Mammals of Indiana

Mammals RUSSELL E. MUMFORD and JOHK O. WHITAKER, JR. of Indiana 07438 BLOONIKGTOIV INDIANA UNIVERSITY PRESS 134 Order / Insectivora • Family / Talpidae Table 44 Ectoparasites and other associates of Condylura cristata (n = 21) from Indiana Parasites Hosts parasitized Parasites Total Average Total Percent Fleas (Siphonaptera) Ctenophthalmus pseudagyrtes 13 0.62 6 28.6 Mites (Acarina) other than chiggers Orycteroxenus canadensis 5,586 266.00 14 66.7 Haemogamasus ambulans 20 0.95 8 38.1 Eadiea condylurae 10 0.48 4 19.0 Androlaelaps fahrenholzi 9 0.43 5 23.8 Pygmephorus sp. 6( + ) 0.29 6 28.6 Pygmephorus identified P. moreohorridus 1 Haemogamasus liponyssoides 4 0.19 3 14.3 Eulaelaps stabularis 1 0.05 1 4.8 Hirstionyssus talpae 1 0.05 1 4.8 Cyrtolaelaps sp. 1 0.05 1 4.8 Macrocheles sp. 1 0.05 1 4.8 Euryparasitus sp. 1 0.05 1 4.8 Anoetidae 1 0.05 1 4.8 star-nosed mole is of grasses or leaves, about 5 to 7 inches in diameter, and a flattened sphere placed above high water, usually in a little hillock or knoll, often beneath a stump or log. We recorded testis size for twelve Indiana star-nosed moles. From this small sample, the four males with the largest testes (12 to 19 mm in length) were collected on 15 February, 22 March, and 28 April. All of seven taken in mid-May, one in June, one in September, and two in November had testes ranging from 3 to 6 mm in length. It would appear from these meager data that copulation takes place mainly in March and April. PARASITES. Wilson (1961) examined one Indiana Condylura from Steuben Countyfor external parasites and found two specimens of the flea Ctenopthalmus pseudagyrtes. We have examined 21 star-nosed moles from Indiana for external and internal parasites; all but one had ectoparasites (Table 44). By far the most abundant fonn (as many as 1,000 per host) was the host-specific hypopial glycyphagid mite Orycteroxenus canadensis. The next most common species, in descending order, were the laelapid mite Haemogamasus amhulans, the flea Ctenopthalmus pseudagyrtes, and the laelapid mite Androlaelaps fahrenholzi. Small numbers of the host-specific myobiid mite Eadiea condylurae and other species of mites were also found, including species of Pygmephorus: P. faini, P. horridus, P. moreohorridus, P. spinosus, and P. whitakeri. None of 18 moles examined harbored endoparasites. DECIMATING FACTORS. Two Indiana specimens were captured by cats and one by a dog. Several others, found dead, were probably dropped by predators. Mumford once found a saw-whet owl perched with a decapitated star-nosed mole in its talons, in southern Michigan. TAXONOMY. The subspecies in Indiana is Condylura cristata cristata (Linnaeus). SELECTED REFERENCES. Hamilton, 1931b; Jackson, 1915. Order CHIROPTERA Family Vespertilionidae Little Brown Myotis Myotis lucifugus (LeConte) Little Brown Bat, Cave Bat Vespertilio gryphus: Allen, 1894 Vespertilio subulatus: Blatchley, 1897 Myotis subulatus: Elliott, 1907 Myestis lucifugus: Van Gorder, 1916 Myotis lucifugus: Hahn, 1909 DESCRIPTION. The little brown myotis is a small, brownish bat with relatively short ears. It is easily confused with several other species of Myotis, or with the evening bat (Nycticeius humeralis), the eastern pipistrelle (Pipistrellus subflavus), and even the big brown bat (Eptesicus fuscus). In the hand, an examination of the teeth will enable one to determine if he is dealing with a species of Myotis, for in this genus there is a space containing two tiny molariform teeth between the canine and the four large, posterior molariform teeth (see Fig. 10). Myotis lucifugus can be separated from M. sodalis (the Indiana myotis), with which it is most often confused, by the fact that sodalis has dark grayish pink, duller fur, smaller feet, and a distinct keel on the calcar (see Fig. 1; keel can be seen in fresh specimens, not in study skins); also in sodalis the hairs on the toes are shorter and do not extend to the tips of the claws. Keen's myotis (M. keenii) is distinguished from other Indiana Myotis by its longer ears and by the tragi, which are frequently curved and pointed (see Fig. 2). The gray myotis (M. grisescens), a rare bat in Indiana, has dorsal fur that is the same color from base to tip; the gray myotis is usually grayish, but females may be brownish in summer. The forearm of grisescens averages about 44 mm in length; and that of other Myotis in Indiana ranges from 35 to 40 mm. The southeastern myotis (M. austroriparius) has a more dense, woollier fur than other species in Indiana; the color above is usually gray, with some individuals in late summer exhibiting a brownish cast; the belly fur is usually white-tipped, but may have a tan to pale brownish tinge on certain individuals. The foot is relatively large in austroriparius and live animals have a pinkish nose (other than M. sodalis and M. austroriparius, other Myotis in Indiana have dark noses). Myotis lucifugus in Indiana ranges in color from tan to chocolate brown; adults have shiny fur, but immatures have duller, darker pelage than adults. Several color variations may occur in a single maternity or hibernating aggregation. We have examined a little brown myotis specimen that has white wing tips and a white venter. White-blotched individuals may occur from time to time, as they do in other species of mammals. 135 136 Order / Chiroptera • Family / Vespertilionidae Little brown myotis. Photo by Merlin Tuttle Weights and measurements are shown in Table 45. The dental formula is I —C — P—M— = 38. 3 13 3 STATUS AND DISTRIBUTION. The little brown myotis is common and is found throughout Indiana (Map 14). It is the second most abundant bat (after the Indiana myotis) in hibernating quarters, but possibly less abundant than the big brown bat and the red bat (Lasiurus borealis) in the summer. Numerous breeding colonies have been found, ranging from Martin, Lawrence, Jackson, and Franklin counties northward (Cope et al., 1961; Humphrey and Cope, 1976). Animals occupying the noncave regions of Indiana withdraw from those areas in the fall, and many little brown bats hibernate in the caves of southern Indiana (see Map 5). The northernmost wintering record is from an old coal mine in Turkey Run State Park (Parke County), where a single male was found 12 February 1962. The largest known winter populations of Myotis lucifugus have been recorded in Coon's Cave and Grotto Table 45 Weights and measurements of Myotis lucifugus from Indiana Males Females Total length (mm) n 39 41 X 83.7 84.3 range 72-94 66-91 SD 6.4 6.2 SE 1 1 Tail length (mm) n 40 41 X 33 33 range 26-43 24-48 SD 4.5 4.6 SE 0.7 0.7 Hind foot (mm) n 39 40 X 8.7 8.5 range 7-11 5-11 SD 1.5 1.4 SE 0.2 0.2 Weight (grams) n 37 36 X 6.15 6.15 range 3.1-10.0 3.2-14.4 SD 1.7 2.5 SE 0.3 0.4 Cave (Monroe County), Big Wyandotte Cave (Crawford County), and Ray's Cave (Greene County), but numerous other caves in southcentral Indiana have occasional winter concentrations of up to 100 or more little brown bats. There is evidently considerable movement of M. lucifugus out of Indiana in the fall, for the numbers known from the breeding colonies are greater