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Bat rabies in Oklahoma.

The early history of the occurrence of bat rabies in the United States has been summarized (Sulkind and Greve, 1954; Venters et al., 1954; Baer and Adams, 1970; Constantine, 1979). The prevalence of bat rabies in the United States is summarized annually in the Journal of American Veterinary Medicine Association (e.g., Blanton et al., 2006; Dyer et al., 2013). Until a review of bat rabies in Oklahoma by Caire (1998), the only summaries of bat rabies in the state were the Oklahoma State Department of Health (OSDH) epidemiological reports and those of Glass (1959), Calhoun (1989), Graves and Spocogee (1992), and Caire (1993). The objective of this paper is to summarize the occurrence of bat rabies in Oklahoma from 1997-2012 and to compare those years to the previous 10 y discussed by Caire (1998).

MATERIALS AND METHODS--The OSDH receives bats from veterinarians, public agencies, and concerned citizens for rabies testing (direct fluorescent rabies antibody test), which occurs within 2-3 days of being submitted. Since 1987, the OSDH has been encouraged to save for identification those bats submitted for rabies testing. Because of changes in personnel during the years 1987-1996, and the poor condition of some bats, not all of the bats submitted to the OSDH during those years were saved. However, most bats submitted from 1997-2012 were saved for identification. Some could not be identified because they were not saved or only brains and heads in poor condition were submitted. Bats were kept frozen at the OSDH until identified by Caire. Nomenclature follows Wilson and Reeder (2005) for scientific and vernacular names except for Parastrellus and Perimyotis, which we used instead of Pipistrellus (following Hoofer et al., 2006). All bats identified from 1987-1996 were incinerated after identification. Most bats from 1997-2012, except those testing positive for rabies which were incinerated, were kept and are deposited in the University of Central Oklahoma Natural History Museum. One exception was a Eumops perotis that tested positive for rabies, submitted in 2010. Because this was the first occurrence of this species in Oklahoma, it was released to the University of Central Oklahoma Natural History Museum where it was accessioned into the collection as a state voucher specimen (Caire and Loucks, 2012).

RESULTS AND DISCUSSION--There have been no summaries of the occurrence of rabies in bats in Oklahoma since Caire (1998). Prior to 1998, Glass (1959) reported rabid Brazilian free-tailed bats (Tadarida brasiliensis) among 1,338 specimens of 13 species he collected in 1956; no species other than T. brasiliensis were reported to have rabies. Calhoun (1989) and Graves and Spocogee (1992), in their reviews of rabies in Oklahoma, did not list which species of bats did or did not have rabies.

Caire (1998) reported 40 bats tested positive for rabies in Oklahoma from 1987-1996, averaging 4 per year and ranging from zero in 1988 to a high of six in 1989 and 1991. The number of rabid bats from 1997-2012 was 63, averaging four bats per year, and ranging from zero in 2008 to 12 in 2009. These values were similar to those from 1987-1996. Over the 26-y period, the number of rabid bats that were identified to species was 94; 32 in the Caire (1998) study and 62 in the current period. One individual was not identified to species because it was not saved. Of the 1,061 bats submitted and tested, the average number of rabid bats per year was 4 (9.7% rabid), ranging from zero in 1988 and 2008 to 12 in 2009.

The following 11 species were tested for rabies and reported by Caire (1998): Nyctinomops macrotis (submitted in 1977), T. brasiliensis, Eptesicus fuscus, Lasiurus borealis, Lasiurus cinereus, Nycticeius humeralis, Parastrellus hesperus, Perimyotis subflavus, Antrozous pallidus, Myotis grisescens, and Myotis velifer. The Eastern red bat, L. borealis, occurred most frequently (261; 72%) among the 363 identified bats. Some could not be identified because only brains were submitted or the bats were not saved. Four species (E. fuscus, L. borealis, L. cinereus, and P. subflavus) had individuals that tested positive for rabies (Caire, 1998).

From 1997-2012, 10 species were tested: E. perotis, T. brasiliensis, E. fuscus, L. borealis, L. cinereus, N. humeralis, P. subflavus, Lasionycteris noctivagans, M. grisescens, and M. velifer. Lasiurus borealis was again the most-common species (301; 50%) among the 607 identified and tested bats. Some could not be identified because only brains were submitted or the bats were not saved. Eight species (E. perotis, T. brasiliensis, E. fuscus, L. borealis, L. cinereus, N. humeralis, P. subflavus, M. velifer) had individuals that tested positive for rabies (Table 1).

