteric microhabitats and desirable resources, (e.g., remnant vegetation, foodstuff for scavengers, sodium pools)(Laurian et al. 2008, Leblond et al. 2007), and road verges may constitute “long, ribbon-like habitats,” that facilitate movement and dispersal (Coffin 2007). North American chiropteran1 species readily exploit these anthropogenic structures, which function as alternative roosts (i.e., diurnal roosts, nocturnal roosts, maternity roosts) and stepping-stone refugia (i.e., transitory roosts) for migratory populations. Forty five microchiropteran species of 19 genera and four families (Mormoopidae, Phyllostomidae, Vespertilionidae, Molossidae) populate the United States (Adams 2003). Of those species, 64% inhabit New Mexico. In North America, many species are categorized as endangered or threatened, have become candidates for these categories, or are considered species of concern (O’Shea and Bogan 2003). In the southeast, 87% of bat species carry special conservation designations (e.g., rare, sensitive, species of concern) somewhere within their range (Laerm et al. 2000 as cited in Menzel et al. 2003). In the western US, more than 45% of chiropterans receive the highest priority status for funding, planning, and conservation actions (Western Bat Working Group 2007, Table 3). Although New Mexico lists only six sensitive species (4 ‘species of concern’ status, 2 ‘endangered’ status; USFWS 2009), the population estimates and distributions of the state’s 29 microchiropteran species are largely unknown. A former Department of the Interior ‘Species of

“Today, the significance of roads has expanded beyond pavement, and their role as links between wildlife corridors is now at the forefront of transportation planning.” Concern’ list identifies Myotis ciliolabrum1, M. evotis1, M. thysanodes1, M. velifer1, M. volans1, M. occultus1, M. yumanensis1, Corynorhinus townsendii1 Lasiurus cinereus1, Euderma maculatum1, Tadarida brasiliensis1, Nyctinomops macrotis1, Eumops perotis1, E. underwoodii2, Leptonycteris curasoae1, L. nivalis1, Choeronycteris mexicana1, and Macrotus californicus2 (Adams 2003). Of these 18 species, 77.7% (14) exploit bridges as either nocturnal or diurnal roosts. Of those 14 species, 13 inhabit New Mexico. As the quantity and quality of natural roost sites dwindle, manmade infrastructures (e.g., mines, buildings, culverts, bridges) function as vital habitat for bat communities worldwide. Interior spaces of these anthropogenic structures proffer physical and thermal characteristics reminiscent of natural roosts and therefore, have become amenable substitutes. The successfulness of the Congress Avenue Bridge (Austin, Texas) has prompted surveys and studies of bridge thermodynamics, microclimates, roosting ecology, species 1 Chiroptera refers to the taxonomic order of mammals consisting of megabats (Old World fruit bats), which constitute the suborder Megachiroptera; and microbats (echolocating “true bats”), which compose the suborder Microchiroptera.

Opposite page: Colony of Mexican free-tailed bats, Tadarida brasiliensis 1


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Opposite page: Female Arizona myotis, Myotis lucifugus occultus, with newborn tucked under wing

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occurrence and activity patterns. A comprehensive, exhaustive literature search generated numerous articles and publications documenting bridge use by bats worldwide. Keeley and Tuttle (1999) list 24 North American species that exploit bridges and “thirteen others likely do” (Tables 1-2) Adams surmises that 93 percent of California’s rare bat species benefit from these structures. Keeley and Tuttle (1999) estimate 4,250,000 bats currently inhabit 211 highway structures. Within the southern United States, estimates approach 33 million bats per 3,600 highway structures (American Association of State Highway and Transportation Officials [AASHTO] 2008).

roost site selection Roost site selection has considerable consequences for the survival and reproduction of bats. A multitude of factors influence the roosting ecology of these mammals; including insect availability, predator avoidance, sociality, thermoregulation and roost structure, location, availability and abundance. The abundance and diversity of available roost structures influence the distributions of most temperate zone species. Menzel et al. (2003) and Bogan et al. (2003) affirm that bat community richness closely parallels the diversity of available roost structures. Cavity quality directly influences reproductive success; either directly via juvenile survival, or indirectly via their subsequent growth and development. Various factors influence postnatal growth rates and survivorship of young; including climate, availability and abundance of foodstuff, roost temperature, age, mother’s nutritional and hormonal condition, parasite loads, colony size, and presumably anthropogenic factors (Allen et al. 2010). Low temperatures may delay gestation, reduce birth size, or slow postnatal growth (Tuttle 1976). Chalinolobus tuberculatus selects for large roost-tree diameters and thicker cavity walls that

precipitate warmer, and more stable microclimatic conditions (Sedgeley 2001). Similarly, Myotis sodalis2 may choose roosts with more solar exposure because elevated temperatures accelerate embryonic and juvenile development (Carter and Feldhamer 2005). The energy requirements of breeding females are extremely high during pregnancy and lactation; approximately 2.5-5 times the level of nonreproductive energy expenditure (McLean and Speakman 1999). Additionally, reproductive females minimize daily torpor (i.e., mechanism to conserve energy) because it can negatively impact reproductive success and mother/offspring fitness (Sedgeley 2001, Kerth et al. 2001, Willis and Brigham 2007). Kerth et al. (2001) conclude that no single optimal roost serves the energetic needs of variable weather conditions or reproductive phases. Kurta et al. (1996) and Callahan et al. (1997) convey that individuals that comprise maternity colonies may exploit several trees to provide the total resources (i.e., cover, correct temperature) necessary (as cited in Carter and Feldhamer 2005). These roosts may be chosen during adverse environmental conditions (e.g., rain, wind, temperature extremes) or loss of primary roost, to reduce parasite loads, access foraging grounds, or to minimize predation. Numerous publications document the presence of maternity colonies and creches (nurseries) within bridges (Davis 1966, Swift 1997, Zahn 1999, Erickson et al. 2002, Wolf and Shaw 2002, Hendricks et al. 2004, Geluso and Mink 2009, Papadatou et al. 2011). Of 27 New Mexico bridges known to provide roosting habitat, more than 41% have documented maternity colonies (Table 4, Map 1). Allen et al. (2010) found that pups born at bridge sites were larger (i.e., body mass, forearm length) and grew faster than those born at cave sites. Therefore, bridge born pups achieved adult size more rapidly, became volant earlier, and thus, acquired a selective advantage compared to cave born pups. Similarly, Geluso et al. (1981) describe bridge inhabiting bats

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exhibiting heavier weights than individuals from caves, including Carlsbad Caverns. Additionally, body condition (i.e., ratio of body mass to forearm length) was greater for females inhabiting bridges (Allen et al. 2010). Bennett et al. (2008) describe the affinity for anthropogenic structures increasing May-August, when maternity colonies appear. These structures are extraordinarily valuable because they provide females with warmer and more stable thermal environments that impart substantial energetic benefits. Maternity roosts are insulated against temperature extremes and have significantly smaller temperature and humidity ranges relative to external, ambient conditions (Sedgeley 2001; Smith and Stevenson, unpubl. data, Figure 1). Females communally roost in maternity colonies, and therefore, social thermoregulation may also play an important role. Although most scientists concur that microclimate strongly influences roost site selection, Willis and Brigham (2007) propose that sociality may preeminently influence roost preferences. Willis et al. (2006) demonstrate preferences to tree cavities with relatively large volumes, which permit individuals to roost collectively. Additionally, colony (group) size was positively correlated with cavity volume. The microclimate within maternity roosts can transform substantially from the metabolic heat of its occupants, increasing internal temperature 5-10 °C above that of unoccupied roosts (Sedgeley 2001). Willis and Brigham (2007) illustrate a significant positive correlation between quantity of bats, maximum daily roost temperature, and energy savings. On average, one individual (normal body temperature) can conserve ca. 9% of its daily energy budget by roosting with conspecifics, which can increase to 53% in a colony of 45. Smith and Stevenson (unpubl. data) document bat occupancy of ten distinctive structures as roosts; including concrete spalls, ¼” crevices in the 2 terminal spans (transverse joint between eastbound and westbound decks), steel drainage pipes, external expansion joints between deck slabs, internal expansion joints above column caps, “open” beams between centermost girders (.639 m), typical “open” beams (1.459 m), bolt cavities within pre-insulated pipes, space between piers and pedestals, and cliff swallow (Hirundo pyrrhonota) nests (Gallery 1). Structures with the greatest volume (i.e., internal and external expansion joints, and open beams) receive the most use (Smith and Stevenson, unpubl. data). Preferred old-growth redwood hollows had greater hollow volumes, greater diameters, and were closer to water than less frequently used trees (Gellman and Zielinski 1996). Similar to concrete bridges, these trees exhibit relatively stable temperatures and humidity, comparative permanence, protection from inclement weather, and spacious internal flight areas.

distance to resources Roost sites adjacent to optimal foraging areas (e.g., lakes, rivers) are energetically ideal, because the foraging of insectivorous bats requires a high rate of energy expenditure. Additionally, females increase food consumption to accommodate the energy demands of reproduction. For Myotis lucifugus1 and M. velifer1, food

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Figure 1. Graph illustrating the stability of internal roost (expansion joint) temperatures compared to ambient. Includes data from 1 May - 2 June 2012, gestation period (top); and from 14 July - 23 August, lactation period (bottom). For both graphs; diamond represents ambient temperature and circle reflects internal roost temperatures.

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Sixty percent, and potentially 89% of North American bat species exploit manmade bridges.

consumption increases approximately 45% from pregnancy to lactation, and between early and mid-lactation, T. brasiliensis1 increases energy intake by 82% (McLean and Speakman 1999). Flight durations (i.e., foraging times) may increase and presumably, influence site selection. Therefore, roost locales that minimize travel distances are preferable for both sexes. Evelyn et al. (2004) document that M. yumanensis1 select roosts close to water and ‘distance to water’ was one characteristic that differentiated roosts from random comparisons. Similarly, a meta-analyses of 56 studies by KalcounisRuppell et al. (2005) affirms that roosts of cavity inhabiting species were closer to water than random parities. M. grisescens2 demonstrates a dramatic correlation between colony location and major bodies of water (Tuttle 1976). Similarly, the activity centers of female M. evotis1 were significantly closer to water than random points, with the probability of use decreasing with distance to available water (Waldien and Hayes 2001). Growth success, percent of low weight pups, and mortality closely correlate with travel distances to foraging areas (Tuttle 1976). Differential energy expenditure relative to travel may contribute to the disparate birth sizes between cave- and bridge- born pups found by Allen et al. (2010). Because water and bridge localities often converge, bats inhabiting bridges have shorter foraging commutes than those inhabiting caves. Shortened commutes and foraging times allow pregnant females to allocate more resources and energy and nutrients to reproductive growth and milk production, respectively.

permanence and roost switching Frequent movements between diurnal shelters occur for various mammalian species (e.g., fox, Vulpes vulpes; skunk, Mephitis mephitis; badger, Meles meles; raccoon, Procyon lotor) (Lewis 1995, Trousdale et al. 2008). In chiropteran species, where interand intraspecific variability exists, the phenomenon of roost switching is not well understood. The relative availability and permanency of roosts may affect roost selectivity and subsequent fidelity. Lewis, and Kunz and Lumsden (2003) illustrate that fidelity corresponds to roost permanency and inversely relates to roost availability. Bats, therefore, exhibit low fidelity for ephemeral, abundant sites (e.g., dead or senescent trees, exfoliating bark) and strong fidelity to relatively rare, permanent structures (e.g., manmade bridges, buildings, caves). Utilization of impermanent roosts may be offset by location, and therefore, lower commuting costs; lower ectoparasite levels; and a relatively high abundance and availability of sites. Thus, bridges may precipitate higher roost fidelity; the benefits of which include lower costs relative to roost searching, site familiarity, and the ability to maintain social relationships (Lewis 1995, ChungMacCoubrey 1999). Trousdale et al. (2008) and Bennett et al. (2008) report high lability of C. rafinesquii2 to abundant, “not exceptionally stable” hollow trees, and site-faithfulness to comparatively permanent caves and manmade structures. Similarly, Brigham (1991) Opposite page: Myotis spp. roosting on open beam 7


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Below: Yuma myotis, Myotis yumanensis; Opposite page: Myotis spp.

affirms that E. fuscus1 populations were “tenaciously loyal” to anthropogenic structures and rock crevices, but noncommittal to tree cavities. Goldingay and Stevens (2009) propose that the rate of collapse of hollow-bearing trees may exceed replacement. Consequently, many native Australian, cavitydependent species are presently listed as threatened. A study by Carter and Feldhamer (2005) indicates that 25-30% of typical ephemeral roosts (i.e., tree snags with exfoliating bark) fell within one year, many within weeks; and most of the exfoliating bark pieces fell within months. Contrastingly, manmade bridges have service lives of approximately 75-100 years. Within and between years, bats show fidelity (Willis and Brigham 2007) to roost sites, patterns consistent with long-term colony stability. Bats demonstrate multiyear fidelity to night roosts, particularly bridges. Lewis (1994) documents Antrozous pallidus1

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returning to night roosts within and between years. Similarly, individual Myotis volans1, E. fuscus1 and M. yumanensis1 have been recaptured within the same bridge 2, 3, and 4 years, respectively, after banding (Pierson et al. 1996). Myotis volans1 and M. lucifugus1 were recaptured 6-10 years later at the same bridges (Ormsbee et al. 2007). Worldwide, numerous studies document the propensity of bats to exploit manmade concrete and timber bridges as roost sites (Table 5-6). Certain concrete properties (e.g., thermal stability, high mean relative humidity) may confer thermoregulatory benefits similar to natural roosts. Structural precast concrete, although not an especially excellent heat conductor or insulator, exhibits thermal stability; the ability to maintain internal temperature within a certain interval, given normal external temperature oscillations. A large thermal mass provides “inertia” against temperature fluctuations; thereby, producing