than the numbers known to hibernate. As with most species of bats in Indiana, population decreases are evident over the past twenty years. Humphrey and Cope (1976) published winter population estimates for four Indiana caves (Table 46). HABITAT. During spring and fall migration and in the summer, the little brown myotis has been encountered by day in buildings, picnic pavilions, on the sides of buildings, beneath shutters, in caves or mines, beneath concrete bridges, and in a wooden covered bridge. At dusk we have shot individuals flying about the borders of Myotis lucifugus • Little Brown Myotis 137 Map 14. The Little Brown Myotis, Myotis lucifugus, in Indiana Table 46 Winter population estimates of Myotis lucifugus in four Indiana caves (from Humphrey and Cope, 1976) Ray's Grotto Wyandotte Coon's Year Cave Cave Cave Cave 1960-61 2,000 — 900 1961-62 2,680 1,525 350 — 1962-63 — — 175 — 1963-64 1,800 — 150 225 1964-65 5,600 1,377 900 — 1965-66 1,300 — 500 350 1966-67 1,800 1, ISO 550 — 1967-68 70 — 140 — 1968-69 350 190 24 — 1969-70 575 266 4 — wooded areas and along streams lined with trees, in Jackson, Tippecanoe, and Vigo counties. Except for one record from a mine, all wintering bats of this species have been found in caves. Of 36 Indiana caves surveyed for wintering M. lucifugus, 27 harbored this bat. Hibernation sites in the caves "were relatively uniform and stable in microclimate, being cool and humid with almost no air flow" (Humphrey and Cope, 1976). All known breeding colonies have been in buildings; the bats are most often found in attics, but also roost behind shutters, behind the large, wooden, sliding doors of barns, and in other protected places. A group of 15 little brown myotis found beneath the loose bark of an American elm on 22 May could have been a nursery colony (Humphrey and Cope, 1976). ASSOCIATED SPECIES. The most common bats that hibernate in caves with the little brown myotis are the Indiana myotis, the big brown bat, and the eastern pipistrelle. Other bats that utilize caves along with the little brown bat are Keen's myotis and the southeastern myotis. Of these species, we have found the little brown bat clustered only with the Indiana myotis and the southeastern myotis. In summer maternity colonies, we have noted the big brown bat and (once) the southeastern myotis with the little brown myotis. Myotis lucifugus has been shot at dusk while foraging with Keen's myotis, the Indiana myotis, the red bat, the big brown bat, and the eastern pipistrelle. A few little brown bats, Keen's myotis, and Indiana myotis roosted in the daytime beneath a large concrete bridge in Turkey Run State Park; a maternity colony of big brown bats also used this bridge. HABITS. The little brown myotis is colonial; large congregations are found in hibernation, and assemblages of females (plus some males) form maternity (nursery) colonies. In summer, relatively few adult males are found, although a few roost in caves and some occupy nursery colonies. It would be most interesting to know where adult males roost in the summer and whether they form groups or become more solitary. From about November to April, the little brown myotis is found almost exclusively in caves. The females leave the caves earlier in the spring than do the males, and begin to congregate in buildings to bear their young; most bats of both sexes have left the caves by 1 June. A very small number of little brown bats (nearly all males) inhabit certain caves 138 Order / Chiroptera • Family / Vespertilionidae during June and July; for example, only four males were found on eighteen June and July visits to twelve caves known to be used for hibernation by M. lucifugus. Evidently, when one finds this species in caves during these months, most of the animals are males. Adult male little brown bats were caught in nets set across the entrance to Donnehue's Cave (Lawrence County) on the following nights: 30 June, 5 bats; 14 July, 1 bat; 26 July, 6 bats. We do not know why M. lucifugus, and other species of bats, can be netted at cave entrances during these months, even when they do not inhabit the caves during the daytime. In August, individuals begin returning to the caves, building up to their winter peak in December. As a rule, hibernating little brown bats do not form huge, compact clusters. A common pattern of roosting groups is to fonn irregular, more or less horizontal rows on the side walls of the caves. This species, however, does hang from cave ceilings and squeezes into both vertical and horizontal crevices. Wintering groups are composed of both sexes, and other species of Myotis may be present in the groups. Many little brown bats also hang singly in the caves; lone individuals may wedge themselves into tiny holes in the cave ceiling just large enough to house a single bat. There appears to be little activity by M. lucifugus in the hibernating quarters from November to March; one male was netted on 2 December at a cave entrance. Emergence from day roosts during the nonhibernating season occurs relatively late in the evening. On 18 June, practically none of the members of a nursery colony left the attic until the light outside was so dim that shooting the bats was impossible. This may explain why so few little brown myotis have been obtained by bat hunting at dusk in Indiana. As with other species of bats, time of emergence is closely correlated with time of sunset and with light intensity. In a study of bat activity at the entrance of Donnehue's Cave, we caught no little brown bats in our nets until after sunset, although both Keen's myotis and pipistrelles entered the nets before sunset. Only 16 of 178 little brown myotis netted were caught within the first hour after sunset. The extensive use of buildings and other man-made structures for roosting and maternity sites by M. lucifugus and other bats certainly reveals how these species have adapted since man came on the scene. One wonders whether little brown bats gathered to produce young in trees (perhaps they still do) or made more use of caves in earlier days. It seems evident that buildings provide excellent conditions for the formation of nursery colonies and the rearing of young. One factor may be the warm temperatures that prevail in tire attics of houses during the summer. (Caves are relatively cool in summer.) Also, houses afford more protection from natural predators. It is of interest that none of the twelve species of bats that occur in Indiana are known to breed in caves in the state. On 18 May, a cage of little brown myotis was removed from a cave for banding. Some of the bats were banded near the cave and released before a rain shower began. As tbe cage with the rest of the squeaking bats was carried one-fourth mile to an automobile, several bats flew about the investigator's head. When he reached the car and paused for a few minutes, four bats circled about and two even lit momentarily on his jacket. All four of these bats were banded and must have been attracted by the noisy animals in the cage. MIGRATION. Since 1946, many thousand little brown myotis have been banded in Indiana to help