Eight species (T brasiliensis, E. fuscus, L. borealis, L. cinereus, N. humeralis, P. subflavus, M. grisescens, M. velifer) were common to both the previous and current study (Table 1). Three species (N. macrotis, P. hesperus, A. pallidus) reported in Caire (1998) were not among those species tested from 1997-2012. From 1997-2012, two additional species, E. perotis and L. noctivagans, were tested for rabies. Four species, E. fuscus, L. borealis, L. cinereus, and P. subflavus, were common to both periods and had individuals that tested positive for rabies.

Over the 26-y period, 970 individuals were identified and L. borealis composed 58% (562) of the 1,061 bats tested. The total number of male and female bats that were submitted and tested for rabies from 1987-1996 was not listed by Caire (1998) due in part to not all the bats being saved by the OSDH; however, the number of rabid specimens of E. fuscus, L. borealis, L. cinereus, and P. subflavus submitted from 1987-1996 were pooled and a significant difference was found to exist in the frequency of rabies between male and female bats: more males than females were rabid (Caire, 1998). From 1997-2012, there was a significant difference (log-likelihood ratio test with Yates continuity correction, Gc = 10.660; P < .0025) between the number of males (253) and the number of females (333) that were submitted; the sex was determined and they were tested for rabies. There was not a significant difference in the total number of male and female bats (24 of 253 males, 9.5%; 36 of 333 females, 10.8%) that tested positive for rabies from 1997-2012 (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 0.6516; P > 0.30).

Of the 170 L. borealis identified from 1987-1996, 46 (27.1%) were males and 124 (72.9%) were females. There was a significant difference in the number of males and females submitted for rabies testing (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 36.5970; P < 0.001). Caire (1998) reported there was a significant difference between the number of males (11/46 males; 23.9%) and females (10/124; 8.1%) that were positive for rabies. Of the 301 L. borealis identified from 1997-2012, five did not have the sex determined because of the condition of the specimen. Of the remaining 296, 98 were males (33.1%) and 198 were females (66.8%). There was a significant difference in the number of males and females submitted for rabies testing (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 34.3133 P < 0.001). There was not a significant difference in the number of male (13/98 males; 13.3%) and female (24/ 198 females; 12.1%) L. borealis that tested positive for rabies (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 0.0211; P > 0.90).

Over the 26-y period, E. fuscus was the second most common species, comprising 24.5% (238) of 970 identified bats. During the reporting period 1997-2012, E. fuscus was more-frequently submitted and tested than in the previous reporting period 1987-1996 (33% and 10%, respectively). Although the actual numbers of male and female E. fuscus were not listed, Caire (1998) reported no significant difference in the number of male and female E. fuscus that tested positive for rabies from 1987-1996. Among the 201 E. fuscus identified and tested for rabies from 1997-2012, 104 were males (51.7%) and 87 were females (43.3%). There was not a significant difference in the number of males and females submitted (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 1.3388; P > 0.20). Among the E. fuscus, 5/104 males (4.8%) and 5/87 females, (5.7%) were rabid. No significant difference was detected between the number of male and female E. fuscus that tested positive for rabies (log-likelihood ratio test with Yates continuity correction, [G.sub.c] = 0.0084; P > 0.95).

Of the bats identified, 95% from 1987-1996 and 91% from 1997-2012 were submitted from April-October (Table 2). This is the time of year when bats are most active in Oklahoma (Caire et al., 1989). June and July, the months when most pups are born, had the highest number of submissions: 60.5% from 1987-1996 and 55.5% from 1997-2012. Over the 26-y period, 92.7% of the bats identified were submitted from April-October, with June and July being the months with the most bat submissions, 57%. Unknown species also were listed by month during the current study period, although unidentified bats were not listed by month in the previous study (Caire, 1998), resulting in a total of 959 bats over the 26-y period.