“chamber” temperatures beneath the bridge that are higher (often by 10 °C or more) and more stable than external ambient conditions (Lacki et al. 2007, Erickson et al. 2002). Timber and concrete demonstrate different behaviors, especially when subject to quick thermal variations. The distribution of temperature remains almost constant within concrete; whereas timber reacts sensitively to thermal variations (Fragiacomo and Ceccotti 2006). Similarly, the relative humidity variations of the environment affect timber and concrete differently. Several properties of timber (elasticity, shrinkage/swelling, mechano-sorption) are dependent on moisture content. Conversely, even for concrete subject to extreme conditions (e.g., long cycles of absorption within water and air drying), the relative inelastic strains are small and occur more slowly than for timber. Therefore, the influence of relative humidity on concrete is

negligible (Fragiacomo and Ceccotti 2006). Thus, timber bridges may proffer numerous microclimates, with varying temperature and moisture parameters. Similarly, the Los Lunas bridge (concrete), which is unique in its availability, abundance and type of roost structures, creates the potential for gradients relative to roost microclimates.

bridge preferences Sixty nine percent, and potentially 91%, of North American bat species exploit manmade bridges. In order of preference; microchiropterans exploit parallel box beam bridges, cast-in-place or pre-stressed concrete girder spans (Keeley and Tuttle 1999, Ferguson and Azerrad 2004). Of concrete cast-inplace with “chambers” on the underside of the bridge; concrete flat bottom; I-beam, parallel concrete or steel girders; and wooden; Adam and Hayes (2000)

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Above: Several Myotis spp. roosting inside metal drainage pipe 11


affirm that concrete cast-in-place bridges support the highest bat densities. A publication by Davis and Cockrum (1963) reports of several extensive studies of bridge colonies by members of the Department of Zoology of the University of Arizona. These authors specify that, “while most bridges apparently furnish sufficient shelter at night, only those of very specific construction provide the necessary conditions to allow them to serve as day-roosts.” Although numerous studies identify affinities to concrete structures, Smith and Stevenson (unpubl. data) and Geluso and Mink (2009) document considerable occupancy of timber bridges. However, bats will not roost within timber structures that have newly applied creosote, an oily wood preservative with a pungent odor (Smith and Stevenson, pers. comm., Lance et al. 2001). Of 36,629 day roosting bats, 99.8% inhabited timber bridges (Geluso and Mink 2009). In these bridges; bats occupy narrow crevices (formed when two adjacent parallel beams separate), spaces above vertical supports at the culmination of spans, narrow space between beams and deck, vertical face of beams, and between guardrail post and outermost beams (Gallery 2). In Montana, a postpartum female Lasiurus cinereus1 with two young roosted in a narrow crevice (4 cm width, 20 cm depth) between two wooden girders 5.3 m above bare ground (Hendricks et al. 2004). Concurrently, a maternity colony of approximately 15 E. fuscus1 occupied the same crevice where the slot was narrower, ca. 1.5 m from the female L. cinereus1. Ferrara and Leberg (2005) report 90.6% of Louisiana’s double-T concrete bridges had bat occupancy. Wolf and Shaw (2002) found 92% occupancy by eight species in the Tucson metropolis of Arizona, including 11,400 T. brasiliensis1 “in 1 bridge at 1 time.” In Montana, 60% (78) of structures (125 bridges, five culverts) surveyed had evidence of bat use; 66 and 12 were designated nocturnal roosts and diurnal roosts, respectively (Hendricks et al. 2004). Percent usage was 75.9% of concrete structures, 37.5% of steel

structures, and 31.6% of wooden structures. Night roost locations were relatively exposed; “typically the vertical face of a girder (concrete or steel) near the abutment with the underside of the deck and in darker areas between girders and close to the intersection with the ground or embankment.” Day roosts were within narrow vertical spaces 3-5 cm (1.25-2”) wide and ≥ 11-20 cm (4.5-8”) in wood or concrete bridges; three maternity colonies (E. fuscus1, Myotis lucifugus1) roosted in wood bridges, one in a concrete bridge. All day roosts in wood bridges were underneath the deck, but five of seven in concrete bridges were in expansion joints between deck sections and near the deck edge. Interviews with researchers from Alabama, Pennsylvania, Massachusetts, New York, Kentucky, North Carolina, Tennessee, West Virginia, Georgia, and Indiana document Myotis leibii2 inhabiting bridge structures of Kentucky, North Carolina, Tennessee, and West Virginia (Erdle and Hobson 2001). One interviewee from North Carolina documents “the largest maternity colony in the southeast this past summer roosting in the expansion joints of a concrete bridge” (Erdle and Hobson 2001). Bennett et al. (2008) and Lance et al. (2001) demonstrate a strong relationship between presence of Corynorhinus rafinesquii2 and construction type. C. rafinesquii2 demonstrates an affinity for large, concrete girder bridges and avoids flat-bottom slab bridges (Bennett et al. 2008). A comprehensive survey of 1,129 bridges throughout South Carolina established new records of occurrence for 10 counties, thereby determining more accurate population distributions. Mean frequency of use was 65.9% and the probability of finding bats beneath an individual bridge was 46-73%, depending on previous year occupancy. These authors, therefore, recommend an inspection interval of 3-5 times annually to determine use. Similarly, Wolf and Shaw (2002) “found some Tucson bridges occupied only in April, and would have thought they were never

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Above: Myotis spp. captured from the Los Lunas bridge 13


occupied had I surveyed only once in March or May.” Ninety percent of the bat species inhabiting the Oregon Coast Range exploit bridges for roosting (Adam and Hayes 2000). Of 17 bridges surveyed in southern New Mexico, 15 (88%) contained day roosting bats and > 47% had maternity colonies with one or more species, including M. lucifugus occultus1, M. yumanensis1, and T. brasiliensis1 (Geluso and Mink 2009). Similarly, bridges are valuable structures for maternity and nursery colonies of Myotis nattereri, a species considered vulnerable throughout Europe and the United Kingdom (Swift 1997). Numerous publications document the presence of C. rafinesquii2 and M. austroriparius2 within bridges (Rice 1957, Lance et al. 2001, Bennett et al. 2008). C. rafinesquii are considered ‘Species of Concern’ throughout their range. In Mississippi, C. rafinesquii2 and M. austroriparius2 are endangered and threatened, respectively. Leptonycteris curasoae yerbabuenae1, listed as endangered by USFWS, occupies bridges of the Huachuca Mountains, Arizona (Hollis 1995). A cursory survey of several structures throughout Los Lunas, Belen, and Albuquerque; information from Geluso and Mink’s (2009) study, and a 20102011 re-examination of those bridges, indicate that bat occupancy rates (8896%) are substantial in New Mexico. This estimate mirrors those of Louisiana and Arizona; southern states which have year-round bat activity, and relatively high species richness and diversity. According to Kunz and Lumsden (2003), deforestation and conversion of native habitats to intensive agriculture and human development constitute the most significant threats to the density and distribution of local bat faunas. Several authors document facultative foliage roosting bats opportunistically exploiting bridge structures, including Lasiurus cinereus1 (Hendricks et al. 2004), L. blossevillii1 (Kunz and Reynolds 2003) and Lasionycteris noctivagans1 (Geluso and Mink 2009, Hendricks et al. 1994). As the availability and abundance of valuable roosts dwindle, occurrences of foliage roosting species exploiting anthropogenic structures may concomitantly intensify. Sgro and Wilkins (2003) demonstrate that bats prefer roosting over roadway segments to roosting over embankment sites (similar to Bernardo, Photo A). At the highest population densities (19 July), bats hung, oftentimes completely exposed, from protruding bolts and along the exterior bevel of the expansion joint. Of four colonies discovered by Chung-MacCoubrey (1999); the third colony, which consisted of ≥ 250 Myotis yumanensis1, roosted within deep, vertical crevices (1.5 cm wide) extending the length of the underside of a small concrete bridge adjacent to the river. Erdle and Hobson (2001) document three primary habitat types of Myotis leibii2; abandoned mine portals and caves, high elevation rock outcroppings, and expansion joints of concrete bridges. Rice (1957) professes that during the winter, Myotis austroriparius2 typically roost over water within the crevices between wooden bridge timbers, storm sewers, road culverts, boat

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Above: Myotis yumanensis roosting in 1/4� crevice in the Los Lunas bridge.

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houses, and the vertical drain pipes of concrete railroad bridges. Bats typically occupy the warmest “chambers,â€? the terminal spans that usually occur over land, proximal to abutments, and often within recesses that are centrally located relative to bridge width (Lacki et al. 2007, Lance et al. 2001, Bennett et al. 2003, Feldhamer et al. 2003, Ferrara and Leberg 2005). Sloping riverbanks or use of fill to stabilize bridge supports often result in end chambers being closer to the ground than center chambers, and occupied chambers occasionally are < 2 m above ground (Lacki et al. 2007). Ferrara and Leberg (2005) describe roosting heights of approximately 2 m, with a tendency to roost closer to ground level than the mean height of the bridge. At the Los Lunas bridge, suitable roosts within crevices and pipes may be 2.28 m, with pipe entrances 1.20 - 2.59 m above ground (Smith and Stevenson, unpubl. data). Dickerman et al. (1981) indicate that Saccopteryx bilineata, a species endemic to Central and South America that occasionally occupies bridges, roosts at heights of 1-10 m. Perlmeter (1996, as cited in Lacki et al. 2007) reports that concrete bridges maintain mean temperatures of 9.3-15.2 °C higher than ambient temperatures. Chambers presumably retain daytime heat, provide shelter, reduce adjacent air flow, and minimize convective heat loss. Central chambers (i.e., those over water) cool more rapidly than end chambers, which are comparatively insulated by proximity to ground and protection from air currents (Perlmeter 1996, Adam and Hayes 2000, Renison 2003 as cited in Lacki et al. 2007). Similarly, Adam and Hayes (2000) conclude that greater use of end chambers correlates to the presence of microsites with favorable thermal characteristics. Erickson et al. (2002) propose

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that bridges attenuate wind and reduce wind chill and dehydration potential.

culvert use Worldwide, culverts provide additional habitat for diurnal roosting species (Table 7). Corynorhinus rafinesquii2 routinely exploits both bridges and culverts (Lance et al. 2001, Bennett et al. 2008). The African bat, Nycteris thebaica, frequently roosts within road culverts and females exhibit considerable roost fidelity, returning to the same culvert or neighboring set of culverts over several years (Monadjem 2005). A new subspecies of the previously monotypic Hylonycteris underwoodi was collected from culverts in Jalisco, Mexico. Additionally, specimens of Micronycteris megalotis, Sturnira lilium, Artibeus jamaicensis, and A. lituratus were obtained from the same culvert (Phillips and Jones 1971). Two common neotropical species; the nectarivorous Glossophaga soricina and the frugivorous Corollia perspicillata were captured in a culvert near Albrook Air Force Base, Canal Zone, Panama (Klite and Kourany 1965). Ellison et al. (2003) document the utilization of culverts by Myotis austroriparius2, M. grisescens2, M. leibii2, M. septentrionalis2, and M. sodalis2. In Texas, Perimyotis subflavus1 (formerly Pipistrellus subflavus) were found overwintering in concrete box culverts along Interstate Highway 35 (Sandel et al. 2001). The Texas Rare Bat Survey by the Texas Parks and Wildlife Department reports the discovery of M. austroriparius2 wintering in culverts (Clark 2003). Fraze and Wilkins (1990) documented approximately 1,000 Tadarida brasiliensis1 on culvert walls, and evidence indicates that parturition may have occurred. These authors determine that T. brasiliensis1 utilizes culverts primarily in spring and summer; however, population increases in March intimate that culverts may additionally function as transitory “stop-over� roosts for migratory colonies. Culverts proffer valuable connectivity of habitats adjacent to highways. They are readily accepted by and frequently employed by slow / low flying bat species. Bats presumably conform to the watercourse or arroyos that travel through culverts. The predominant species exploiting culverts, for both roosts and commuter routes, are those adapted to dense environments. Their wing morphology (i.e., maneuverable flight) and echolocation pulses enable these species to fly and detect small obstacles within small spaces, such as culverts. Boonman (2011) documents cross sectional area as the most important determinant of culvert use, which was positively correlated with activity. A study by Boonman (2011) reports that 85% of 54 culverts were used by bats.

nest use The microclimatic properties critical to avian species parallel those of microchiropterans; and include wind, radiation, air temperature and humidity. These parameters directly influence thermoregulatory demands; therefore, nest structure and placement (e.g., orientation, shelter from inclement weather and wind, elevation, materials) become vitally important. These structures, when abandoned or unoccupied, provide ancillary roost habitat for chiropterans worldwide; including several North American species (Eptesicus fuscus1, Tadarida brasiliensis1, Myotis velifer1, M. yumanensis1) (Table 8). Shulz (in press) calculates an incidence rate of 3.9 bats per 100 Hirundo ariel nests. This author