researchers learn something about movements, longevity, and other aspects of this bat's natural history. Data from the recapture of banded individuals have not been spectacular, but some information has resulted from this work. Humphrey and Cope (1964) reported the movements of 47 little brown myotis from a maternity colony at Thorntown (Boone County) over a six-year period. Banded bats were recaptured in eight caves in Indiana and two caves in Kentucky, at distances up to 200 miles from the banding site. The general direction of movement was mostly due south-north. Half of the recoveries were made in four caves in Monroe County, Indiana, about 70 miles south of the breeding colony. Band recoveries from studies by Mumford reveal roughly the same type of movement—a general southward movement in fall and a general northward Myotis lucifugus • Little Brown Myotis 139 movement in the spring. In both of these banding studies, a few bats were retaken to the east and west of the usual line of movement, indicating a certain amount of sporadicmovement in all directions. Humphrey and Cope summarized the recapture of 845 little brown bats away from the banding site. They found that the net distance of autumn migration ranged from 10 to 455 kilometers and averaged 100 kilometers. Few data are available regarding the speed of migration, but Humphrey and Cope (1964) provided some information. Three bats from Thorntown (Boone County) were captured in Big Wyandotte Cave (Crawford County) 20, 32, and 36 days after being banded; the distance traveled was 132 miles. Two others moved 88 miles to Donnehue's Cave (Lawrence County) in 12 and 13 days. All these periods are maxima, so that the bats may actually have covered these distances much faster. Later, Humphrey and Cope reported movements of 60 kilometers in one night. Fall migration from the breeding colonies appears to take place from late July to midOctober. At a Boone County colony, 450 bats were present 14 August, 32 on 29 September, and 3 on 10 October (Humphrey and Cope, 1964). Spring migration data are less abundant, but it seems that bats leave the hibernating sites in the caves the last half of April and may be migrating throughout most of May, or even during the first week of June. Some intensive studies of the spring buildup of maternity colony populations would shed more light on spring movements. In late summer and early fall, relatively large numbers of little brown bats, and other bats, gather about certain cave entrances between dark and midnight. (There is similar spring behavior involving smaller numbers of bats.) Such gatherings of bats can be netted at the cave entrances for banding programs, and offer to ihe investigator the opportunity to handle fairly large samples of several species. Most of the bats that make up such gatherings are transients, evidently on migration, and on days following this "swarming" activity at the cave entrances only a few bats will be found inside the caves. Swarming is known to occur at Big Wyandotte, Donnehue's, and Ray's caves in Indiana; it probably also takes place at other, as yet undiscovered, sites. A good example of the pattern of swarming in Myotis lucifugus was obtained by Wilson Baker at Big Wyandotte Cave. On the nights of 3, 4, 5, 6, and 7 September 1962, he captured, respectively, 63, 231, 229, 306, and 117 little brown bats. Of these, 627 were males and 319 were females. Three isolated nights of netting at Ray's Cave resulted in the following catches of Myotis lucifugus: 301 on 2 September 1962; 246 on 3 September 1964; 205 on 8 September 1971. Smaller numbers were taken at Donnehue's Cave, the largest catches being 61 on 26 August 1970 and 41 on 16 September 1970. Other species of bats netted with Myotis lucifugus at these sites included M. sodalis, M. keenii, M. austroriparius, M. grisescens, Eptesicus fuscus, Pipistrellus subflavus, and Lasiurus borealis. The Indiana myotis was most abundant at Big Wyandotte and at Ray's Cave; Keen's myotis was most numerous at Ray's Cave. HOMING. Cope et al. (1961) carried out homing experiments with a summer colony of little brown myotis at Brookville (Franklin County) using radioactive bands. Forty females were banded and released 20 air miles due west of Brookville between 9:00 and 9:34 P.M. From 75 to 80 percent of these bats returned to the original roost within 4 hours 55 minutes and 6 hours. Unfortunately, the ages of the bats used in the experiment were not stated; one might expect adults to home more successfully and at a greater speed than immatures. Four bats that did not return to the original roost were located in another roost just across the street, within 7 hours 45 minutes. Cope et al. (1958) reported on homing of little brown myotis at a nursery colony at Tunnelton (Lawrence County). Groups of a total of 364 bats (67 subadults of both sexes and 297 adult males and females) were released at air mile distances of 120 miles (north-northwest), 93 miles (north-northwest), 66 miles (north-northwest), and 4 miles (west) from Tunnelton. The colony was next visited 34 days later. From 120 miles, 20 percent of the females and 25 percent of the males were back; the release from 120 miles was composed of 110 adult females and 8 adult males. The highest proportion (31 percent) of bats returning had homed from 66 140 Order / Chiroptera • Family / Vespertilionidae miles north-northwest; the release was made up of 51 adult females, 8 adult males. Only 1 bat homed from 4 miles west, where the release consisted of 32 subadult females and 22 subadult males. Homing from the fourth release (93 miles) involved 18 percent of 120 adult females and 13 subadult males. Although there is an imperfect correlation between the number of adults in the samples, the mileage traveled, and success of homing, it may be significant that adults homed more successfully than subadults. In another homing experiment at this same colony, none of 15 subadults released 82 miles south or 40 subadults released 107 miles northeast had homed three, twelve, or thirteen months later. LONGEVITY. From banding between 1952 and 1959, and recaptures analyzed in 1974, there were 100 little brown myotis that had attained a minimum age of 5 years. Of these animals 69 were females and 31 were males; it should be mentioned however, that many more females than males were banded. The complete breakdown of these recaptures (in minimum years of age) is as follows: 4 males and 8 females at least 5 years of age; 4 males and 20 females recaptured 6 years after banding; 10 males and 11 females at least 7 years old; 3 males and 16 females at least 8 years old; 7 males and 12 females recaptured at 9 years; 1 male and 2 females at least 10 years of age; 1 male recaptured 11 years after banding; 1 male at least 12 years of age. It is quite probable that some bats banded during the latter part of this longevity study are still alive; thus the percentage of recaptures will ultimately increase and additional data will be forthcoming. Also, we are reporting minimum ages; many bats could have been several to many years old when banded. There is a suggestion that