During June and July, most female bats are either nursing or pregnant in Oklahoma. In June and July from 1987-1996 (Table 2), 203 bats were submitted, with 176 being L. borealis (86.7%). Of the 176, 27 were females carrying pups; 71 total pups averaging three pups per female and ranging from 1-5. As reported by Caire (1998) for the 1987-1996 period, none of the female bats submitted with pups, nor any of the pups, submitted from 1997-2012 tested positive for rabies. Lasiurus borealis was also the most-frequently submitted species (Table 2) in June and July from 1997-2012 (226/346 bats submitted, 65.3%). Of the 226, 14 were females carrying pups: 54 total pups averaging four per female and ranging from 1-4. One possible explanation of why L. borealis are more-frequently found by the public and submitted to the OSDH for rabies testing is that the females of this tree bat are sometimes blown from tree roosts on windy days and might not be able to regain flight from the ground due to the additional weight of the pups (Sparks and Choate, 2000).

Rabid bats probably do occur in all Oklahoma counties. From 1997-2012, the 63 bats reported as rabid by the OSDH were from 28 counties in Oklahoma, with 39.7% from three counties: Tulsa, 13; Oklahoma, six; and Cleveland, six. Caire (1998) reported that the 138 rabid bats from 1967-1996 were from 42 counties with 65 (47%) in the same three counties: Tulsa, 31; Oklahoma, 25; and Cleveland, nine. Over the 26-y period, rabid bats have been reported from 49 of Oklahoma's 77 counties, with the majority of them being from the same three counties. These three counties contain the largest urban centers and overall are more-heavily populated, and encounters between bats and people are more likely in populated areas than in rural regions.

Based on data compiled from the OSDH from 2003-2012 for all mammals that tested positive for rabies in Oklahoma, bats comprised a smaller percentage, 5.2%, than did skunks and other wildlife, 68.6%; livestock (cattle, horses, goats), 13.6%; and dogs and cats, 12.6%. However, Dyer et al. (2013) indicated that most human rabies cases in the United States are due to bat exposures. The infection of humans with rabies from bats in Oklahoma is an infrequent occurrence. Krebs et al. (2005) and Dyer et al. (2013) reported one individual in Oklahoma who died in 2004 from bat rabies. However, this was after the man had received a liver transplant from a donor who had died of rabies after an encounter with a rabid T brasiliensis. The only other record of a human death from a bat rabies variant in Oklahoma was an individual who died in 1979 without a known animal bite exposure (Messenger et al., 2002). How he contacted the virus is unknown; however, he was a spelunker and could have contacted a bat in a cave.

The incidence of rabies in bats submitted to the OSDH in Oklahoma has not varied much over the last 26 y. Continued monitoring of bat rabies in Oklahoma should provide information related to variations and frequency of rabies among bat species in Oklahoma as well as its etiology and epizoology within the state. The identification of bats submitted to state health departments will continue to add to our knowledge of bat ecology and distributions. Without the annual identification of bats at the OSDH, from 1987-1996 new state records of N. macrotis would have gone unnoticed as well as records of the endangered species M. grisescens. Likewise, from 1997-2012 we identified the first record of E. perotis in Oklahoma and additional records of M. grisescens. Other uncommon species (e.g., L. cinereus and L. noctivagans) also show up among the bats submitted to state health departments. As suggested by Caire (1993), a clear understanding of bat rabies in Oklahoma remains incomplete and will remain so until an unbiased study of rabies in natural populations can be made. Biologists are encouraged to continue to monitor the occurrence of bat rabies in other states and on a regional scale.

We thank the individuals at the Oklahoma State Department of Health for their cooperation, the time spent preserving the bats for identification, and the retrieval of data from early records. The manuscript was improved because of comments from G. M. Caddell.

LITERATURE CITED

BAER, G. M., and D. B. ADAMS. 1970. Rabies in insectivorous bats in the United States, 1953-65. Public Health Reports (1896-1970) 85:637-645.

BLANTON, J. D., J. W. KREBS, C. A. HANLON, and C. E. RUPPRECHT. 2006. Rabies surveillance in the United States during 2005. Journal of the American Veterinary Medical Association 229:1897-1911.

CAIRE, W. 1993. Bat rabies in Oklahoma. Oklahoma State Department of Health Epidemiology Bulletin 93:4-5.

CAIRE, W. 1998. Bat rabies in Oklahoma. Proceedings of the Oklahoma Academy of Science 78:59-65.