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mentions the same location of the same species over two consecutive years may indicate permanent residency. A study by Schulz and Hannah (in press) documents Murina florium, listed as vulnerable (i.e., fewer than 50 specimens within Australia), predominately exploiting Sericornis citreogularis nests. In fact, these nest roosts accounted for 64% of total roosts found. The only documented roosts of Kerivoula papuensis, listed as rare, are those within the abandoned nests of Sericornis spp. and Gerygone mouki. The threatened species, M. florium, may inhabit the nests of Oreoscopus gutturalis at a frequency rate of 3.33% (1 in 30 nests) (Schulz 1997). The enclosed mud nests of Hirundo ariel provide roosting habitat for two rare Australian bat species, Chalinolobus dwyeri and Nyctinophilus timoriensis (Hyett 1980, Hyett and Shaw 1980, Schulz 1997). Sharma (1986) concludes that Kerivoula picta utilizes the nests of Ploceus philippinus due to the scarcity of other amenable roost sites. Buchanan (1958) documents Tadarida brasiliensis1 and Myotis velifer1 occupying Petrochelidon pyrrhonota nests. All nests from which specimens were collected contained both Tadarida1 and Myotis1 communally, with the exception of one nest which contained only two Myotis1. Buchanan reports 20 Tadarida1 and five Myotis1 within one nest, and 17 individuals from four additional nests. A note by Jackson et al. (1982) describes M. velifer1 occupancy ratios (no. individuals to no. nests surveyed) of 11/28 (39%) and 7/29 (24%) at two concrete box culverts, respectively. These authors surmise the abundance of M. velifer1 in the arid southwest “may well be due to their preference for ‘crevice’ roosts and their opportunism shown here by their use of birdbuilt ‘crevices’ on the walls of man-made ‘caves’.” A cursory survey of one NMDOT bridge (No. 7339, Route NM 346) suggests that incidence rates are comparable to those of Jackson et al. (1982). Only Chalinolobus morio, a species endemic to Australia, has been recorded hibernating in birds’

nests (Richards 1983); specifically, those of H. ariel. Interestingly, H. ariel is the only species that constructs an enclosed bottle-shaped mud nest, a structure indistinguishable from those built by Petrochelidon pyrrhonota. This convergent similarity suggests that P. pyrrhonota nests possess comparable thermal qualities and may, likewise, function as suitable winter roosts for several southwestern bat species. Although relative prevalence of this occurrence may be low, we have documented nest occupancy into December (mean temperature, 5 °C; wunderground.com). The U.S. Migratory Bird Treaty Act (MBTA; 16 U.S.C. 703-712; Ch. 128; July 13, 1918; 40 Stat. 755) protects North American Hirundo and Petrochelidon species. NMDOT’s environmental commitments address potential impacts to these species; however, there are no commensurate regulatory policies to safeguard bats. If construction occurs during the nesting season (March-July), NMDOT will ensure that a migratory bird nest survey is conducted and unoccupied bird nests are removed. Bats that occupy H. rustica nests lay nearly prostrate within the nest cup and those within P. pyrrhonota nests (gourdshaped enclosed structures) are typically concealed and undetectable without a borescope or fiberscope. Therefore, a survey to identify swallow activity will not discern bat occupancy. Jackson et al.’s 1982 ‘note’ (The Southwestern Naturalist Notes Section; documents brief, notable field observations) provides not only the most contemporary, but the only estimates of occupancy rates within the United States. The distribution of Hirundo and Petrochelidon species encompasses most of North America; therefore, this occurrence may have widespread implications. We recommend either 1. an evaluation of swallow nest occupancy be a component of the statewide bridge survey, or 2. an adequate proportion of nests be inspected during the applicable phase of development (i.e., Environmental Investigations, Analysis of Alternatives; Federal

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Highway Administration 2009).

bridges as hibernacula In temperate North America, colder months signify lower ambient temperatures and the concomitant reduction of insect prey. To circumvent this ‘energetic bottleneck,’ migratory species relocate to warmer environments, whereas other species hibernate and remain relatively inactive. Hibernating bats select ambient temperatures which allow them to drop their rate of metabolism to very low values, and maintain a temperature differential with the environment of only a few tenths of a degree Celsius (1- 2 °C above ambient), a state that has been referred to as ‘‘thermo-conformity’’ (Arlettaz et al. 2000). In the southwestern United States, certain species have been found active throughout winter. Captures by O’Farrell and Bradley (1970) indicate that Pipistrellus hesperus1, Myotis californicus1, and Antrozous pallidus1 are active year-round in Nevada. P. hesperus1 and M. californicus1 were netted at temperatures from -8 to 33 °C. “There appears to be an alternative to hibernation or migration, at least in the warmer areas of the southwest” (O’Farrell and Bradley 1970). M. californicus1 exhibits intraspecific plasticity relative to winter activity patterns, which vary from sustained hibernation to intermittent dormancy. O’Farrell et al. (1967) report approximately 11% of P. hesperus1 were netted at temperatures of 5 °C or below, indicating considerable activity at low ambient temperatures. Similarly, M. californicus1 were active at ambient temperatures between 2 and 6 °C. In southern and central New Mexico (Bernalillo, Eddy, Grant, Hidalgo and Socorro counties), Geluso (2007) captured 12 species November March (hibernation period); with P. hesperus1, M. californicus1, and A. pallidus1 captured every month; and L. noctivagans1 and T. brasiliensis1 netted every month except January. Ambient temperatures at time of capture vary from 1- 22 °C. Also within

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New Mexico (Catron county), L. noctivagans1 and P. hesperus1 were captured at temperatures of -2 °C and 1 °C, respectively (Barbour and Davis 1969). Geluso (2008) documents the mass exodus (tens of thousands) and return of T. brasiliensis1 from Carlsbad Caverns in November, February, and March, and ≥ 100 individuals in December and January. Nonmigratory populations of T. brasiliensis1 from the southeastern United States, California, and southern Oregon are year-round residents. In these locales, this species typically roosts within buildings, forming maternity and winter colonies in warm and cooler months, respectively (Kunz and Reynolds 2003). Scales and Wilkins (2007) report that, in downtown Waco, urban roosts were in use 21.4% of monitored winter days, demonstrating continued occupancy of these roosts through winter. These roosts may satisfy physiological requirements that caves do not; a tolerable thermal environment and proximity to foodstuff that remains available year-round. In California, overwintering or hibernation roosts typically occur from late fall-early spring. In many instances, these sites are also used as diurnal roosts the remainder of the year (Erickson et al. 2002). Hoffmeister (1970) describes winter residence of M. velifer1, M. thysanodes1, M. californicus1, M. subulatus (Myotis leibii)2, L. noctivagans1, P. hesperus1, L. cinereus1, L. ega2, A. pallidus1, T. brasiliensis1, T. molossa (Nyctinomops macrotis)1, M. waterhousii (Macrotus californicus)2, E. fuscus1, P. townsendii (Corynorhinus townsendii)1, and E. perotis1 in Arizona. Menzel et al. (2003) document a citation of P. subflavus1 hibernacula in culverts, and houses. Benson (1947, as cited in Bernardo and Cockrum 1962) reports eight hibernating Tadarida brasiliensis1 inside the crevice of a bridge, 6.5 miles southeast of Wilbur Springs, California on 1 December. Wolf and Shaw (2002) denote that T. brasiliensis1 and P. hesperus1 were present year-round, and document winter colonies of T. brasiliensis1 in Tucson bridges. Corynorhinus rafinesquii2 have been observed


hibernating in an abandoned concrete structure in northcentral Mississippi (Martin et al. 2011). Several bridges within DeSoto National Forest were used as roosts during winter (2008), and C. rafinesquii2 hibernates in cisterns, wells, and culverts in the northern part of their range (Martin et al. 2011). Cel’uch and Ševčík (2008) identify a hibernation colony of 10,000 Nyctalus noctula that occupy a road bridge in Germany. Weaver (2012) documents T. brasiliensis1 occupying D’Hanis Bridge (Texas) in November, December, January and February of 2010-2011 and 2011-2012, with an estimated population of 226,350 bats. D’Hanis Bridge exhibited temperature and humidity values similar to ambient; minimum and maximum temperatures were 7.45 and 19.31 °C from December January. This author notes 3 additional bridges in Hays County (2) and Travis County (1), Texas with overwintering populations of T. brasiliensis1, indicating the northward expansion of this species’ winter range. Similarly, we documented T. brasiliensis1 occupancy year-round in the Bernardo bridge in 2012 and 2013. Temperatures within bat roosts of the Los Lunas bridge (ca. 33 miles north) were monitored from May 2012 to March 2013 (Smith and Stevenson, unpubl. data). December temperatures vary from -3 to 13.9 °C, mean of 4.6 °C; whereas January temperatures vary from -8.2 to 12.2 °C, mean of 2.4 °C. These parallel those temperatures at which Eptesicus fuscus1 (-10 to 20 °C) and Myotis spp. a (-9 to 20) hibernate (Webb et al. 1996); however, many species require thermally stable hibernacula with < 10% oscillation. Additionally, temperatures below 0 °C may exceed the temperature threshold for most southwestern chiropterans (5 - 20 °C, Webb et al. 1996). Bats were absent from the Los Lunas bridge between late October and mid-March; therefore, temperatures may not equally characterize the Bernardo bridge ‘winter roost’. Webb et al. (1996) document bats hibernating at ambient temperatures between -10 and 21 °C, which correspond to New Mexico’s ambient temperatures in December (-11.39 to 18.67) and January (-15.39 and 17). Weaver’s (2012) microclimatic values indicate T. brasiliensis1 bats are selecting colder, less stable environments during winter in central Texas. Hibernation is characterized by torpor bout duration, lower body temperature (may fall to 2 °C), and metabolic suppression. Arousals and the return to euthermy (normal body temperatures) are energetically expensive. For Myotis lucifugus1, each arousal of several hours duration costs 108 mg of fat, the equivalent of 67 days of torpor (Speakman and Thomas 2003). At a natural rhythm of approximately one arousal every 12-15 days, arousals constitute 85% of an individual’s fat depletion through the winter. Therefore, additional arousals due to disturbance (e.g., human activity, bridge construction and maintenance) may reduce their energy supply to the point where survival of the individual is not possible.

conclusion Globally, bat populations are declining. Habitat loss and modification, climatic change, roost availability and disturbance, pesticides and pollution, disease, and human development (e.g., wind turbine facilities, urbanization) cumulatively contribute to population level impacts. Despite their decline, the increase of public awareness and conservation concern, and the

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occurrence of North America’s most devastating wildlife threat, white-nose syndrome; bats remain among the most neglected and misunderstood animals. “The available evidence suggests that bats are essential to ecological balance and forest health, fulfilling the same roles by night as birds do by day” (Lacki et al. 2007). Chiropterans are intrinsic to healthy ecosystems, community integrity, and vital ecological processes. They provide valuable ecosystem services; including arthropod suppression, seed dispersal, pollination (Kunz et al. 2011) and are often considerable contributors to a country’s mammalian diversity. Boyles et al. (2011) estimate the loss of North American bats to the agricultural industry at approximately $3.7 - $53 billion annually. Worldwide, roost availability and abundance are critical elements limiting chiropteran populations.

(O’Shea and Bogan 2003). Bats spend over half their lives within their roost environment, and possess life history traits (e.g., longevity, low reproductive rates) that influence their ability to overcome population declines. Thus, quality roost sites are paramount to the survival and reproduction of an individual, and fundamentally the species. It is evident that manmade bridges function as important components of the roosting ecology and habitat of North American bats. The identification and protection of these key roosting sites can be monumental relative to the longterm viability and conservation of endemic bat populations. North America’s transportation system comprises 582, 976 bridges longer than 6 m (20 ft); of these, 83% traverse waterways. The U.S. Federal Highway Administration estimates an additional 12.5 million

“While insectivorous bats lack the readily definable market value of domestic animals, they have great value both as critical components of ecosystems and for the ecological services they provide.” North American bats are secondary cavity nesters; they rely on preexisting crevices and cavities. Processes of cavity creation (e.g., wood decay, fire, insect activity, excavation) and loss (e.g., deterioration, tree fall) determine the availability and abundance of quality roosts (Sedgeley 2001). In undisturbed forests, these processes achieve an approximate equilibrium; in urban environments, these ecosystem dynamics are absent. Consequently, synanthropic species are further limited by roost availability and abundance than rural or forest inhabiting species. Additionally, bats may occupy specific roosts that provide critical microclimates for different aspects (e.g., night roost, maternity roost, transient roost, hibernaculum). Therefore, only a small subset of available roosts may be suitable and large segments of regional populations may be restricted to few specific roosts during critical times of the year

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smaller structures (e.g., box culverts, drainage structures) that correlate to approximately one structure per quarter mile (400 m). An increasing proportion of national infrastructure currently exceeds or approaches its terminal service life (Doyle et al. 2008); approximately 30% of bridges have classifications of “deficient” (16% structurally deficient, 13.6% functionally obsolete) (Forman et al. 2003). Construction of deficient or new structures offers an exceptional opportunity to communicate and mitigate important biotic effects relative to bat populations (Box 1). Numerous states have become environmental stewards, actively engineering and/or retrofitting bridges to accommodate bat colonies. In the Pacific Northwest, concern relative to the importance of bridges and the sensitive status of several regional bat


Arizona myotis, Myotis lucifugus occultus, a subspecies of the little brown myotis

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species were the impetus for the protection of abandoned wooden bridges on federal lands (United States Department of Agriculture Forest Service and United States Department of the Interior Bureau of Land Management 1994). Similarly, the diminution of available roost sites and the concurrent decline of North American bat populations was the incitement for the Oregon Department of Transportation’s (ODOT) proactive Bat Habitat Enhancement environmental performance standards (EPS). The ODOT’s standard outlines objectives and guidelines, “to maintain, replace, or improve roosting on bridges over waterways” (Barbaccia 2011). The Indiana DOT has developed a ‘Habitat Conservation Plan’ for the endangered Myotis sodalis2 as part of the improvement of transportation facilities around Indianapolis International Airport. The 2012 Indiana Bat Programmatic Biological Assessment and Programmatic Conservation Memorandum of Agreement between the Kentucky Transportation Cabinet, the Federal Highway Administration, and the U.S. Fish and Wildlife Service provides a standardized approach for biological assessments and contributes to the USFWS’s statewide conservation efforts for the Indiana bat (Federal Highway Administration 2013). The Texas, Florida, Georgia, Indiana, Arizona, and California Departments of Transportation are correspondingly developing canonical procedures to determine occupancy, minimize disturbance, and preserve existing or potential roosting opportunities during the management, maintenance, and demolition of bridges.

concerns & recommendations The concept of ‘road ecology’ (i.e., the ecosystem-level effect of