males may live longer than females, as Hitchcock (1965) has pointed out for this species in Canada. HUMAN USES OF BATS. Lyon (1931) uncovered the interesting possibility that people in southern Indiana used bat oil for rheumatism; he thought that Myotis lucifugus was probably the species involved. Hibernating bats do attain great deposits of subcutaneous fat in early fall and this fat has a low melting point. Over the past twenty years, many thousands of bats have been removed from Indiana caves for physiological research. This has resulted in conflicts between bat banders and physiologists, and has, more importantly, been an additional decimating factor on hibernating bat populations. Bats have been much used as research animals by various scientists conducting life history, food habits, parasite, homing, longevity, and other studies. Bats from Indiana were also used in various radiation experiments. (It was at one time thought that bats might be immune to radiation!) Bat guano has been used in other parts of the country (and the world) for fertilizer. We have found no indication that Indiana residents have made use of this commodity. Mumford once removed 49 gallons of bat guano from the attic of a house where a maternity colony of little brown bats was present; this represented only about onethird of the droppings present in the attic. In most Indiana caves, large deposits of bat guano are not present. Many caves housing large numbers of bats have streams of water flowing through them; these streams apparently carry the droppings away and they do not accumulate. Also, much more guano is deposited by summer colonies of bats than by wintering colonies, and no large summer colonies are known in Indiana caves. MOLT. We have little data regarding molt in little brown myotis. Three adult females collected on 10 July were molting heavily (two were undergoing a complete body molt and one of these two was still lactating). An immature female taken with the adults was also undergoing a complete body molt. Two of four males (age unknown) taken 30 June had complete body molts in progress. Another unaged male taken 22 June was molting heavily. An immature male and 7 males and 3 females of undetermined age taken 30 August showed no evidence of molt. FAT STORAGE. Adults evidently build up fat deposits earlier in the fall than do young of the year. An adult female taken 7 August 1957 was so fat she appeared to be pregnant when captured and examined. Two immature females taken with her had no body fat, and an adult male had moderate fat; an unaged male taken the same day had no fat. An immature male taken 20 August had no fat; 3 Myotis lucifugus • Little Brown Myotis 141 unaged males the same day had moderate fat, and 2 unaged females had heavy fat. Nine unaged males and 2 unaged females taken 31 August had no fat. Three females and 1 male taken 3 to 26 September were very fat; another male was medium fat on 10 September. More data are needed to determine the timing of the deposition of body fat during the prehibernating period. The sample collected 31 August may have been composed of migrants, for it was netted at the mouth of a cave. Perhaps migrants have not yet accumulated their winter fat when they pass through Indiana. On the other hand, nonmigrants that may move short distances to winter quarters may take on large amounts of fat rather early in the fall. Migration would deplete stored fat, and excess fat might be extra weight for bats to transport on migration. Adult females may accumulate body fat earlier than any other sex and age class. Males may be slower (and more active) because copulation occurs in the fall (and winter). Immatures probably gain fat reservoirs more slowly because of the energy expended in growing and perhaps there is thus less physiological potential to add fat rapidly during tlie first fall of their lives. FOOD. Sixteen stomachs of little brown bats from Indiana were examined by Whitaker (1972b) for food analyses (Table 47). The food of this species from May through September was more similar to that of the red bat than to that of the big brown bat, with Lepidoptera and Trichoptera the foods most often eaten. Cicadellidae and Delphacidae (Homoptera) were also important, but Coleoptera formed only 12.4 percent of the total volume. A single coleopterous larva formed the entire contents of one stomach, and it and crickets were the only nonflying items eaten. The Lepidoptera eaten were small forms as compared to those eaten by the red bat. REPRODUCTION. The events leading up to copulation are unknown for most species of bats; further study may reveal whether or not there is courtship or other preeopulatory Table 47 Foods eaten by Myotis lucifugus (n = 16) from Indiana (from Whitaker, 1972b) Percent Percent Food Item Volume F requeney Lepidoptera (moths) 21.6 31.3 Trichoptera (caddis flies) 13.1 25.0 Diptera (flies) 11.9 31.3 Cicadellidae (leafhoppers) 11.6 43.8 Delphacidae (planthoppers) 8.8 6.3 25.0 Coleopterous (beetle) larvae 8.8 6.3 6.3 Ichneumonidae (Ichneumon flies) 3.8 12.5 Carabidae (Ground beetles) 3.4 18.8 Reduviidae (Assassin bugs) 2.8 12.5 Scarabaeidae (June beetles) 2.5 6.3 Coleoptera (Unidentified beetles) 2.2 18.8 Tipulidae (Craneflies) 1.9 12.5 Hemerobiidae (Hemerobiids) 1.9 6.3 Chironomidae (Midges) 1.6 12.5 Cerambycidae (Long-homed beetles) 1.6 6.3 Formicidae (Ants) 1.3 12.5 Chrysomelidae (Leaf beetles) 0.9 6.3 Chrysomelidae, Diabrotica 6.3 undecimpunctata 0.9 6.3 Nitidulidae (Nitidulid beetles) 0.9 6.3 Miridae (Leaf bugs) 0.6 6.3 Gryllidae (Crickets) 0.3 6.3 Insects (Unidentified) 0.3 100.2 6.3 142 Order / Chlroptera • Family / Vespertilionidae behavior. Little brown bats in Indiana have been observed copulating on 20 August (one pair, in a maternity colony), on 30 August (at least ten pairs, in a maternity colony), on 16 November (a pair in a cave), and on 11 January (a pair hanging from a cave ceiling; the male of this pair hanging from the cave ceiling grasped the fur on the back of the female's head with his teeth and pulled her head back slightly while copulating with her). Sperm are stored in the uterus of the female until spring, when fertilization occurs. In April, females begin gathering in maternity (nursery) colonies, where the young are born and reared. Most nursery colonies have been located in the attics of houses, but some are also known in barns, churches, and other buildings. Some nursery sites are dark and poorly ventilated; others (in barns) are well ventilated and well lighted. Daytime temperatures in some of the attics used as nurseries are well in excess of 100 degrees Fahrenheit. This heat evidently has no adverse effect on the bats and it is possible that they choose such sites to promote rapid growth of the young. On the hottest days, the bats tend to hang more in the open places in the attic (exposed rafters, faces of chimneys); they utilize crevices more when temperatures are cooler. In colonies used by the bats for many years, we have seen "stalactites" of crystallized urine hanging from the rafters. In other colonies, deposits of bat droppings at least a foot deep have