CAIRE, W., and L. S. LOUCKS. 2012. The greater mastiff bat, Eumops perotis (Chiroptera: Molossidae) in Oklahoma. Southwestern Naturalist 58:102.

CAIRE, W., J. D. TYLER, B. P. GLASS, and M. A. MARES. 1989. Mammals of Oklahoma. University of Oklahoma Press, Norman, Oklahoma.

CALHOUN, J. 1989. Annual review of animal rabies, Oklahoma, 1988. Oklahoma State Department of Health Epidemiology Bulletin 89:1-4.

CONSTANTINE, D. G. 1979. An updated list of rabies-infected bats in North America. Journal of Wildlife Diseases 15:347-349. Dyer, J. L., R. Wallace, L. Orciari, D. Hightower, P. Yager, and J. D. Blanton. 2013. Rabies surveillance in the United States during 2012. Journal of the American Veterinary Medical Association 243:805-815.

GLASS, B. P. 1959. Recovery of rabies virus from the Mexican freetail bat in Oklahoma. Proceedings of the Oklahoma Academy of Science 39:83-84.

GRAVES, T., and M. SPOCOGEE. 1992. Rabies in Oklahoma, 1991. Oklahoma State Department of Health Epidemiology Bulletin 92:1-3.

HOOFER, S. R., R. A. VAN DEN BUSSCHE, and I. HORACEK. 2006. Genetic status of American pipistrelles (Vespertilionidae) with description of a new genus. Journal of Mammalogy 87:981-992.

KREBS, J. W., E. J. MANDEL, D. L. SWERDLOW, and C. E. RUPPRECHT. 2005. Rabies surveillance in the United States during 2004. Journal of the American Veterinary Medical Association 227:1912-1925.

MESSENGER, S. L., J. S. SMITH, and C. E. RUPPRECHT. 2002. Emerging epidemiology of bat-associated cryptic cases of rabies in humans in the United States. Clinical Infectious Diseases 35:738-747.

SPARKS, D. W., and J. R. CHOATE. 2000. Distribution, natural history, conservation status, and biogeography of bats in Kansas. Pages 173-228 in Reflections of a naturalist: papers honoring Professor Eugene D. Fleharty (J. R. Choate, editor). Fort Hays Studies, Special Issue 1, Hays, Kansas.

SULKIND, S. E., and M. J. GREVE. 1954. Human rabies caused by bat bite. Texas State Journal of Medicine 50:620-621.

VENTERS, H. D., W. R. HOFFERT, J. E. SCATTERDAY, and A. V. HARDY. 1954. Rabies in bats in Florida. American Journal of Public Health 44:182-185.

WILSON, D. E., and D. M. REEDER (Editors). 2005. Mammal species of the world: a taxonomic and geographic reference. 3rd ed. Smithsonian Institution Press Washington, D.C.

Submitted 13 January 2014.

Acceptance recommended by Associate Editor, Troy A. Ladine, 2 May 2014.

WILLIAM CAIRE, * LYNDA SAMANIE LOUCKS, AND PHOEBE L. BROWN

Biology Department, University of Central Oklahoma, Edmond, OK 73034 (WC, LSL)

4304 Rancho Drive, Oklahoma City, OK 73119 (PLB)

* Correspondent: wcaire@uco.edu
TABLE 1--Oklahoma bat species identified and the
number and percentage of each species reported
as rabid from 1987-2012.

Species                     1987-1996

                             n    rabid n     %

Eumops perotis                0      0       0
Tadarida brasiliensis        14      0       0
Eptesicus fuscus             37      4      10.8
Lasiurus borealis           261     23       8.8
Lasiurus cinereus             8      3      37.5
Nycticeius humeralis         20      0       0
Parastrellus hesperus         1      0       0
Perimyotis subflavus          9      2      22.2
Antrozous pallidus            1      0       0
Lasionycteris noctivagans     0      0       0
Myotis grisescens             4      0       0
Myotis sp.                    0      0       0
Myotis velifer                8      0       0
Unknown                      74      8      10.8
Total                       437     40       9.15