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roads and traffic) has become increasingly popular, and mitigation measures (e.g., wildlife crossing structures; amphibian tunnels, culverts, overpasses, underpasses; EISs) have become important initiatives to increase permeability and habitat connectivity for wildlife inhabiting transportation corridors (Forman et al. 1997). Throughout Europe, Asia, Australia, and North America; hundreds of mitigation structures, or “ecoducts” exist; and concomitantly, numerous documents illustrating their success. However, recommendations and subsequent development of wildlife passageways target cervids, large carnivores, amphibians, and rodents (Animal Road Crossing, arc-solutions.org; Huijser et al. 2007; Clevenger and Huijser 2011). Few recommendations relative to enhancing connectivity for wildlife include bats and fewer still address the effect of infrastructural development on these mammals. Roads may affect bats via 1. vehicle collisions, 2. destruction or degradation of roosts and foraging areas, and 3. severance of critical commuter and migratory routes (Berthinussen and Altringham 2012, Semrl et al. 2012). Bats employ a network of roosts, flight paths, and foraging areas within the landscape. These routes parallel linear landscape elements including treelines, hedgerows, minor roads, and watercourses (Graphic 1). The home ranges of temperate insectivorous bat species typically extend 0.5 - 5 km from their roost, with most species exhibiting high fidelity to roosts, foraging sites, and the commuting flyways that connect them. The probability of vehicle collisions directly relates to landscape structure, and secondarily, to road features (Medinas et al. 2012) and bat ecology predictors. The highest incidence of road casualties occur where roads cross bat flyways (travel corridors), especially at junctions with forest edges and tree alleys (Lesiński 2007). Medinas et al. (2012) indicate that high quality habitats, roost proximity, activity level, and traffic volume are primary elements of

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road casualties. The vulnerability of specific species correlates to flight (i.e., altitude, wing morphology, maneuverability) and foraging ecology (i.e., aerial insectivore versus gleaner, flying time, number of foraging bouts). Species with low flight altitudes that forage close to vegetation or ground (< 10 m), and that orient via guiding landscape structures are most susceptible to vehicle collisions. Mortalities may be high because most species cross at heights that place them in the paths of vehicles. Where tree canopy is high (> 20 m) and within 10 m of the highway, bats cross above traffic; where the canopy is low (< 6 m), bats cross lower and closer to traffic and therefore, probability of vehicle collisions increase (Russell et al. 2009). A study by Zurcher et al. (2010) demonstrates that Myotis sodalis2 alters direction and exhibits antipredator avoidance behavior when vehicles are present. Leziński (2007) determines that juveniles were killed significantly more often than adults, with increases in mortality in late summer (i.e., intense dispersal of young bats). Gaisler et al. (2009) document similar fatality patterns, with increases in July coincident with the occurrence of volant young; and late September - early October, which may correlate to mating and autumnal migration. Leziński et al. (2011) denote that more than half of road casualties occurred in July and August. Medinas et al. (2012) report that female mortality was particularly high in summer; this period encompasses births and lactation, a critical phase in the chiropteran life cycle. This may impact population viability, because newborns are dependent on mothers for survival. Kerth and Melber (2009) attribute lower reproductive success and smaller foraging areas of female Myotis bechsteinii to a major roadway that restricted habitat accessibility. In bats, both reduced reproductive success and increased mortality will profoundly affect local colony size and overall population size. Medinas et al. (2012) affirm “the number of killed


bats reported in our study is at least of the same order of the magnitude of the one often reported for wind farms,” and “bat road-kills should be also an issue of high concern for long-term bat population viability.” These authors emphasize the importance of further studies to clarify the role of bridges in road fatalities, since they may be important structures enhancing or reducing the probability of bat-vehicle casualties. Keeley (National Roads Authority 2005) asserts, “the most significant impact of road construction upon bats is in the clearance phase of the scheme, namely tree-felling, the removal of hedgerows and other vegetation...” These landscape features are important habitat components; commuting routes, essential sources of insect prey, and potential roosts for both crevice- and foliage-roosting species. Additionally, roads may bisect natural flyways, and therefore, perform as barriers or filters to movement, restricting bats from accessing critical resources (e.g., foraging or roosting sites). This has the potential to influence

To minimize collisions and mortalities, alternative roosting structures should be positioned away from roadways. Traditional wooden bat boxes and “condos” do not replicate the thermal dynamics of concrete infrastructures, and novel concrete forms (RD Wildlife Management; Los Lunas, New Mexico) are not yet proven. Therefore, we propose installation of prototype concrete boxes to determine occupancy and successfulness. We recommend both 1. the installation of roosts attached to bridges, and 2. the establishment of free-standing structures within right-of-ways adjacent to these bridges, particularly where bridges span roadways. By affixing temperature and humidity recorders (LogTag Recorders Ltd., New Zealand), we can compare internal microclimates to those of actual concrete bridge roosts (Los Lunas bridge study site; Smith and Stevenson, unpubl. data). The consideration of landscape elements (i.e., flight paths, foraging areas and roosts) prior to construction

“Preservation of critical habitat is as important as preservation of a particular number of individuals.” the abundance and distribution of individuals and populations, both locally and regionally (Bennett and Zurcher 2013). These authors demonstrate that gaps in tree lines, hedgerows, and tree canopies can render commuting routes unsuitable for bats. To improve landscape permeability and connectivity, Bennett and Zurcher (2013) recommend the restoration (e.g., replanting shrubs and trees in gaps), enhancement (encourage interlinking tree canopies through pruning, trimming, and coppicing), and establishment of linear features; including tree lines, hedgerows, and fence lines. Lüttmann (2012) further emphasizes that the incorporation of fence lines and canopy cover at safe altitudes (> 4 m) provide guiding elements and barriers that prevent low flying species from entering traffic.

and/or demolition, and the inclusion of appropriate mitigation measures, will reduce the severity of significant impacts on bat populations. Artificial habitats; including mines, railroad tunnels, and bridges are oftentimes closed, destroyed, or altered without first surveying. Anthropogenic activities (restoration, reinforcement or demolition of structures; landscape modification) that modify habitat parameters and thermal conditions (i.e., temperature, humidity) of hibernacula and roost sites, may cause mortality or site abandonment (Erdle and Hobson 2001). The magnitude of impact varies considerably relative to the period at which these activities are conducted. The periods of pre-volancy (i.e., young are being reared and are unable to fly) and hibernation are

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Graphic 1. Landscape level illustration of the network of flight paths that bats use daily. Different species employ different paths to travel across the landscape. Some bat species (blue) fly from their roost along the lanes, hedges, and close above water. Some species (orange) disperse along all types of guiding structures to all corners of the landscape. Some species (pink) fly from their roost high above the landscape to their feeding area high above the water. Adapted from Limpens et al. (2005).

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Box 1. Several possible mitigation measures to reduce any negative consequences of bridge activities include (National Roads Authority 2006a, Sétra et al. 2009)

i. examination of structure prior to demolition and maintenance activities; if bats are present, exclusion procedures should be established ii. proper timing of demolition and maintenance activities iii. provision of a purpose built structures as an alternative roost(s) adaptations of bridges to “bat friendly” structures; incorporation of roosts into new construction, addition of retrofitted roosts iv. provision of bat boxes; these structures should allow colonization by numerous species and be initiated prior to commencement of construction ( > 6 months) v. addition of lighting to inhibit bat entry onto roads; light intolerance may dissuade bats from transversing commuter routes and therefore, traffic vi. addition of close planting of tall vegetation or barriers (> 4 m) to encourage higher flight paths and hence, fly over roadways (i.e., hopover, Graphic 2)

Graphic 2. With a hop-over, the flight path traverses the roadway at a safe height (Limpens et al. 2005).

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particularly sensitive as individuals are powerless to escape. Similarly, the elimination or diminution of natural environments (e.g., forests, wetlands, hedged farmland, uncultivated land in the right of way) to construct or facilitate these activities can effectively destroy foraging areas and/or create open expanses that constitute physical barriers preventing the movement of bats (Sétra et al. 2009). We became cognizant of the impacts of bridge maintenance activities (i.e., expansion joint replacement, vegetation removal, resurfacing) and the deficiencies of current survey techniques. We have found bats exploiting structures not previously identified (i.e., concrete spalls, insulation bolts, space between timber beams and deck, behind insulation board). Therefore, surveys to determine occupancy prior to demolitions may fail to notice bats utilizing uncommon structures and locations; and thus, timing of demolitions, bat exclusion, and maintenance activities are critical considerations during planning. To minimize negative impacts on bat populations, a comprehensive, statewide bridge survey should be conducted with primary emphasis on those slated for construction or replacement. A survey of New Mexico’s wooden and concrete transport infrastructures would provide guidance for the strategic planning and prioritization of projects and the most appropriate techniques relative to bat species management, opportunities and/or exclusion. Any individual conducting bat surveys should possess a thorough understanding of life history characteristics, various species affected and their ecological requirements. Occupancy can oftentimes be difficult to detect without external signs (e.g., guano, urine staining), and therefore, special efforts to confirm presence / absence may be required. In Ireland, all bat species receive legal protection via the Wildlife Act, 1976; the Wildlife (Amendment) Act, 2000; and Annex IV of the Habitats Directive. Similarly, all species are strictly protected in the United Kingdom via the Wildlife and Countryside Act, 1981; Wildlife Order, 1985; and the Habitats and Species Directive 92/43/EEC. Therefore, the documents ‘Best Practice Guidelines for the Conservation of Bats in the Planning of National Road Schemes,’ ‘Guidelines for the Treatment of Bats During the Construction of National Road Schemes,’ (National Roads Authority 2005, 2006a) and ‘Design Manual for Roads and Bridges’ (Highways Agency et al. 1999) provide the most comprehensive, exemplar recommendations relative to ‘bats in bridges’. At present, we advocate commencing construction when bats are least vulnerable (i.e., after juveniles have dispersed and individuals have not yet commenced hibernation) - prior to parturition (April - May) or following latesummer dispersal (September - October). Current recommendations propose maintenance conducted between mid- October and early April will minimize disturbance. However, we document bat occupancy through consecutive winters and therefore; conclude that bats are overwintering, with bridges functioning as hibernacula (Bernardo bridge). We believe this to be the first documentation of

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a bridge hibernaculum within central New Mexico. The unpredictability of this occurrence necessitates surveys prior to any works to ensure the absence of bats from bridges October through March.

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APPENDIX


Appendix A. Comprehensive mitigation strategies and guidelines to minimize impacts to local bat communities. Given the potential impacts from construction and operational activities, we recommend the employment of appropriate and practicable mitigation and compensation measures to minimize these impacts on bat communities. Due to the unpredictability of winter activity (as far north as Bernalillo county), bridge hibernation (Socorro county, No. 5983) and swallow nest use; every bridge and any proximate works necessitate initiatory surveys to leastwise confirm presence/absence. Species differ relative to sensitivity and exploitation (e.g., roost choice, travel corridors, flight heights, foraging strategy) of the landscape. Therefore, any mitigation measures should endeavor to accommodate the species with the most sensitive requirements or conservation status. 1. Specialist Surveys The survey and assessment of bat occupancy in bridges requires expertise, experience and objectivity. “It is comparatively easy to determine use of a site by bats, but absence is more difficult to prove. It requires greater effort to demonstrate beyond reasonable doubt that bats are not present or likely to be present (Bat Conservation Trust 2007).â€? Surveyors must be mindful and respectful, to prevent roost abandonment and accidental injury to or mortality of bats. Additionally, the surveyor must be competent in identifying bats (may require capture and handling to determine/confirm species) and their respective habitat. The specialist must be able to characterize the existing environment and evaluate its significance. Where mitigation measures are necessary, the specialist must be capable of assisting the design and implementation of these measures. Surveys should identify existing and potential roost sites, map important foraging areas and record principal commuting routes. This valuable information facilitates a strategic plan to be formed to protect local bat populations. All bat surveys should be undertaken at the appropriate time of year to collate the information required (i.e., summer surveys to detect maternity roosts and winter surveys to detect hibernating bats). A preliminary landscape analysis (maps.google.com) can identify probable roost locations and landscape features favorable to the presence or transit of bats (mature forests, large trees, small fields, presence of water and watercourses) prior to site survey(s). Additionally, searches performed prior to site surveys can provide information on the likelihood of vulnerable species being present (i.e., New Mexico species of concern, T & E species) . Surveys to document overall impacts should include right-of-way property; including land necessary for accommodation access, machinery, construction staging, and post-construction maintenance. Caves and mature trees proximate to sites have the potential for various roost types; including maternity roosts, transitional roosts, bachelor roosts, and hibernacula. All species (including those that occupy nearby natural roosts) are vulnerable to poor watercourse management, and removal of treelines and vegetative cover (vulnerable to predators, open spaces are more lit, no protection from wind/rain). Surveys must be sufficient to characterize the local environment and to provide defensible and robust impact predictions. Any sites supporting bat colonies of considerable conservation value may require more exhaustive inspections and documentation. This ensures the proper evaluation of impacts and mitigation measures. To determine movements and the connection of critical areas, surveys may require a broader survey zone. We recommend, when feasible, that comprehensive surveys identify the extent of all significant roosts and breeding sites within 1 km (National Roads Authority 2006b, SĂŠtra 2009). To ensure reliability, several visits across biological seasons should occur (2-3 times per year). If impracticable, the optimum time for surveys is summer. While bats are active throughout the night, peak activity occurs at dusk and before dawn; and surveyors should address bat activity during these time frames to provide comprehensive information of site utilization. The most effective detector survey period is June - August, which will provide information on maternity roosts. Earlier studies (April and May) and later studies (September) will yield information on alternative roosts (National Roads Authority 2006b). Mist netting to capture and identify local bat species that may or may not be identified with bat detectors may be appropriate in certain circumstances (i.e., where detailed information on specific species is required, or where species of concern or high conservation value may occur). In enclosed areas (e.g., bridges), harp-trapping may be employed to confirm the presence of species. 1.1 Standard survey A standard survey to establish presence/absence, assess probability or severity of impact(s), and acquire information to recommend mitigation and/or compensation measures should include: 1.1.1 Date 1.1.2 Site description (includes both location and structure information)

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1.1.3 Proposed activity (demolition, repair, maintenance) 1.1.4 If bats are presently, or have been, within the structure 1.1.4.1 Inspection of existing infrastructure 1.1.4.1.1 Structural fissures (cracked or spalled concrete, damaged or split beams, split or damaged timber railings, et cetera) 1.1.4.1.2 Crevices (expansion joints, space between parallel beams, spaces above supports piers, et cetera) 1.1.2.1.3 Alternative structures (drainage pipes, bolt cavities, open sections between support beams, swallow nests, et cetera) 1.1.4.2 Cursory inspection of natural structures and trees in proposed activity “footprint.� The presence of bats in trees or rock crevices can be difficult without external signs (presence of guano, sounds of bats). Occupancy can be established by examination of suitable crevices, cavities, limb fractures, and loose bark. Specialist equipment (e.g., rope access, borescope) may be required in certain circumstances (advanced survey). 1.1.5 Species present 1.1.6 Roost information including type (e.g., diurnal, nocturnal), location, characteristics