been noted. In some cases, the weight of accumulated droppings has caused the ceiling of the room below to sag or break through. As many as 1,000 or more females may occupy a single maternity colony. The 50 nursery colonies examined by Humphrey and Cope ranged in size from about 20 to about 3,000 bats (females and young), but most colonies contained from 300 to 1,200 bats. Humphrey and Cope also noted that the larger nursery colonies were nearer the caves, and in northern Indiana (where there are no hibernacula) the colonies were smaller. Maternity colonies of Myotis lucifugus usually averaged considerably larger than maternity colonies of other species of bats that breed in Indiana. Most nurseries of little brown myotis were within a few hundred meters of a pond or stream, to which the bats flew when leaving the colony at dusk. In the maternity colonies, bats may hang from the rafters, on the sides of brick chimneys, inside the walls or between the roofing and the attic ceiling. It is not usually easy to catch or observe all individuals in a colony at any one time. Young bats are frequently found dead on the attic floor; some may become separated from adult females and succumb without being retrieved, and some probably are born dead or die and fall to the floor. From time to time a dead bat is found hanging from a wire or cord in which it has become entangled in flight. Maternity colonies usually contain a single species of bat, but we have found big brown bats sharing a building with little brown myotis on seven occasions, and we found a single southeastern myotis (a transient?) in a little brown myotis colony on 1 October. Once selected, breeding sites are evidently used for years; one such colony in Jackson County was in continuous use for at least sixty-five years, and Cope et al. (1961) suggests a colony in use for a hundred years. Small numbers of adult males may inhabit the maternity colonies. One colony contained 2 percent males on 20 April, and another contained 3 percent males on 30 April. At a third colony, 7 males and 121 females were present on 29 May. As the season progresses, a higher percentage of males sometimes occupies the colony. One colony contained 32 percent males in late August, 84 percent the first of October, and 96 percent the second week of October (Cope et al., 1958). Whether the colonies are staging areas for males in preparation for fall migration, or whether the gathering of males in the nurseries facilitates mating is unknown, but copulation has been observed there as early as August. More intensive studies are needed to determine what percentage of these males are animals reared there during the summer and how many are transients. Many animals from a maternity colony tend to return each year to that colony, but banding studies have also revealed that individuals may occupy different maternity colonies in different years. In general, it is felt that many adult females return to the same maternity colonies year after year throughout their lives. Some males also may be found in a particular maternity colony more than one year. One male was banded at Tunnelton (Lawrence County) in a breeding colony on Myotis lucifugus • Little Brown Myotis 143 30 August 1954 and released at Terre Haute (Vigo County) that night. He was again in the Tunnelton colony on 4 July 1957 and on 25 July 1962. He was also captured twice during the fall at Big Wyandotte Cave (Crawford County), on 7 September 1961 and 13 September 1963. Another male banded 7 August 1957 at Tunnelton was not caught again there but was found 9 August 1958 and 14 August 1963 in a maternity colony at Thorntown (Boone County), 95 miles north-northwest. A male banded 18 June 1953 in a nursery colony near Cortland (Jackson County) and released at Freetown (8 miles west) was back at Cortland 2 July 1953. On 14 June 1954, he was taken again from Cortland and released near Kossuth (Washington County), and was found at Cortland on 31 August 1954; this time he was released at Bedford (Lawrence County), but was found again at Cortland on 18 August 1961. The return of adult female little brown myotis to a particular maternity colony has been determined from banding studies. From 18 June 1953 to 17 August 1962, there have been ten visits made to a maternity colony near Cortland. One female M. lucifugus has been present in this colony over a span of seven years, six females over a span of eight years, and nine females over a span of nine years. Since the colony has not been visited since 1962, we do not know whether some females actually used the colony for longer periods. Two males were found here over periods of seven and eight years, but seven males are known to have been present in a single year and three for two years. Only four little brown bats banded as young of the year in this colony have been recaptured there in later years; these were found one, two, two, and nine years later. Cope and Hendricks (1970) reported on the fate of twelve "well-established little brown bat nursery colonies in Indiana; some of these had been active for at least ten years. Of these colonies, checked from June to August 1969, three were no longer present, four showed declines of 48 to 96 percent, and five retained their former levels. A single young is generally produced each year by the adult female; occasionally twins may be born. Parturition usually occurs in late May, in June, or in early July. The young are capable of flight about four weeks after birth. One newborn little brown myotis weighed 1.4 grams, had a total length of 48 mm, a forearm length of 12.5 mm, and a wingspan of 94 mm. The young in a nursery colony are not all born at the same time; twelve young from Fountain County (21 June) ranged from newborn to well-furred individuals and seemed to represent three rather distinct developmental stages on the basis of their morphological characteristics. Ten of them had forearm lengths from 12 to 30.5 mm (average 21 mm). Two adult females collected from this colony on the same date each contained a single fetus, measuring 20 and 21 mm respectively in crown-rump length. At a maternity colony in Pulaski County (2 July) sixteen young had forearm lengths of 15 to 34 mm (average 26 mm). On 21 June, eleven young from a colony in Clay County had forearms ranging from 13.5 to 21 Table 48 Ectoparasites of Myotis lucifugus (n = 84) from Indiana Parasites Parasites Total Average Hosts Parasitized Total Percent Fleas (Siphonaptera) Myodopsylla insignis Mites (Acarina) other than chiggers Macronyssus crosbyi Acanthophthirius lucifugus Chigger Mites (Trombiculidae) Euschoengastia pipistrelli Ticks (Ixodides) Ornithodoros kelleyi 109 7 0.01 0.10 0.01 1.30 0.08 5 1 20 2 1.2 6.0 1.2 23.8 2.4 144 Order / Chiroptera • Family / Vespertilionidae mm (average 18.2 mm). And nine young from Kosciusko County (1 July) had forearms from 12 to 28 mm (average 17.9 mm). Growth of young bats is relatively rapid; some reach adult proportions and weights and are capable of flight by early July. For example, one immature male weighed 6.4 grams and had a 36-mm forearm by 1 July. PARASITES AND DISEASES. Only 