Species                     1997-2012

                             n    rabid n     %

Eumops perotis                1      1      100
Tadarida brasiliensis        14      2       14.3
Eptesicus fuscus            201     12        6
Lasiurus borealis           301     37       12.3
Lasiurus cinereus            15      6       40
Nycticeius humeralis         40      1        2.5
Parastrellus hesperus         0      0        0
Perimyotis subflavus         10      2       20
Antrozous pallidus            0      0        0
Lasionycteris noctivagans     1      0        0
Myotis grisescens             3      0        0
Myotis sp.                    2      0        0
Myotis velifer               19      1        5.3
Unknown                      17      1        5.9
Total                       624     63       10.1

Species                     Total

                              n     rabid n     %

Eumops perotis                  1     1       100
Tadarida brasiliensis          28     2         7.1
Eptesicus fuscus              238    16         6.7
Lasiurus borealis             562    60        10.7
Lasiurus cinereus              23     9        39.1
Nycticeius humeralis           60     1         1.7
Parastrellus hesperus           1     0         0
Perimyotis subflavus           19     4        21
Antrozous pallidus              1     0         0
Lasionycteris noctivagans       1     0         0
Myotis grisescens               7     0         0
Myotis sp.                      2     0         0
Myotis velifer                 27     1         3.7
Unknown                        91     9         9.9
Total                       1,061   103         9.7

TABLE 2--The number of Oklahoma bat species identified
each month, 1987-2012. Abbreviation for each species is:
Ep, Eumops perotis; Tb, Tadarida brasiliensis; Ef,
Eptesicus fuscus; Lb, Lasiurus borealis; Lc, Lasiurus
cinereus; Nh, Nycticeius humeralis; Ph, Parastrellus
hesperus; Ps, Perimyotis subflavus; Ap, Antrozous
pallidus; Ln, Lasionycteris noctivagans; Mg, Myotis
grisescens; Ms, Myotis septentrionalis; Mv,
Myotis velifer; Un, Unknown.

Year    January   February   March   April   May    June

1987      --         --       --     1 Ef    1 Lb   20 Lb
          --         --       --     1 Lb     --     --
1988      --         --       --      --      --    1 Ef
          --         --       --      --      --    3 Lb
          --         --       --      --      --    1 Nh
1989      --         --       --      --      --    7 Lb
          --         --       --      --      --     --
1990     1 Ef        --       --     1 Lb    2 Ef   1 Ef
          --         --       --     1 Tb    1 Lb   6 Lb
          --         --       --      --     1 Lc    --
1991      --         --      1 Tb    1 Lb    1 Ef   1 Ef
          --         --       --      --     3 Lb   9 Lb
          --         --       --      --      --    1 Nh
1992      --         --      1 Ef     --     1 Lb   1 Ef
          --         --      1 Lb     --      --    24 Lb
          --         --       --      --      --    1 Tb
1993     1 Nh        --       --     1 Lb    1 Mg   17 Lb
          --         --       --      --     2 Lb   3 Ef
          --         --       --      --     1 Tb    --
1994      --         --       --      --     1 Ef   1 Lb
          --         --       --      --     2 Lb    --
          --         --       --      --      --     --
          --         --       --      --      --     --
1995      --         --       --      --     1 Lb   6 Lb
          --         --       --      --     1 Nh    --
          --         --       --      --     1 Ps    --
1996      --         --      1 Ps    1 Lb     --    10 Lb
          --         --       --      --      --     --
          --         --       --      --      --     --
          --         --       --      --      --     --
1997     1 Ef        --      1 Ef    1 Lb    1 Ef   4 Lb
          --         --      1 Mv     --      --    1 Nh
          --         --       --      --      --     --
1998      --         --       --     1 Ef    1 Nh   1 Ef
          --         --       --      --      --    1 Nh
          --         --       --      --      --    6 Lb
1999      --         --       --      --     1 Ef   4 Ef
          --         --       --      --     1 Lb   17 Lb
          --         --       --      --     1 Un   1 Mg
          --         --       --      --      --    1 Mv
          --         --       --      --      --    1 Nh
          --         --       --      --      --    1 Tb
          --         --       --      --      --    1 Ps
          --         --       --      --      --    2 Un
2000     1 Un       1 Ef     2 Ef     --     1 Ef   1 Ef
          --         --      2 Nh     --     2 Lb   7 Lb
          --         --       --      --      --    2 Un
          --         --       --      --      --     --
2001     1 Ef       1 Ef      --     1 Ps    1 Lb   3 Ef
          --         --       --      --      --    7 Lb
          --         --       --      --      --     --
2002      --         --       --      --      --    5 Ef
          --         --       --      --      --    12 Lb
          --         --       --      --      --    1 Un
          --         --       --      --      --     --
          --         --       --      --      --     --
          --         --       --      --      --     --
2003      --         --      1 Lb    1 Lb    3 Ef   3 Ef
          --         --       --      --     1 Lb   23 Lb
          --         --       --      --     1 Mv   1 Lc
          --         --       --      --      --    1 Un
2004     1 Ef        --       --     1 Ef    1 Ef   3 Lb
          --         --       --      --     2 Lb    --
          --         --       --      --      --     --
          --         --       --      --      --     --
2005    1 M sp       --       --     1 Lb    2 Ef   2 Ef
          --         --       --     1 Un    2 Lb   10 Lb
          --         --       --      --      --     --
2006     1 Ef       1 Ef      --     1 Ef    1 Ef   6 Ef
          --         --       --     1 Lb    3 Lb   4 Lb
          --         --       --      --      --    2 Nh
2007      --        1 Nh     1 Nh    2 Ef    1 Ef   5 Ef
          --         --       --     1 Nh    1 Lb   27 Lb
          --         --       --      --      --     --
2008      --        1 Ef     1 Ef    1 Ef    1 Ef   1 Ef
          --         --       --      --      --    3 Lb
          --         --       --      --      --     --
2009     1 Nh       2 Ef     1 Ef    1 Ef    1 Ef   3 Ef
          --         --       --     1 Ps     --    3 Lb
          --         --       --     3 Tb     --    1 Nh
          --         --       --      --      --     --
2010     1 Nh        --      1 Ef    1 Lb    3 Lb   2 Ef
          --         --      1 Lb     --      --    16 Lb
          --         --       --      --      --     --
2011      --         --       --      --     1 Ef   5 Ef
          --         --       --      --     1 Lb   10 Lb
          --         --       --      --     1 Mv    --
2012     1 Ef        --      1 Lb    1 Ef    3 Ef   14 Ef
          --         --       --     1 Lb    9 Lb   11 Lb
          --         --       --     2 Tb     --    2 Nh
          --         --       --      --      --     --
Total     11         7        17      30      67     350