1.1.7 Intensity (e.g., number of bats, time and duration of use) 1.1.8 Photographs to support written documentation

1.2 Advanced survey Survey effort should be proportionate to survey purpose (i.e., to obtain adequate results for specific objectives) and may identify: 1.2.1 Information from standard survey, 1.2.2 Species whose distribution includes site (identify potential for species of conservation concern), 1.2.3 Any features of particular ecological or conservation significance, 1.2.4 Specific roost sites (confirmed and potential) that occur in close proximity to site; detailed inspection of potential tree roosts identified by standard survey, 1.2.5 Any watercourses, flyways, crossing points, or foraging areas that may be impacted by construction and clearance activities, 1.2.6 Potential site-specific mitigation, compensation or enhancement measures, 1.2.7 Colony type and sex. Sexual segregation does occur within habitats of various species. Therefore, the occupation of habitats by males/females should be identified. This may become important (e.g., impact the selection of trees for felling) because a site that sustains females would be more significant than one that support males, 1.2.8 Identify time of survey with respect to biological season. Bat activity may differ between certain periods due to variations in availability of prey, recruitment of juveniles, or the availability of suitable roost sites. For example, summer roosts may not provide the appropriate microclimates necessary for hibernation. Therefore, a survey done outside the breeding season may impart a false impression of the site’s importance, 1.2.9 Bat activity surveys. Appropriate during warmer months (April - September) and at dusk emergence and/or dawn re-entry, and may include documentation of active foraging and commuting habitats. emergence times and locations, intensity (estimate of population), species assessment via manual/automated bat detectors, and camera/video equipement (FLIR, infrared). 2. Health and Safety Recommendations

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2.1 Rabies Before working near known roosts, maintenance crews should be encouraged to avoid disturbing the bats as much as possible, and taught not to handle them. These procedures will minimize the direct exposure of crews to bats and eliminate the possibility of personnel being bitten, which would then necessitate post-exposure rabies treatment. 2.2 Histoplasmosis Bird and bat guano are classic reservoirs for H. capsulatum, the fungus that causes histoplasmosis, a systemic infection primarily of the respiratory tract. Outbreaks have been associated with demolition and earth-moving activities that aerosolize topsoil and dust (e.g., bridge reconstruction and demolition, jack- and air-hammering, waste disposal). Employees should wear personal protective equipment and employ dust-suppression techniques when working in areas potentially contaminated with bird and/or bat droppings (Huhn et al. 2005). 3. Appropriate Time Schedule “A simple project with a defined design concept that will be ready for construction within two years may be programmed for the first two years of the STIP [Statewide Transportation Improvement Program]. The activities conducted in year one can encompass preliminary engineering, environmental certification, right-of-way acquisition, and final design. Year two can include construction” (New Mexico State Highway & Transportation Department 2009). As previously mentioned, Bennett et al. (2008) recommend an inspection interval of 3-5 times annually to determine use. Lengthy time intervals between biological investigations and construction increase the probability of occupancy; and therefore, the unreliability of those evaluations. (Chart 1). 3.1 Exclusion Specific mitigation measures to exclude must be in situ prior to demolition or maintenance activities. 3.1.1 Migratory birds Exclusion practices to prohibit migratory birds with ≥ 3/4” netting does not exclude bats. Any indication of bats necessitates the installation of 3/8” netting. If bats are present, installation should not occur May - August when bats are exceptionally vulnerable to disturbance. If hibernating bats are present, installation can not occur late November - early March. 3.1.2 Bats Bats must be excluded with one-way valves and professional foam sealants and/or 3/8” exclusion netting. Exclusion may occur September - April. If bats are present, installation of one-way valves must occur prior to netting. Winter exclusion must entail a survey to confirm either, 1. bats are absent or 2. present but active (continuously active - not intermittently active due to arousals from hibernation). 3.2 Maintenance Maintenance activities include cleaning activities, preventative maintenance to preserve and lengthen service life, technical and specialized repairs and stream channel maintenance. These activities can involve the operation of support vehicles and equipment, pavement repair, welding and grinding operations, and associated pollutants, which may impact nearby bat colonies. 3.2.1 Minor maintenance activities (e.g., wing wall repair or underpinning foundations) typically have minor or no impact on bats. However, more substantial maintenance operations, including replacement or strengthening of structures above water level, should entail a bat assessment (Bat Conservational Ireland 2010). If bats are present, exclusion procedures should be implemented prior to maintenance activities.

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3.2.2 Application of foam products without prior surveying or professional exclusion can entomb bats. We advocate the preclusion of foam sealants unless a qualified consultant is present and confirms the absence of bats. 3.2.3 Some maintenance activities (e.g., sealing cracks and crevices) may entomb bats or cause the abandonment of non-volant young. Additionally, these activities can create noise pollution, vibrations, and modify thermal conditions of roosts; and consequently, may promote roost abandonment. 3.2.4 Night-time maintenance activities can affect bats. Light, odors and noise can delay or discourage bats from emergence, or potentially, cause site abandonment. Activities adjacent to flyways and roosts should be avoided, especially when bats are most vulnerable (mid-April - end of July). If operations are inevitable, we recommend the installation of very localized lighting in the worksite zone, avoiding surrounding areas to reduce the barrier effect. The temporary erection of noise barriers and/or light screens may also be considered. Temporary infrastructure (stockpile areas, roads for construction traffic) should be constructed at a distance from roosts (Sétra 2009). 3.2.5 Vibrations from noise disturbances within 0.5 miles of a known or suspected hibernaculum may cause arousal from hibernation. The disturbance to hibernating bats reduces the probability of survival because arousals and the return to euthermy (normal body temperatures) depletes imperative fat

reserves (i.e., energy supply).

4. Mitigation Measures This section provides methodology to eliminate, minimize and if possible, remedy significant adverse effects. Where elimination is not possible, measures can be implemented to alleviate the severity of impacts (barriers, plantings). Where impacts cannot be avoided or lessened, it may be possible to compensate for these impacts or restore some aspect of the natural environment to an approximation of its previous condition (planting of native trees and shrubs to compensate for the loss of hedgerows or woodland, habitat creation in areas adjacent to the road (including wetlands), connecting bridges or passages to help link fragmented habitats). 4.1 Avoidance Avoidance of an area, structure, or site with a significant bat presence remains the best mitigation measure for the protection of bats. The significance of the roost may determine the appropriate mitigation measures. 4.2 Guidelines for clearance activities Those species that exploit bridges may equally benefit from proximate tree cavities and natural rock crevices. Therefore, clearance activities within the right of way may cause the destruction of secondary roosts and diminish habitat integrity (e.g., loss of foraging areas and structures that compose flyways). “A break, even a few metres long, in the linear structures that form flyways is likely to reduce or prevent access to foraging areas or more remote roosts” (Sétra et al. 2009). 4.2.1 Conservation of landscape structures The removal of trees and vegetation decreases the availability and abundance of foraging habitat and provisional roosts for individuals occupying bridges, as well as, local bat assemblages that select trees as primary roost sites. Linear landscape features form an important component of the commuting routes for bats, as well as essential foraging sites. Hedgerows and treelines function as “roads,” and migration or long-distance flights may be dependent on these discernable landscape features. USFWS (2007) proposes the protection of land via conservation easement or deed restriction to offset the loss of suitable habitat, particularly near waterways / riparian areas (stream corridors). We advocate native tree plantings to create future habitat and travel corridors and restore connectivity and landscape permeability. Additionally, the control of invasive plant species can further create quality habitat.

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4.2.1.1 Where trees of importance to bats are situated along the periphery of the construction footprint, the potential of retaining these trees should be outlined in the EIS and discussed with relevant personnel prior to site clearance (National Roads Authority 2005). 4.2.1.2 If possible, retain treelines and vegetation adjacent to watercourses. 4.2.1.3 Confine operational activities on watercourses to one side of the channel to minimize damage to the wildlife corridor 4.2.1.4 Protect and identify (tags, flags, et cetera) trees and shrubs to be retained during operational activities 4.2.1.5 Implement a management scheme where removal of important trees and shrubs are replanted with similar or native species within the same year of activities. To minimize the potential for vehicle collisions, roadside plant species should not attract insects and hence, indirectly bats. SĂŠtra (2009) recommends an approximately 10 m wide strip without woody vegetation on either side of the roadway. 4.2.2 Tree-felling Maternity colonies form from late May onwards and remain relatively cohesive through mid- to late August. Single young, born June-July, are non-volant (not capable of flight or evasive action, wholly dependent on mothers) for several weeks and thus, are extremely vulnerable to disturbance by human activities (restoration, reinforcement or demolition of structures). 4.2.2.1 Tree-felling can commence 15 September and continue to 15 March. These months coincide with periods of volant young and hibernation, with the assumption that bats are not hibernating within the site footprint. 4.2.2.1.1 If bats are present, felling should not occur 15 April - 15 September to ensure the protection of maternity colonies and nonvolant juveniles. 4.2.2.1.2 Tree removal activities within 5 miles of a known or suspected hibernaculum can occur between 15 November and 15 March and between 15 September and 15 March within 10 miles of the hibernaculum (USFWS 2007). 4.2.2.2 Immediately prior to felling, trees should be examined for presence/absence of bats and/or other bat activity. A bat specialist should comprehensively inspect trees during daylight hours. Bats rarely roost openly, and are most commonly present within tight spaces and crevices; therefore, a borescope or fiberscope may be necessary for definitive presence/absence. The National Roads Authority (2006b) further recommends a night time detector survey, which should occur from dusk through dawn to ensure that bats do not re-enter the tree. An inspection “confirms the status of the tree only at the time of inspection and where there is a delay of one day or greater the tree must be re-assessed.â€? 4.2.2.3 If exclusion procedures must be implemented, the bat specialist will provide direction relative to the necessary actions and appropriate time periods. 4.2.2.4 If a roost tree must be felled outside the optimum season, the bat specialist must endeavor to remove any bats to safety. Live bats may be released once all tree-felling has concluded. If any deceased bats are found, we encourage the documentation (date, site information) and submittal of carcasses to RD Wildlife Management for species identification and distribution information. 4.2.2.5 We encourage the implementation of mitigation measures to compensate for the loss of tree roosts. 4.2.3 Vegetation removal Bats orient themselves by, and fly parallel to, linear landscape elements; including tall vegetative cover, transition zones (i.e, edge habitat), minor roads and waterways. Bats exploit these linear features to

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locate and capture prey and to safely commute between roosts and feeding areas. The removal of vegetative cover and tree-line, loss of mature trees and disturbance of wetlands all affect the availability of vertebrate prey and foraging sites for bats. A gap of 10 m or more (> 32’) may cause some species to abandon both the commuting route and roosting site.

4.3 Lighting restrictions Artificial light may affect the activity of some species and tolerance may vary from complete avoidance to no effect. Lighting can modify bats’ roosting sites, commuting routes and foraging areas, especially along waterways. Bats commute and forage along dark wildlife corridors (e.g., rivers, canals) and consequently shy away from highly illuminated sections. Therefore, illuminated structures can impede their flight to suitable feeding areas. Lighting along waterways should be avoided at all times. In addition, buffer zones (dark zones) should be included adjacent to waterways. Intensely lit areas may impair bats’ vision and cause disorientation; inhibit movements and prevent access to roosts and foraging areas. Bats may abandon a roost that is illuminated or may considerably delay their emergence. This can reduce foraging times (miss peak levels of insect activity, decrease foraging bouts) and thus, body weight, reproduction and winter survivability. Therefore, it is essential that lighting plans for a development site and around known roosts take into consideration the exit points, flight

paths and foraging areas for bats and ensure the these areas are not illuminated. Light intolerance can be beneficial to dissuade some species from using customary commuting routes that cross traffic and thus, increase the potential for bat-vehicle collisions. Lights positioned close to identified crossing points can deter bats when placed at 10 m intervals.

4.4 Provision of safe crossing structures / points Crossing structures are site-specific movement corridors positioned over roadways that bisect important habitat. Green bridges and underpasses benefit European bat species by permitting them to cross roadways close to established commuting points (Sétra 2009). In certain circumstances, these structures offer a solution to allow bat movement from nearby roosts to foraging areas and alternative roosts. To encourage employment of these structures, plantings should direct bats towards the bridge or underpass. Bats instinctively follow these linear elements to crossing points. Planting of linear corridors should happen early in the construction phase and, where possible, should comprise relatively mature plants, both to ensure these are established quickly and, ideally, so that bats discover these routes before project completion. Valuable treelines or vegetation loss during the clearance phase can be detrimental to bat communities. The installation of hedges/shrubs can direct bats to safe crossing points. The time requirement for plants to grow reduces the effectiveness of this measure. Temporary netting or fencing can therefore be installed to avoid gaps. Hop-overs are created by means of tall trees or 6 m high wire netting with dense low vegetation to encourage bats to fly higher (Graphic 2). Overpasses, when connected to landscape structures, can also direct bats over roadways. The addition of plants along the length of one side, or an opaque windbreak (between 1.5 m and 3 - 4 m) will further enhance this measure (Graphic 3). Additionally, underpasses with lines of vegetation can encourage bats to employ these structures to cross roadways. Wire netting or a screen can also direct bats to enter the tunnel or fly over traffic. The consensus relative to width is “the wider the passage the more it will be used by bats” (Sétra 2009). A height of 4.5 m and width of 4 - 6 m has been recommended to ensure accessibility for all species. The optimal size for culverts is 3 m, with a minimum of 1.5 m (Graphic 4). Simple removable overpass structures consist of wires or wire netting stretched horizontally between two mast on either side of the roadway. These may be installed either temporarily, to appraise the suitability of a location before installing more substantial structures, or as a permanent mitigation measure (Graphic 5).