1 little brown myotis of 80 examined for rabies tested positive (Whitaker and Miller, 1974). We found external parasites on 25 of 84 little brown bats examined, and internal parasites in 14 of 92 specimens. Two mites, a chigger, a tick, and a flea were the external parasites (Table 48). Internal parasites found were nematodes and trematodes (as yet unidentified). Wilson (1961) reported additional data on parasites, including a flea, Nycteriodopsylla chapini, not found in our sample. The little brown bat harbors relatively few external parasites. Of the six species found on this bat to date, only the chigger is very common. The flea Nycteriodopsylla chapini is normally found on the big brown bat; Wilson's (1961) specimen from the little brown bat probably represents a case of straggling. Myodopsylla insignis is often found on Myotis lucifugus in summer colonies, and Wilson collected 207 fleas of this species in 22 Indiana counties. Ornithodorus kelleyi is primarily found on the big brown bat, but Wilson reported 49 ticks of this species on little brown bats from 7 counties. Whitaker (1973) recovered 8 specimens of the mite Macronyssus crosbyi, 1 of Acanthophthirius sp., 109 of the chigger Euschoengastia pipistrelli, 7 ticks, Ornithodoros kelleyi, and 1 flea, Myodopsylla insignis, from 84 little brown bats. The Acanthophthirius has since been described as new, A. lucifugus Fain and Whitaker, 1976. DECIMATING FACTORS. Little is known about the natural predators of Myotis lucifugus but some bats of this species are undoubtedly taken by hawks, owls, and snakes. Martin (1961) reported on the probable predation on bats in Ray's Cave by the prairie vole, but the bat remains noted were not identified to species. One day when Mumford was releasing banded little brown myotis outside a cave, a blue jay actively pursued one of the released bats for some distance but evidently was unable to catch it in flight. Some years ago, the floor of a low passage in Big Wyandotte Cave was littered with bat wings, said to have been the remains of bats (a species of Myotis) captured and eaten by a domestic cat. The worst known enemy of bats is man, who uses various methods in attempting to drive nursery colonies from buildings. We know of the use of tear gas, sulphur candles, moth balls, automobile exhaust fumes, and heavy grease. DDT and cyanide gas have been used in some roosts to eliminate the bats. The most successful way to prevent bats from entering attics is to close all entrance holes that the animals can use; this is sometimes difficult, if not impossible, but it is the most satisfactory and permanent method. The remodeling of houses, churches, and other buildings has eliminated many summer colonies. Vandalism has resulted in the loss of bats at various seasons. Fire, sticks, stones, and BB guns have all been used to kill and mutilate bats in their roosts. During hibernation, the mere disturbance of bats can be harmful to them; if they are awakened and fly about, they use up some of their stored fat reserves. Repeated disturbances may cause them to deplete so much fat that they cannot live through the winter. Researchers have probably caused considerable damage to bats. We know, for example, that large numbers of bats have been removed from some Indiana caves and other roosts for physiological studies. Others have been collected for scientific specimens or to be examined for food habits, parasites, or other data. Moderate collecting of specimens by scientists who know the species and keep abreast of the status of various bats in Indiana does relatively little harm. But in 1949, a cluster of several dozen of the relatively rare southeastern myotis was removed from a cave for specimens. This cave has housed only a few of that species each winter since then. Now that the Indiana myotis is on the national endangered species list and is rather common as a hibernator in some Indiana caves, it becomes more important that those people working with bats are able to identify the various species readily. The unjustified removal of excessive numbers of this species Myotis austroriparius • Southeastern Myotis 145 (either accidentally or intentionally) could deal a damaging blow to its future in Indiana. We also feel that bat banding could have been a factor in the decline of some bat populations in the state. Banding was begun in Indiana, at Big Wyandotte Cave, in 1946. Numerous workers have banded bats since 1946, and the disturbance caused by catching, confining, banding, and otherwise causing the animals to waken during hibernation may have produced casualties, although we have no proof of this. Also, it is well known that bands applied to the wings of bats can cause irritation, inflammation, the formation of scar tissue, and possibly infection. We cannot evaluate the effect of banding on the well-being of bats, but it is possible that deaths have occurred as a result of banding activities. In more recent years, we have been concerned about man's increased use of insecticides and the possible detrimental effect on bats. All bats that occur in Indiana feed exclusively on insects and thus are subject to high levels of insecticides that may be present in their food. We do know that the decline in the numbers of some bats in the state has been dramatic since 1951, but we still cannot pinpoint the reasons for these decreased populations. TAXONOMY. The subspecies in Indiana is Myotis lucifugus lucifugus (LeConte). SELECTED REFERENCES. Anthony and Kunz, 1977; Barbour and Davis, 1969; Belwood and Fenton, 1976; Benton and Scharoun, 1958; Cagle and Cockrum, 1943; Cope, Baker, and Confer, 1961; Cope and Hendricks, 1970; Cope, Koontz, and Churchwell, 1961; Cope, Mumford, and Wilson, 1958; Davis and Hitchcock, 1965; Fenton, 1970, 1977; Humphrey, 1964; Humphrey and Cope, 1964, 1976; Kunz, Anthony, and Rumage, 1977; Walley, 1970. Myotis subulatus: Hahn, 1908b (in part) Myotis austroriparius: Miller and Allen, 1928 DESCRIPTION. Most Indiana specimens of the southeastern myotis have a gray dorsum and a whitish venter. These animals closely resemble gray-pelaged gray myotis in color, but the gray myotis has a longer forearm, dorsal fur the same color from base to tip, and less white (more grayish) on the venter. The hairs on the dorsum of the southeastern myotis are more woolly and have dark bases and paler tips. Brown-pelaged individuals are best separated from other Myotis in Indiana by forearm length and (in life) by pink noses. Color in Indiana Myotis austroriparius appears to be best correlated with season; animals tend to be grayish in winter and spring and brownish in summer. Most of our brownish-pelaged specimens were collected in August. For example, seven captured on 4 August were quite variable in color; threewere brown above and tan below, and four were grayish above (some with brownish Southeastern myotis. Photo by Bruce J. Hayward patches). All were in heavy molt. Four older specimens, taken by Hahn from 2 August to 9 August 1907, show the same variations. In Southeastern Myotis Myotis austroriparius (Rhoads) Rhoads' Bat 146 Order / Chlroptera • Family / Vespertilionidae addition, three taken 20 August 1969 were all molting; two were undergoing nearly complete molts. We suspect that the brownish color is related to the reproductive cycle, as it seems to be in the gray myotis we have examined from Kentucky, with females becoming quite brownish during the summer. Most of our brownish southeastern myotis taken in August are females. Weights and measurements are shown in Table 49. The dental formula is I — C — P — M — = 38. 