Year    July    August   September

1987    5 Lb     2 Lb       --
        1 Mg     2 Ps       --
1988    8 Lb     1 Ef      1 Lb
        1 Lc     1 Lb      1 Lc
         --      1 Nh       --
1989    4 Lb     1 Ap      3 Lb
         --       --       1 Lc
1990    1 Ef     2 Ef      3 Lb
        15 Lb    3 Lb      2 Mv
         --       --       1 Ps
1991    1 Ef     1 Ef      1 Ef
        4 Lb     4 Lb      1 Nh
        2 Nh     1 Nh       --
1992    2 Lb     2 Ef      7 Lb
        1 Mv     1 Lb      2 Lc
         --      1 Mg      1 Nh
1993    10 Lb    1 Tb      1 Ph
        1 Nh      --        --
        1 Tb      --        --
1994    12 Lb    1 Ef      1 Ef
         --      1 Lb      3 Lb
         --      1 Nh      1 Lc
         --      1 Tb       --
1995    10 Lb    1 Ef      1 Ef
        1 Ps     1 Lb      1 Lb
         --      1 Nh       --
1996    1 Ef     1 Ef      2 Lb
        6 Lb     1 Lb      1 Nh
        1 Mv      --        --
        2 Ps      --        --
1997    6 Lb     1 Lb      1 Lc
        1 Nh     1 Mv       --
        1 Mg      --        --
1998    4 Lb     1 Tb       --
         --       --        --
         --       --        --
1999    4 Ef     3 Ef      1 Lb
        4 Lb     2 Lb       --
        1 Nh     1 Un       --
        1 Tb     1 Nh       --
         --       --        --
         --       --        --
         --       --        --
         --       --        --
2000    2 Ef     3 Ef      3 Ef
        4 Lb     1 Lb      1 Lb
        1 Mv     2 Nh      1 Ps
         --      2 Un       --
2001    4 Lb     3 Ef      1 Ef
        3 Lc     2 Lc      1 Ps
         --      1 Nh       --
2002    3 Ef     2 Ef      1 Ef
        1 Lb     2 Lb      3 Lb
        1 Nh     1 Nh      1 Ps
         --      1 Mv      1 Lc
         --      1 Mg       --
         --      1 Ps       --
2003    5 Lb     1 Ef      1 Ef
         --      3 Lb       2Lb
         --      1 Mv      1 Nh
         --       --        --
2004     --      3 Ef      1 Ef
         --      1 Lb      1 Mv
         --      1 Tb       --
         --      1 Mv       --
2005    2 Ef     2 Ef      2 Ef
        6 Lb     1 Nh      1 Lb
         --      1 Tb       --
2006    2 Ef     1 Ef      1 Mv
        2 Lb     1 Nh      1 Un
         --      1 Un       --
2007    2 Ef     1 Ef      2 Ef
        1 Lb     1 Lb      1 Lc
        2 Nh      --        --
2008    7 Ef     2 Ef      1 Ef
        5 Lb      --        --
        2 Lc      --        --
2009    1 Ef     5 Ef      11 Lb
        13 Lb   1 M sp     1 Lc
        1 Nh     2 Nh      1 Nh
         --       --       1 Ps
2010    2 Ef     3 Ef      2 Ef
        2 Lb      --       1 Lb
        1 Lc      --        --
2011    2 Ef     5 Ef      2 Ef
        4 Lb      --       1 Mv
         --       --       1 Tb
2012    3 Ef     2 Ef      1 Lb
        2 Lb      --        --
        1 Nh      --        --
         --       --        --
Total    199     105        87