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4.5 Provisions of alternative roosts * Where sizeable maternity roost(s) are lost, alternative structures (bat houses, boxes) can be built to expiate these impacts. The structure should adequately support the displaced colony with respect to size and thermal value. This measure should also include suitable boxes to satisfy the general requirements of different bat species present year-round. Alternative roosts may provide critical short-term roosting opportunities and should be employed to complement any mitigation measures to protect roosts. The institution of alternative roosts may reduce the impacts to local bat populations and provide safe roosting sites for colonies where natural sites are not available. The erection of these roosts should be initiated prior to commencement of operations (site construction/ demolition) and must be appropriately sited (adjacent to suitable foraging areas). Boxes should be monitored for acceptance and seasonal occupancy, and those that remain vacant for > 2 years should be relocated. Boxes should be installed on bridges with soffit > 1 m above highest recorded water levels, which permits bats to safely drop into flight from roosts. To improve the effectiveness of alternative roosts, we recommend installation two years prior to project commencement. To minimize the potential for vehicle fatalities, particularly for bridge roosts over roadways, we alternately recommend the erection of purpose-built structures that are built within the right-of-way.

* To date, there is no bat box that recreates the thermal capacity, conductivity and microclimatic conditions of bridge roosts and novel concrete forms (RD Wildlife Management) are not yet proven. We propose installation of prototype concrete boxes to determine occupancy and successfulness. We currently have commitments to test concrete boxes within the following structures; bridge over the Pecos River, east of Hagerman (NM 249, Chaves County) and the Galisteo bridge (SR 41, Santa Fe County). Additionally, we will be installing several boxes on bridges within southern AZ (pers. comm. Sandy Wolf on behalf of AZDOT).

4.6 Integral roosts Numerous states have become environmental stewards, actively engineering or retrofitting bridges to accommodate bat colonies. Construction of new bridges offers an exceptional opportunity to incorporate bat roosts at minimal cost (Keeley and Tuttle 1999). At the conceptual design phase, bat specialists can communicate with and advise engineering contractors on appropriate dimensions to accommodate many different species. To minimize vehicle fatalities, we recommended that ‘integral’ roosts not be incorporated into section(s) that will span traffic lanes. The addition of ancillary, nonfunctional drainage pipes with sealed tops would provide integral roosts with minimal or no vandalism.

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Hibernating

J†

D†

Hibernating

F†

Hibernating

Starting to become active

N†

M†

Mating, adding fat stores for winter

Feeding on warmer nights/returning to spring roosts

A

O†

Leaving roosts

Females returning to maternity roosts

M

S Young volant ((ying) and foraging

J

A Females lactating and nursing young

Pregnant females in maternity roosts/early births (e.g., pallid bat births occur late May)

J Young born

Chart 1. Chart for appropriate activity timetables relative to life cycle. Red indicates ‘Stop,’ period when bats are exceptionally vulnerable to disturbance. Yellow represents “Caution,” bats may be vulnerable depending on species and their location within New Mexico. Green indicates “Go,” period when bats should not be present and therefore, not impacted by activities. † The ‘Go’ phase may require a standard survey to confirm absence. Due to the unpredictability of winter activity, bats may be present and/or hibernating.

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Graphic 3. Overpass design features that encourage bats (Setra 2009).

Graphic 4. Vegetation positioned at entrance to encourage bats to fly through the structure (Setra 2009).

Graphic 5. Temporary overpass structures (Setra 2009).

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43

TABLES & MAP


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Table 1. North American bat species known to occupy highway bridges; summarized from Keeley and Tuttle 1999, Hendricks et al. 2004, MacGregor and Kiser 1998, and Kunz and Reynolds 2003.

Species

Inhabits New Mexico

Antrozous pallidus, pallid bat

yes

Choeronycteris mexicana, Mexican long-tongued bat

1, 5

Corynorhinus rafinesquii, Rafinesque’s big-eared bat Corynorhinus townsendii, Townsend’s big-eared bat

yes no

1, 3, 4

yes

Eptesicus fuscus, big brown bat

yes

Lasiurus cinereus, hoary bat

yes

Lasionycteris noctivagans, silver-haired bat

yes

Leptonycteris curasoae, lesser long-nosed bat

1, 4, 5

yes

Macrotus californicus, California leaf-nosed bat

no

Myotis austroriparius, southeastern myotis

no

Myotis californicus, California myotis

yes

Myotis ciliolabrum, western small-footed myotis

yes

Myotis evotis, long-eared myotis

yes

Myotis grisescens, gray myotis

no

4, 5

Myotis leibii, eastern small-footed myotis

2

no

Myotis lucifugus, M. l. occultus; little brown myotis, Arizona myotis

yes

Myotis septentrionalis, northern myotis

no

Myotis sodalis, Indiana bat

2

no

4, 5

Myotis thysanodes, fringed myotis

yes

3

Myotis velifer, cave myotis

yes

Myotis volans, long-legged myotis

yes

Myotis yumanensis, Yuma myotis

yes

Nycticeius humeralis, evening bat Nyctinomops macrotis, big free-tailed bat

no 3

yes

Perimyotis subflavus (formerly Pipistrellus subflavus), tri-colored bat

yes

Pipistrellus hesperus, western pipistrelle

yes

Tadarida brasiliensis, Mexican free-tailed bat

yes

1

New Mexico Listed and Sensitive Species

2

Center for Biological Diversity petition to list, Endangered Species Act

3

Bureau of Land Management Sensitive Species

4

US Fish & Wildlife Endangered Species

5

IUCN Red List, threatened, endangered, or vulnerable species <http://www.iucnredlist.org/>

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Table 2. North American bat species that have the potential to occupy highway bridges; summarized from Keeley and Tuttle 1999; New Mexico distributions from Adams 2003.

Species

Inhabits New Mexico

Corynorhinus townsendii virginianus, Virginia big-eared bat 4

no

Corynorhinus townsendii ingens, Ozark big-eared bat

no

Euderma maculatum, spotted bat

4

yes

3

Idionycteris phyllotis, Allen’s lappet-browed bat Leptonycteris nivalis, Mexican long-nosed bat

1, 3

yes

1, 4, 5

yes

Mormoops megalophylla, Peter’s ghost-faced bat

yes

Myotis auriculus, southwestern myotis

yes

Myotis keenii, Keen’s myotis

no

Eumops glaucinus, Florida mastiff bat

no

Eumops perotis, western mastiff bat (greater bonneted bat)

yes

Eumops underwoodii, Underwood’s mastiff bat

no

Molossus molossus, Pallas’ mastiff bat

no

Nyctinomops femorosaccus, pocketed free-tailed bat

yes

1

New Mexico Listed and Sensitive Species

2

Center for Biological Diversity petition to list, Endangered Species Act

3

Bureau of Land Management Sensitive Species

4

US Fish & Wildlife Endangered Species

5

IUCN Red List, threatened, endangered, or vulnerable species <http://www.iucnredlist.org/>

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Table 3. Western Bat Working Group Priority Matrix and the Natural Heritage Program classifications for bridge roosting species that inhabit New Mexico, including four species that have the potential to occupy these structures (Idionycteris phyllotis, Mormoops megalophylla, Eumops perotis and Nyctinomops femorosaccus; Keeley and Tuttle 1999).

Species

WBWG Priority Matrix †

Natural Heritage Program ‡

Antrozous pallidus, pallid bat

Low

G5 / S5

Choeronycteris mexicana, Mexican long-tongued bat

High

ND

Corynorhinus townsendii, Townsend’s big-eared bat

High

G4 / S2

Eptesicus fuscus, big brown bat

Low

G5 / S5

Euderma maculatum, spotted bat

Medium

G4 / S3

Lasiurus blossevillii, western red bat

High

G5 / S2

Lasiurus cinereus, hoary bat

Medium

G5 / S3

Lasionycteris noctivagans, silver-haired bat

Medium

G5 / S5

Leptonycteris curasoae, lesser long-nosed bat

High

G3 / S1

Leptonycteris nivalis, Mexican long-nosed bat

-

-

Myotis auriculus, southwestern myotis

Medium

G4 / S4

Myotis californicus, California myotis

Low

G5 / S5

Myotis ciliolabrum, western small-footed myotis

Low to Medium *

G5 / S5

Myotis evotis, long-eared myotis

Low to Medium *

G5 / S4

Myotis lucifugus occultus, Arizona myotis

Medium

G5 / S3

Myotis thysanodes, fringed myotis

Medium

G4G5 / S5

Myotis velifer, cave myotis

Low to Medium *

G5 / S4

Myotis volans, long-legged myotis

Low to Medium *

G5 / S5

Myotis yumanensis, Yuma myotis

Low

G5 / S5

Nyctinomops macrotis, big free-tailed bat

Low to Medium *

G5 / S2

Perimyotis subflavus (formerly Pipistrellus subflavus), tri-colored bat §

-

-

Pipistrellus hesperus, western pipistrelle

Low

G5 / S5

Tadarida brasiliensis, Mexican free-tailed bat

Low

G5 / S2

Idionycteris phyllotis, Allen’s lappet-browed bat

High

ND

Mormoops megalophylla, Peter’s ghost-faced bat

Medium

ND

Eumops perotis, western mastiff bat (greater bonneted bat)

Medium

ND

Nyctinomops femorosaccus, pocketed free-tailed bat

Medium

G4 / S1

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† Western Bat Working Group (WBWG) comprises a steering committee, coordinators from 13 western states, and special committees to address specific bat conservation issues. The Regional Priority Matrix provides states, provinces, federal land management agencies, and interested organizations and individuals an understanding of the overall status of bat species inhabiting western North America. ‘G’ represents Global Status, ‘S’ represents State Status, ‘ND’ indicates No Data. ‘High’ ranking indicates that these species should be the highest priority for funding, planning, and conservation actions; these species are imperiled or at high risk of imperilment. ‘Medium’ ranking indicates a level of concern that should warrant further evaluation, research and conservation efforts targeted at both preserving the species and reducing possible threats. ‘Low’ ranking indicates that most of the existing data support stability for populations of the species and that the overall status of the species is secure; conservation actions would still apply for these species, but limited resources should fund ‘High’ or ‘Medium’ ranked species. ‡ University of New Mexico, Natural Heritage Program Numerical Ranking System; 1, critically imperiled or extremely rare; 2, imperiled or very rare; 3, very rare or found in a restricted range; 4, apparently secure; 5, demonstrably secure * Indicates different priority listings for different ecoregions within New Mexico; all associated species above are ‘Low’ in Regions 9 and 10 (9, temperate steppe; 10, tropical / subtropical steppe) and ‘Medium’ in Regions 7 and 8 (7, Arizona - New Mexico mountains semidesert; 8, tropical / subtropical desert). Map of ecoregions from Adams 2003. § Classification systems predate the occurrence of Pipistrellus subflavus in New Mexico (Geluso et al. 2005).

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Table 4. New Mexico bridges known to provide roosting habitat for North American bat species.

NMDOT Bridge Number

Road No.

Milepost

Construction Type

Species Present

Maternity Colony

2503 1, 2

NM 187

31.0

Timber

M. l. occultus, M. yumanensis, T. brasiliensis

Yes

2507

1

NM 187

26.7

Timber

M. l. occultus, M. yumanensis, T. brasiliensis

Yes

1832

1, 2

NM 51

2.2

Timber

M. yumanensis

Yes

1832

2

NM 51

2.2

Concrete

T. brasiliensis

1669

1, 2

NM 187

13.7

Timber

M. yumanensis

Yes

1667

1, 2

NM 187

16.0

Timber

T. brasiliensis, A. pallidus, Myotis spp.

Yes

1669

1

NM 187

13.8

Timber

M. yumanensis, T. brasiliensis

Yes

2591

1, 2

NM 185

14.3

Timber

M. yumanensis, T. brasiliensis

Yes

1831

2

NM 51

3.0

Timber

Myotis spp.

Yes

2509

1

NM 187

25.4

Concrete

M. yumanensis, T. brasiliensis

1660

2

NM 187

23.7

Timber

Myotis spp., T. brasiliensis

2510

2

NM 187

20.5

Timber

Myotis spp.

8540

2

US 550

1.6

Concrete

Myotis spp., T. brasiliensis

7925 / 7926

Bridge Blvd.

-

Concrete

Myotis spp., T. brasiliensis

7898

2

Central Ave.

-

Concrete

Myotis spp., T. brasiliensis

6224

2

NM 500

1.7

Concrete

Myotis spp., T. brasiliensis

7339

2

NM 346

2.3

Concrete

Myotis spp.

8580

2

US 60

167.8

Concrete

Myotis spp.

8286

2

2

US 60

167.6

Concrete

Myotis spp.

5983 / 5984

2

I-25

175.3

Concrete

Myotis spp., T. brasiliensis

5977 / 5976

2

I-25

174.2

Concrete

T. brasiliensis (prior to construction)

6791

2

I-25

83.9

Concrete

T. brasiliensis

1832

2

NM 51

2.2

Timber

A. pallidus, T. brasiliensis

1831

2

NM 51

3.1

Timber

A. pallidus

7544

2

NM 6

18.5

Concrete

T. brasiliensis

7453

2

NM 6

35.6

Concrete

M. l. occultus, M. yumanensis, M. velifer,

Yes

Yes

T. brasiliensis, Myotis spp. Bosque del Apache Union County

3

4

-

-

Concrete

Myotis yumanensis

-

-

Concrete

Perimyotis subflavus

1

Geluso and Mink 2009

2

Smith and Stevenson, pers. obs.

3

Chung - MacCoubrey 1999

4

Geluso et al. 2005

49

Yes


Map 1. New Mexico bridge locations with documented bat colonies.

50


Table 5. Worldwide documentation of chiropteran species that exploit bridges as diurnal roosts.