3 13 3 STATUS AND DISTRIBUTION. The first Indiana specimens of Myotis austroriparius were taken by Walter Hahn near Mitchell (in what is now Spring Mill State Park) in 1906. We have examined 13 extant specimens taken by Hahn, who had identified 12 as Myotis subulatus (= Myotis keenii) and 1 as Myotis lucifugus. Miller and Allen (1928) first realized the true identity of 4 of Hahn's specimens after they almost described them as a new species. Lyon (1936) reported no new records. The southeastern myotis is rare and evidently occurs most commonly in southcentral Indiana, where it is known from Crawford, Greene, Lawrence, and Washington counties (Map 15); for all but Lawrence County, these are single records. There has been a decided decrease in Indiana wintering populations since 1949. The species is present throughout the year, but there is no record of breeding in the state. HABITAT. Except for a female found in a breeding colony of little brown bats in a building, all Indiana records of the southeastern myotis are from caves. ASSOCIATED SPECIES. The only other bat we have found clustered with Myotis austroriparius was M. lucifugus. On 28 December 1955 there were 27 southeastern myotis and 2 little brown myotis in a compact group in Donnehue's Cave; not far away a single southeastern myotis was in a clump of 10 little brown myotis. In this same cave, on 30 December 1959, a southeastern myotis was clustered with 6 little brown myotis. One southeastern and 2 little brown myotis occupied a small hole in the ceiling of DonTable 49 Weights and measurements of Myotis austroriparius from Indiana Males Females Total length (mm) n 43 13 X 83.3 87.2 range 70-91 80-95 SD 5.5 4.6 SE 0.8 1.3 Tail length (mm) n 42 13 X 34.4 35.8 range 27-40 30-41 SD 4.1 3.0 SE 0.6 0.8 Hind foot (mm) n 41 13 X 8.8 9.6 range 7-10 9-11 SD 1.1 0.8 SE 0.2 0.2 Tragus (mm) n 32 7 X 7.0 7.0 range 5.5-9.0 5.2-8.0 SD 0.8 0.9 SE 0.1 0.3 Forearm (mm) n 35 10 X 38.1 38.1 range 32-41 35.5-40.0 SD 1.9 1.5 SE 0.3 0.5 Weight (grams) n 27 16 X 6.5 7.2 range 4.1-9.2 5.1-9.1 SD 1.1 1.3 SE 0.2 0.3 aldson's Cave, 30 December 1959. During the year, Indiana myotis, gray myotis. Keen's myotis, big brown bats, eastern pipistrelles, and silver-haired bats have been found in caves with southeastern myotis. The caves that harbor the largest numbers of southeastern myotis contain practically no Indiana myotis, perhaps suggesting a subtle habitat preference between the species. POPULATIONS. We have no record of southeastern myotis for Indiana between 17 Myotis austroriparius Southeastern Myotis 147 Map 15. The Southeastern Myotis, Myotis austroriparius, in Indiana December 1907 (Hahn specimen) and 8 November 1947 (R. G. Prasil specimen). Before 1953, all records were from the caves at Spring Mill State Park, mostly from Bronson's Cave or Donaldson's Cave. A few southeastern myotis can still be found in the caves at Spring Mill. We have not adequately checked the several "wet" caves there often enough to determine the status of this bat in the park. There were about 50 individuals observed in Bronson's Cave on 7 February 1949; the following day, 40 of these were collected. The largest number recorded since in Bronson's Cave was 8 on 30 March 1966. A cluster of about 25 was located in Donaldson's Cave on 23 November 1951 and about two-thirds of them were caught and banded; 6 were observed in the same spot the next day. Since then, the maximum number seen in this cave was 3, on 6 March 1954. On 8 January 1954, at least 9 southeastern myotis were found in Donnehue's Cave at Bedford. This cave has been visited by us more than a hundred times over the past twenty years. Small numbers of the southeastern myotis still use the cave and the species has been found there every month of the year except July. The maximum number found on a visit was 28, on 28 December 1955; the second largest number was 19, on 12 February 1955. This bat has been reported once in each of four other caves: Ray's Cave (Greene County), a female, 7 December 1953; Salt Petre Cave (Crawford County), about 15 on 23 February 1953; Endless Cave (Washington County), 1 male, on 28 January 1961; Big Wyandotte Cave (Crawford County), 2 females, 12 February 1966. HABITS. During the hibernation period, Myotis austroriparius has been found in compact clusters of up to 50 individuals, hanging from the ceilings or on the side walls of caves. Most clusters were on side walls, but the largest was on a ceiling. Singles and small groups are usually found, especially today, since the population decline. Solitary individuals, usually males, often wedge themselves into tiny holes in the cave ceiling. Some of these holes are just large enough for a single bat. We collected at least three southeastern myotis from one such hole, and observed others; more than once we have also found a single southeastern myotis in another tiny, favored roosting hole. Holes and crevices too large for a single bat to fill have been found occupied by a southeastern myotis and 1 to 4 little brown myotis, or by up to 3 southeastern myotis. On one occasion, we found a single southeastern myotis roosting inside a hollow stalactite. Of the four largest clusters of southeastern myotis found, two were in dry portions of the cave where no water was present below them, one was in a completely dry cave (Salt Petre Cave), and one was over a gravel bar at the edge of the cave stream. Singles and small groups have been observed both in dry passages and hanging directly over water. During the winter these bats are readily awakened and are more active than are other Myotis. On 23 November 1951, we found an active cluster of southeastern myotis because of their squeaking and probably would have missed them otherwise. Individuals dis- 148 Order / Chiroptera • Family / Vespertilionidae lodged from die ceiling or a crevice may escape by taking flight. Our attempts to catch two on 13 April were unsuccessful. One of the bats, in a crevice in the ceiling, immediately squeezed itself into a tiny hole when it was touched with a stick. Repeated careful prodding caused it to drop from the crevice and fly away. The second was inside the hollow core of a broken stalactite, but also dropped out and flew when touched with a stick. On 11 January 1949, a cluster of about 35 to 40 southeastern myotis was observed on the ceiling of a cave. One, dislodged from die edge of the group witii a stick, fell to the floor, but the rest immediately flew farther back into the cave. On the night of 23 November 1951, 16 southeastern myotis were banded in Donaldson's Cave. About 15 minutes later one of these was recaptured in Bronson's Cave, which is one-half