Year    October   November   December

1987      --         --         --
          --         --         --
1988      --         --         --
          --         --         --
          --         --         --
1989     2 Lb       1 Lb        --
         1 Lc        --         --
1990     1 Ef       1 Ef        --
         3 Mv        --         --
          --         --         --
1991     1 Ef        --         --
         1 Lb        --         --
         1 Nh        --         --
1992     1 Lb       1 Lb       1 Ef
          --         --         --
          --         --         --
1993     3 Lb       1 Lb        --
         1 Nh        --         --
          --         --         --
1994     2 Ef       1 Lb        --
         1 Mg        --         --
          --         --         --
          --         --         --
1995     1 Ef       1 Tb        --
         1 Tb        --         --
          --         --         --
1996     1 Nh       1 Ef        --
          --        1 Lb        --
          --        1 Mv        --
          --         --         --
1997     1 Mv        --         --
         1 Lb        --         --
         1 Un        --         --
1998      --        1 Lb       1 Ef
          --        1 Un        --
          --         --         --
1999     1 Un        --        1 Ef
          --         --         --
          --         --         --
          --         --         --
          --         --         --
          --         --         --
          --         --         --
          --         --         --
2000     1 Lb       2 Lb       3 Ef
         2 Ps        --         --
          --         --         --
          --         --         --
2001     1 Ef       1 Ef        --
         3 Mv        --         --
          --         --         --
2002     1 Ef       1 Lb       1 Ef
         1 Mv       1 Mv       1 Nh
          --         --         --
          --         --         --
          --         --         --
          --         --         --
2003      --        1 Nh        --
          --         --         --
          --         --         --
          --         --         --
2004     1 Nh       1 Ef        --
         1 Ln       1 Lc        --
          --         --         --
          --         --         --
2005     1 Ef       1 Ef        --
          --        1 Lb        --
          --         --         --
2006     1 Ef       1 Ef       1 Tb
          --         --         --
          --         --         --
2007     1 Mv       1 Ef        --
         1 Nh       1 Nh        --
          --         --         --
2008     1 Ef        --         --
          --         --         --
          --         --         --
2009     1 Ef        --         --
         1 Lb        --         --
          --         --         --
          --         --         --
2010     1 Ef       1 Ep       1 Ef
          --         --         --
          --         --         --
2011     1 Tb        --         --
          --         --         --
          --         --         --
2012     3 Ef        --         --
         1 Lc        --         --
         1 Nh        --         --
         1 Tb        --         --
Total     51         25         10
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Author:Caire, William; Loucks, Lynda Samanie; Brown, Phoebe L.
Publication:Southwestern Naturalist
Article Type:Report
Date:Dec 1, 2014
Words:4032
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