North America

Information

Antrozous pallidus

AZ, Davis and Cockrum 1963 (concrete highway bridge; open expansion joint; railroad bridge); AZ, Silver Creek Bridge, ca. 14 miles NE Douglas on US 80, Cochise Co.; Canon del Oro Bridge, ca. 10 miles NW Tucson on State 84, Pima Co.; Hwy bridge, 7.3 miles S. St. David on US 80, Cochise Co.; Hwy bridge, 12.1 miles S. St. David on US 80, Cochise Co.; Hwy bridges on Collidge Dam road, S side San Carlos Lake, Graham Co.; Santa Cruz River Bridge, ca. 5 miles NE Nogales on State 82; Santa Cruz Co.; Davis 1966; CA (I-beam bridge) Krutzsch 1946; OR, maternity sites at various bridges and fidelity is evident from year to year, Ferguson and Azerrad 2004; CA, Erickson et al. 2002; from 11 western states (99), Ellison et al. 2003; AZ, Ellison et al. 2003; AZ, Tucson (1), Wolf and Shaw 2002 (concrete); AZ, Hoffmeister 1970 (type not identified); states not identified, [van Zyll de Jong 1985, Lewis 1994, Pierson et al. 1996, Bell 1980, Hermanson and O’Shea 1983, Brown et al. 1997, Tatarian 1999 as cited in Ferguson and Azerrad 2004]; southern AZ, Davis 1969

Choeronycteris mexicana

AZ or NM, Ellison et al. 2003

Corynorhinus rafinesquii

LA, Ferrara and Leberg 2005 (double-T concrete bridges); MS; DeSoto National Forest; Chickasawhay District, DeSoto District; Homochitto National Forest (7), Trousdale and Beckett 2002 (concrete); SC (4), Loeb and Zarnoch 2011 (girder-type); LA (32), Lance et al. 2001, (girdertype bridges); Bennett et al. 2003; 14 southeastern states (12), Ellison et al. 2003; SC (73), Bennett et al. 2008 (concrete; 6 multi-beam, 32 T-beam); states not identified, Keeley and Tuttle 1999; NC, McDonnell 2001 as cited in Bennett et al. 2008; MS, Martin et al. 2011 (18), maternity roosts; MS, DeSoto NF (36), Trousdale and Beckett 2004

Corynorhinus townsendii

CA, Fellers and Pierson 2002 (type not identified); LA, Lance and Garrett 1997 as cited in Trousdale et al. 2008; C. t. townsendii, CA, Erickson et al. 2002; from 20 western states, Ellison et al. 2003; states not identified, Keeley and Tuttle 1999; CA, Pierson and Rainey 1998

Eptesicus fuscus

AZ, Davis and Cockrum 1963 (concrete highway bridge; open expansion joint; railroad bridge); LA, Ferrara and Leberg 2005 (double-T concrete bridges); AZ; Hayward’s Bridge on old Tucson-Nogales highway, ca. 7 miles N of US Border at Nogales, Santa Cruz Co.; Highway bridge, 7.3 miles S. St. David on US 80, Cochise Co.; Highway bridge, 12.1 miles S. St. David on US 80, Cochise Co.; Davis 1966; CA, 2 m east of Bohemia (I-beam highway bridge), Green Valley Falls (I-beam highway bridge), Krutzsch 1946; MT, Hendricks et al. 2004; 2005 (10; 6 concrete, 4 timber) (10); CA, Erickson et al. 2002; from 44 states, Ellison et al. 2003; AZ, Ellison et al. 2003; SC (10), Bennett et al. 2003, 2008 (concrete); FL (35), Gore and Studenroth 2005; states not identified, Keeley and Tuttle 1999; LA, Lance et al. 2001 (type not identified); IL, Feldhamer et al. 2003 (concrete); AZ, Tucson (9), Wolf and Shaw 2002 (concrete); SD, Swier 2003 (concrete); IN, Duchamp et al. 2004; AL, Highway 106 (expansion joint), Richardson et al. 2009

Lasionycteris noctivagans

AZ, Tucson (1), Wolf and Shaw 2002 (concrete)

Lasiurus cinereus

MT, 4.8 km S of Columbus, Hendricks et al. 2004 (wooden girder bridge); unknown state, Ellison et al. 2003

Leptonycteris curasoae

AZ, Ellison et al. 2003; AZ, Hollis 1995

Macrotus californicus

AZ, Davis and Cockrum 1963 (concrete highway bridge; open end); CA, Erickson et al. 2002; southwest (AZ, CA, NV) (4), Ellison et al. 2003

51


Table 5 continued.

North America

Information

Myotis austroriparius

FL, Rice 1957 (drain pipes of railroad bridges, timber bridges), Waccasassa River Bridge on Highway 24; unknown state, majority of reports from FL (15), Ellison et al. 2003; SC (1), Bennett et al. 2003, 2008 (concrete); FL (8), Gore and Studenroth 2005; states not identified, Keeley and Tuttle 1999; LA, Lance et al. 2001 (type not identified); AR, Feldhamer et al. 2003 (expansion joints of concrete bridge); AR, Saugey et al. 2001 (maternity colony of highway bridge); NC, McDonnell 2001

Myotis californicus

CA, Krutzsch 1954 (between supporting beams of wooden bridge); CA, Erickson et al. 2002; from 10 western states, primarily AZ, CA, CO, NV, Ellison et al. 2003; AZ, Tucson (2), Wolf and Shaw 2002 (concrete)

Myotis ciliolabrum

MT, Hendricks et al. 2004, 2005 (2 wooden); CA, Erickson et al. 2002; from 16 western states, primarily SD, WY, CO, ID, Ellison et al. 2003

Myotis evotis

12 western states, primarily CO, MT, OR (9), Ellison et al. 2003; IL, Feldhamer et al. 2003 (concrete)

Myotis grisescens

14 southcentral and southeastern states, primarily MO, AR, AL, KY, Ellison et al. 2003; states not identified, Keeley and Tuttle 1999

Myotis leibii (M. subulatus)

WV (bridge expansion joint), Johnson et al. 2011; Frankfort, KY, J. MacGregor, Kentucky Department of Fish and Wildlife Resources, Frankfort, KY, unpubl. data (guardrail crevices of concrete bridge); NC and SC, O’Keefe and LaVoie 2011 (guardrail crevices of concrete); state not identified, Erdle and Hobson 2001(expansion joints); TN, Erdle and Hobson 2001 (expansion joints); KY, MacGregor and Kiser 1998, 1999 (maternity colonies bridge expansion joints); state not identified, Barbour and Davis 1969, Tuttle 1964 as cited in Johnson et al. 2011, Hitchcock 1955 as cited in Barbour and Davis 1969; AZ, Davis and Cockrum 1963 (concrete highway bridge; expansion joint); NC, Fletcher 2002; WV, Fletcher 2002

Myotis lucifugus

MT, Hendricks et al. 2004, 2005 (highway bridge) (4; 2 concrete, 2 timber); IL, Feldhamer et al. 2003 (concrete); SD, Swier 2003 (type not identified); MT, Bitterroot Valley, Bailey 1926; M. l. occultus; CA, Blythe, Stager 1943 (timber bridge); CA, Erickson et al. 2002; CA, Erickson et al. 2002

Myotis septentrionalis

LA, Ferrara and Leberg 2005 (double-T concrete bridges)

Myotis sodalis

IN (concrete), Mumford and Cope 1958 as cited in Barbour and Davis 1969; from 24 eastern states, primarily IN, KY, NY, PA, Ellison et al. 2003; states not identified, Keeley and Tuttle 1999

Myotis thysanodes

CA, Erickson et al. 2002; from 10 western states, (34), Ellison et al. 2003

Myotis velifer

AZ, Davis and Cockrum 1963 (concrete highway bridge; open expansion joint, open end); AZ; Hayward’s Bridge on old Tucson-Nogales highway, ca. 7 miles N of US Border at Nogales, Santa Cruz Co.; Highway bridge, 7.4 miles NE Pima on US 70, Graham Co., Davis 1966; CA, Erickson et al. 2002; from 7 western states (67), Ellison et al. 2003; states not identified, Keeley and Tuttle 1999; AZ, Tucson (10), Wolf and Shaw 2002 (concrete); AZ, Tylor 2013

Myotis volans

13 western states (23), Ellison et al. 2003

Myotis yumanensis

AZ, Davis and Cockrum 1963 (concrete highway bridge; open end); CA, Stager 1943 (timber bridge); CA, Erickson et al. 2002; from 12 western states, primarily AZ, CA, OR (62), Ellison et al. 2003; AZ, Tucson (2), Wolf and Shaw 2002 (concrete); AZ, Graham County, Barbour and Davis 1969

Nycticeius humeralis

from 15 states (10), Ellison et al. 2003; FL (10), Gore and Studenroth 2005; states not identified, Keeley and Tuttle 1999 52


Table 5 continued.

North America

Information

Perimyotis subflavus

LA, Ferrara and Leberg 2005 (double-T concrete bridges); MS, Homochitto NF, Trousdale and Beckett 2002 (unknown bridge type); SC (26), Bennett et al. 2003, 2008 (concrete); states not identified, Keeley and Tuttle 1999; LA, Lance et al. 2001 (type not identified); IL, Feldhamer et al. 2003 (concrete); NM, Geluso et al. 2005 (concrete); NC, McDonnell 2001

Pipistrellus hesperus

CA, Erickson et al. 2002; from 7 western states, Ellison et al. 2003; states not identified, Keeley and Tuttle 1999; AZ, Tucson (15), Wolf and Shaw 2002 (concrete); AZ, Tylor 2013

Tadarida brasiliensis

TX, Allen et al. 2009 (pre-caste concrete highway bridge); AZ, Davis and Cockrum 1963 (concrete highway bridge; open expansion joint; railroad bridge); TX; Fraze and Wilkins 1990, Sgro and Wilkins 2003 (concrete bridge; expansion joint); CA, SE of Wilbur Springs, Benson 1947; CA, Lakeside, Mitchell 1956 as cited in Bernardo and Cockrum 1962 (unknown bridge type); AZ (St. David, Cochise Co.; Canon del Oro, N Tuscon; Nogales), Bernardo and Cockrum 1962 (unknown bridge type); AZ, Silver Creek Bridge, ca. 14 miles NE Douglas on US 80, Cochise Co.; Highway bridge, 7.3 miles S. St. David on US 80, Cochise Co.; Hayward’s Bridge on old Tucson-Nogales highway, ca. 7 miles N of US Border at Nogales, Santa Cruz Co.; Highway bridges on Collidge Dam road, S side San Carlos Lake, Graham Co.; Davis 1966; CA, Green Valley Falls (I-beam highway bridge), San Pasqual Creek bridge (unknown bridge type) Krutzsch 1946, 1955; TX, Keeley and Keeley 2004 (concrete bridge); CA, Merced Co. (timber bridge), Geluso et al. 1981; NV, (expansion joint of concrete bridge) Hirshfeld et al. 1977; CA, Stager 1943 (timber bridge); CA, McCracken and Gassel 1997; CA, Erickson et al. 2002; from 18 states, primarily AZ, NM, OK, TX (324), Ellison et al. 2003; SC (1), Bennett et al. 2008 (concrete); FL (100), Gore and Studenroth 2005; states not identified, Keeley and Tuttle 1999; AZ, Tucson (32), Wolf and Shaw 2002 (concrete); TX (3), Keeley and Keeley 2004 (concrete), (3) Turmelle 2009; AR (4), Saugey et al. 2001 (type not identified, 1 day roost in a narrow space beneath a metal platform on the State Highway 70 bridge); AZ, Tylor 2013; TX, Winter Garden region (5), Cleveland et al. 2006; TX, Balcones Escarpment (2), Horn and Kunz 2008

species not identified

AZ, Holbrook, Leroux Wash Bridge (Arizona Game and Fish Department); WA, Methow Valley; Morse Flat Bridge, McFarland Bridge; AZ, Phoenix, 40th St. and Camelback Road (Lowery 2012)

International

Carollia perspicillata

southeastern Brazil, Mendes et al. 2011

Rhynchonycteris naso

southeastern Brazil, Mendes et al. 2011

Saccopteryx bilineata

Guatemala, (unknown bridge type), Dickerman et al. 1981

Mormopterus francoismoutoui

Mascarene Islands (expansion fissures of concrete bridge), Goodman et al. 2008

Nycteris grandis

Africa; Guinea (bridge under road), Fahr et al. 2006

Rhinolophus hillorum

Africa; Guinea (small concrete bridge under road), Fahr et al. 2006

Rhinolophus hipposideros

Crimea (Ukraine), Ševčík et al. 2011; Portugal, Reis and Rufino 2012; Austria, Reiter et al. 2012

Rhinolophus dentii

Jones 1992, Grubb et al. 1999; as cited in Kock et al. 2002

Pipistrellus kuhli

Portugal, Reis and Rufino 2012

53


Table 5 continued.

International

Information

Myotis daubentonii

southern Ireland, Flavin et al. 2001; Ireland, Masterson et al. 2008; Slovakia (2), Cel’uch and Ševčík 2008 (2.5 cm wide and 15 cm deep vertical crevice between bridge beams); Ireland, Carden et al. 2010 (“frequently found in bridges); Ireland; Charles Bridge (2) (5 arch sandstone), Upper Lake (5 arch sandstone), Newcastle Bridge (arch limestone and sandstone), Drumcar Bridge (arch stone bridge w/ concrete base), O’Daly’s Bridge (6 arch sandstone), Rampart (single arch limestone), Donaghpatrick Bridge (6 arch sandstone and concrete); Italy, Abruzzo (4), Russo 2002

Pipistrellus pipistrellus

northeast Scotland, Rydell et al. 1996 (unknown bridge type); Kosicka kotlina basin, Cel’uch and Ševčík 2008 (within drainage tubes and crevices between concrete girders; UK, Yorkshire Dales National Park, Senior et al. 2005 (type not identified); Portugal, Reis and Rufino 2012

Pipistrellus pygmaeus

Portugal, Reis and Rufino 2012

Rousettus aegyptiacus

Lavrenchenko et al. 2004 (colony of 10,000 in 20 m long bridge made of 20 concrete sections)

Barbastella barbastella

Portugal, Reis and Rufino 2012

Hipposideros ruber

Ethiopia, Lavrenchenko et al. 2004 (under a bridge ceiling)

Hipposideros caffer

Ethiopia, Lavrenchenko et al. 2004 (type not identified)

Miniopterus africanus

Ethiopia, Lavrenchenko et al. 2004 (type not identified)

Tadarida teniotis

Huelva, Spain (expansion joints), Ibanez and Perez-Jorda 1998; Portugal, Reis and Rufino 2012

Plecotus auritus

Portugal, Reis and Rufino 2012

Plecotus austriacus

Portugal, Reis and Rufino 2012

Eptesicus isabellinus

southern Iberian Peninsula, Papadatou et al. 2011 (highway bridge); Portugal, Reis and Rufino 2012

Eptesicus serotinus

Portugal, Reis and Rufino 2012

Myotis nattereri

central Scotland, Swift 1997 (unknown bridge type); Ireland, Masterson et al. 2008; Ireland, Drumcarra Bridge, Donaghpatrick Bridge (arch sandstone and concrete)

Myotis myotis

country not identified, Cel’uch and Ševčík 2008 (maternity colonies in hollow chambers)

Myotis macropus (formerly M. adversus) New South Wales, Law and Urquhart 2000, Law et al. 2001; Australia, Thomson 1998 (timber) Myotis blythii

Italy, Molveno, Chirichella et al. 2003 (in cracks); Slovakia, Cel’uch and Ševčík 2008 (vertical shafts and narrow pipes)

Mops midas

South Africa, Smithers 1983 as cited in Dunlop 1999 (“report of roosting habits shows them roosting head up, packed together tightly, in the joints of a concrete bridge; Bavaria, Germany, nursery colony with hollow space of concrete bridge, Zahn 1999

Mops condylurus

Mozambique, Reside 2007 (type not identified)

Emballonura semicaudata

Fiji (100 individuals under a bridge of unknown type), Hutson et al. 2001

Nyctalus noctula

Germany, Cel’uch and Ševčík 2008 (hibernation colony); Slovakia, Cel’uch and Ševčík 2008

Nyctalus leisleri

Portugal, Reis and Rufino 2012

Nycteris macrotis

Gambia, Kock et al. 2002 (unidentified bridge)

Rhinolophus landeri

Gambia, Kock et al. 2002 (unidentified bridge)

Carollia perspicillata

country not identified, Mendes et al. 2011

Trachops cirrhosus

country not identified, Mendes et al. 2011

Desmodus rotundus

country not identified, Mendes et al. 2011

Species not identified

Hong Kong (unknown bridge type), Hutson et al. 2001 54


Table 6. Worldwide documentation of chiropteran species that exploit bridges as nocturnal roosts. Nocturnal roosts possess several functions; including energy conservation, protection from predators and inclement weather, locations for information transfer, sociality, and consumption and digestion of prey.

North America

Information

Antrozous pallidus

AZ, Continental highway bridge No. 1, 3.7 miles N of Continental, Pima Co., C 1; OR, Lewis 1994 (wooden bridge, cement bridges with I-shaped support beams, cement bridges with ‘box’ support structures); state and bridge type unidentified Lewis 1994, Barbour and Davis 1969, Keeley and Tuttle 1996, Pierson et al. 1996 from Ferguson and Azerrad 2004; CA, Erickson et al. 2002; British Columbia, Rambaldini 2006 (open girder concrete highway bridge)

Eptesicus fuscus

OR; Adam and Hayes 2000, Waldien and Hayes 1999; CA, Erickson et al. 2002; SD, Swier 2003 (stone bridge); OR, Portland (2), Perkins 2003

Corynorhinus townsendii

OR, Adam and Hayes 2000, Waldien and Hayes 1999; C. t. townsendii, CA, Erickson et al. 2002

Lasionycteris noctivagans

OR, Adam and Hayes 2000

Myotis californicus

OR, Adam and Hayes 2000; OR, Waldien and Hayes 1999; CA, Erickson et al. 2002; CA, Krutzsch 1954

Myotis ciliolabrum

CA, Erickson et al. 2002

Myotis evotis

OR, Adam and Hayes 2000, Waldien and Hayes 1999; CA, Erickson et al. 2002; OR, Waldien 1998; ID, Nez Perce National Forest, Perkins 1992 (concrete)

Myotis lucifugus

OR, Adam and Hayes 2000, Waldien and Hayes 1999; CA, Erickson et al. 2002; OR, Portland (2), Perkins 2003; M. l. occultus, CA, Erickson et al. 2002

Myotis thysanodes

OR, Adam and Hayes 2000, Waldien and Hayes 1999; CA, Erickson et al. 2002

Myotis volans

OR, Adam and Hayes 2000, Waldien and Hayes 1999; CA, Erickson et al. 2002

Myotis yumanensis

OR, Adam and Hayes 2000; Sinaloa, Mexico, Miller and Allen 1928 as cited in Barbour and Davis 1969; OR, Waldien and Hayes 1999; CA, Erickson et al. 2002

Tadarida brasiliensis

NV, (expansion joint of concrete bridge) Hirshfeld et al. 1977; CA, Erickson et al. 2002

species not identified

WA, Methow Valley; McFarland Bridge; AZ, Preacher Canyon; Highway 260 Bridge (Arizona Game and Fish Department)

International

Eptesicus isabellinus

Iberian Peninsula (highway bridge), Papadatou et al. 2011

Nycteris thebaica

Swaziland, South Africa (culvert, beneath roads and railway tracks, underside of a low bridge over a dry stream, Monadjem et al. 2009

Myotis daubentonii

55

Italy, Russo 2002


Table 7. Worldwide documentation of chiropteran species that exploit culverts.

North America

Information

Choernycteris mexicanus

AZ, Keeley and Tuttle 1999

Corynorhinus rafinesquii

NC, Roby et al. 2011; LA, Lance et al. 2001, Bennett et al. 2008; MS, Martin et al. 2011 state not identified, Harvey 1997

Myotis austroriparius

states not identified, Keeley and Tuttle 1999; FL, Rice 1957; southeastern states (15), Ellison et al. 2003

Myotis grisescens

states not identified, Keeley and Tuttle 1999; state not identified, Ellison et al. 2003

Myotis leibii

from 16 states, Ellison et al. 2003

Myotis septentrionalis

from 31 states, Ellison et al. 2003

Myotis sodalis

from 24 eastern states, Ellison et al. 2003

Myotis velifer

TX, Jackson et al. 1982, Keeley and Tuttle 1999

Perimyotis subflavus

state not identified, Moore 1949 as cited in Barbour and Davis 1969

Tadarida brasiliensis

states not identified, Keeley and Tuttle 1999; TX, Fraze and Wilkins 1990

Myotis californicus mexicanus

Mexico, north Guadalajara, Jones et al. 1970

Hylonycteris underwoodi

Mexico, Jalisco, Phillips and Jones 1971

Micronycteris megalotis

Mexico, Jalisco, Phillips and Jones 1971

Sturnira lilium

Mexico, Jalisco, Phillips and Jones 1971

Artibeus jamaicensis

Mexico, Jalisco, Phillips and Jones 1971

Artibeus lituratus

Mexico, Jalisco, Phillips and Jones 1971

International

Macrophyllum macrophyllum

Guatemala, Dickerman et al. 1981

Glossophaga spp.

Guatemala, Dickerman et al. 1981

Glossophaga soricina

Panama, Klite and Kourany 1965

Carollia spp.

Guatemala, Dickerman et al. 1981

Carollia perspicillata

Panama, Klite and Kourany 1965

Rhynchonycteris naso

Guatemala, Dickerman et al. 1981

Myotis nattereri

central Scotland, Swift 1997 (unknown bridge type)

Myotis daubentonii

Italy, Abruzzo, Russo 2002; Netherlands, Boonman 2011

Nycteris macrotis

Gambia, Kock et al. 2002 (3); Mozambique, Reside 2007

Nycteris thebaica

Swaziland, Monadjem 2005

Miniopterus schreibersii

Australia, Sydney, White 2007

Rhinolophus hillorum

Africa; Guinea, Fahr et al. 2006

56


Table 8. Worldwide documentation of chiropteran species that exploit birds’ nests.

North America

Information

cliff swallow, Petrochelidon pyrrhonota

Eptesicus fuscus; TX, Jameson (1959) as cited in Barbour and Davis 1969 Tadarida brasiliensis; TX, Buchanan 1958

barn swallow, Hirundo rustica

Eptesicus fuscus; AZ, O’Shea and Vaughan 1999 Myotis velifer; TX, Buchanan 1958; TX, Jackson et al. 1982 (H. rustica) in concrete box culverts; Milstead and Tinkle 1959; Tinkle and Patterson 1965; Pitts and Scharninghausen 1986 as cited in Kunz and Lumsden 2003; Keeley and Tuttle 1999 Myotis yumanensis; AZ, O’Shea and Vaughan 1999

cave swallow, Petrochelidon fulva

Myotis velifer; southwestern US, Jackson et al. 1982, Pitts and Scharninghausen 1986, Ritzi et al. 1998 as cited in Kunz and Lumsden 2003

International †

fairy martin, Hirundo ariel

Tadarida australis, Hyett and Shaw 1980 Mormopterus planiceps; Hyett 1980, Hyett and Shaw 1980 Nyctophilus geoffroyi; Hall and Richards 1979, Hyett 1980, Hyett and Shaw 1980, Lumsden and Bennett 1995, Maddock and Tidemann 1995, Schulz 1997 Nyctophilus gouldi, Schulz 1997 Nyctinophilus timoriensis; Hyett 1980, Hyett and Shaw 1980 Chalinolobus dwyeri; Hyett 1980, Hyett and Shaw 1980, Hoye and Dwyer 1995, Schulz 1997 Chalinolobus gouldii, Shulz 1997 Chalinolobus morio; Hyett 1980, Hyett and Shaw 1980, Holsworth 1986, Lumsden and Bennett 1995, Shulz 1997 Myotis spp., Schulz 1997 Miniopterus schreibersii; Schulz 1997 Vespadelus troughtoni, Schulz 1997

grey-crowned babbler, Pomatostomus temporalist

Chalinolobus gouldii; Brown et al. 1978, Beruldsen 1980, Pizzey 1980

fernwren, Oreoscopus gutturalis

Murina florium; Shulz and Hannah, in press; Schulz 1997

yellow-throated scrubwren, Sericornis citreogularis

Murina florium; Shulz and Hannah, in press Species not identified, Allen 1939 Species not identified, Schulz 1997 Species not identified, Schultz 1997 Kerivoula papuensis, Schulz 1995

white-browed scrubwren, Sericornis frontalis

Species not identified, Allen 1939

Nectariniidae

Kerivoula spp. (Vespertilionidae); Sclater 1900, Shortridge 1934, Roberts 1951, Rosevear 1965, Brossett 1966, Sharma 1986, Skinner and Smithers 1990

57


Table 8 continued.

International â€

Information

brown gerygone, Gerygone mouki

Kerivoula papuensis, Schulz 1995

white-rumped swiftlet, Collocalia spodiopygia

Taphozous australis, Schulz 1991 Species not identified; Harrison 1976, Tarburton 1986

Baya weaverbird, Ploceus philippinus

Kerivoula picta, Sharma 1986

†All references taken from Schulz 1997

58


59

PHOTO GALLERY


60


gallery 1

1 3

2

1 bolt cavity Myotis spp. utilizing a unique location , bolt cavities with pre-insulated pipes

2 drainage _pipe When drainage pipes are clogged, they function as roosts

3 _concrete spall Myotis spp. roosting in the void created by concrete spalling

61


4_ concrete spall Myotis spp. roosting in the void created by concrete spalling

5 _open beam Myotis spp. roosting in the "open" beams between centermost girders

6 internal expansion joint T. brasiliensis colony in left corner of internal expansion joint between centermost girders, "double beam"

5

4 6

62


7 9

8

7 _internal expansion joint T. brasiliensis colony roosting in internal expansion joints above column caps. Crevice is completely full of roosting bats that they are roosting outside the crevice

8 internal expansion joint Even outside the crevice , bats are protected from diaphragm "walls"

9 crevice crevice formed between concrete slabs, approximately 1/4" width

63


10_ wasp's nest Myotis spp. roosting in a wasp's nest built within the framework of a deteriorated swallow nest

11 _swallow nests Bats (visible) exploiting two abandoned swallow nests.

12 swallow nests Bats roosting inside intact swallow nests

11

10 12

64


13 15

14 13 insulated pipe collar A significant amount of guano is present in these structures, but further visibility is minimal

14 concrete slab bridge Individuals roosting behind blue insulation board. Typically, this bridge (slab concrete with corrugated metal on the underside of deck) would be immediately identified as incapable of supporting bats

15 crevice between box beams High occupancy rates force bats to roost in the ends immediately above riprap, appoximately .45 m from ground to roost entrance (Bridge No. 5983)

65


16 Bridge No. 5983 roosts are location between concrete "beams" and the space above the column cap (inaccessible)

17 _Bridge No. 8580 this bridge serves as an important night roost, indicated by significant urine staining and guano below roost

18 Bridge No. 8540 bats occupy crevice between center beams, entire length of bridge is roosting habitat

17

16 18

66


gallery 2

1 3

2

1 double beam Bats roost in narrow crevices where parallel beams have separated

2 _"wedge" Bats use spaces above vertical supports between spans

3 _above beam In this picture, bats are present in the narrow space between beams and deck

67


4_ guardrail post Bats roost between guardrail posts and outermost beams

5 open beam Five Myotis sp. roosting on the vertical face of an open beam; at least 3 individuals (only ears and heads are visible)_ are emerging from between the deck and beam (area corresponds to picture 3)

6 above beam Several individuals roosting between beam and deck; space was made available due to shims

5

4 6

68


gallery 3

1 2

1 consequence of tarring during hibernation period Group of bats dead from maintenance activities at bridge functioning as a hibernacula

2 _vehicle fatalities Numerous bats on roadway due to vehicle collisions

69


3_ mortalities Mortalities from unknown cause occurred in late January, another conclusive indication that bats are using this bridge as a hibernacula

4 mortalities same event as 3

3 4

70


71

LITERATURE CITED


72


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