mile from Donaldson's Cave, with which it has, however, an underground connection. One or two individuals have quickly taken flight when the beam of a flashlight was shined on them. These experiences and others suggest that Myotis austroriparius does not tolerate disturbances and is likely to change its roosting site quite readily, even in hibernation quarters. On 23 February 1953, there were clusters of 7 and 8 on die side wall of Salt Petre Cave; a single bat was carefully taken from one cluster for a specimen, but none of the bats flew. On 7 March 1953, not a bat of this species could be found in the cave (which is easy to census for bats), nor have we seen M. austroriparius there again. After we captured and banded 16 of a cluster of about 25 one night, we returned the next day and found only 6 hanging at the same spot. However, in two instances we have found groups in two successive winters hanging at identical spots; such roosting spots may be particularly favorable to the species for hibernation. Seasonal Activity. We obtained some information on this aspect of behavior from our work at Donnehue's Cave. Periodic netting of the entrance to the cave throughout a year resulted in our capturing a single southeastern myotis on 18 November and one on 4 February. These captures indicate that diis bat may move about during the winter, especially when the temperature is above freezing. That the bats mentioned above left Salt Petre Cave between 23 February and 7 March also points up this possibility. Daily Activity. Myotis austroriparius emerges from its daytime roosts in caves relatively late in the evening. Our Donnehue s Cave data with regard to this behavior were interesting. We summarized the comparative times of netting the first southeastern myotis, the first little brown myotis, and the first bat on seven evenings, recording only animals that struck the inside face of the nets (and thus were presumably leaving the cave). On the six occasions when we captured both M. lucifugus and M. austroriparius, the latter was caught consistently later than the former by 24 to 186 minutes. Myotis lucifugus was not always the first species netted each evening, and the time between the netting of the first bat of any species and the first M. austroriparius ranged from 34 to 186 minutes, for seven dates. A southeastern myotis was netted as late as 12:16 A.M. (26 August) and we captured 11 others between 10:00 P.M. and 12:05 A.M. (26 August to 14 October). One was captured on 4 February at 7:17 P.M., one on 11 May at 9:44 P.M., two on 30 October at 8:45 P.M. and 9:10 P.M., 1 on 18 November at 8:42 P.M. and 2 on 20 August at 9:12 and 9:37 P.M. MIGRATION. Although numerous southeastern myotis have been banded in Indiana, band recoveries have shed no light on movements of the species. We have no proof that migration occurs; no individuals have been collected or observed north of the known wintering distribution in the state. The bats obviously move into and out of the hibernating caverns periodically and the female found in a little brown bat nursery colony probably did not hibernate there. Perhaps this species is more sedentary than other Myotis. LONGEVITY. The relatively small number of southeastern myotis recaptured after banding revealed little regarding the life-span of the species. Of 21 animals recaptured, 16 were at least 1.5 years old, 2 were at least 2.5 years of age, 1 was at least 3, and 2 were at least 4 years old. SEX RATIOS. Of 62 southeastern myotis collected in caves during nine months of the year, 48 were males and 14 were females (77.4 percent males); no specimens were Myotis grisescens • Gray Myotis 149 taken in May, September, or October. Of 25 collected in February, only 5 were females (20 percent), but of 15 taken in August, 6 were females (40 percent). Of 104 southeastern myotis banded in caves (every month except May, June, July) 61 were males and 43 were females (58.6 percent males). For Donnehue's Cave, where 80 of the 104 were banded, 45 (56.3 percent) were males; also, 11 males and 5 females were collected there, giving a total of 58.3 percent males for this cave. FOOD. We have examined no bats of this species which had food in the stomachs. REPRODUCTION. We have no direct evidence that the southeastern myotis breeds in Indiana, although adults have been captured in each month of the year. Elsewhere, females each produce two young in May. We netted a female on 4 August that evidently had been nursing young; she had areas devoid of hair around each nipple. A group of about 25 southeastern myotis observed on 23 November was quite active and noisy; one female had moisture, which could have been semen, about the vulva. J. B. Cope saw a pair that "seemed to be copulating" on 7 March. Males with scrotal testes have been collected or examined in March, April, and August; those with the largest testes were taken in August. In this species, each testis descends to a conspicuous position on either side of the base of the tail. PARASITES. We examined 6 southeastern myotis for parasites and found the mite Olabidocarpus whitakeri, the chigger Euschongastia pipistrelli, and some unidentified intestinal trematodes (Whitaker and Mumford, 1971a; Whitaker, 1973). Olabidocarpus whitakeri was described by McDaniel and Coffman (1970) from material collected from three southeastern myotis taken from Donnehue's Cave, 20 August 1969. DECIMATING FACTORS. Man is probably the major enemy of the species. One southeastern myotis was found dead in a cave on 30 December, hanging by one foot from the ceiling. On the night of 18 November, during netting of Donnehue's Cave, a screech owl struck the mist net immediately after a southeastern myotis was caught; it appeared that the owl was pursuing the bat, or possibly attempting to take it from the net. A southeastern myotis found in a cave on 7 April had the tips of both ears somewhat shortened and devoid of pigment, a condition suggesting frostbite. TAXONOMY. Myotis austroriparius mumfordi Rice (1955) was described from Indiana specimens taken at Spring Mill State Park, but LaVal (1970) has since concluded that the species is monotypic. Thus, the current name for the species is Myotis austroriparius (Rhoads). SELECTED REFERENCES. Barbour and Davis, 1969; Foster, Humphrey, and Humphrey, 1978; LaVal, 1970; Rice, 1955, 1957; Sherman, 1930. Gray Myotis Myotis grisescens Howell Large-winged Bat, Howell's Bat Myotis velifer: Hahn, 1908b Myotis grisescens: Howell, 1909a DESCRIPTION. This relatively large, gray myotis can be separated from other species of the same genus in Indiana by its forearm length (42-45 mm), dorsal hair the same color from base to tip, and the attachment of the wing membrane to the tarsus instead of to the side of the foot. In Indiana, only Myotis austroriparius has the grayish dorsal color ofM. grisescens; we suspect that the two species exhibit roughly the same seasonal color variations. For example, an adult female M. grisescens taken on 9 August and examined by Howell was described by him as being "sepia above, with a russet tinge" and "hairbrown" below. This specimen (now lost) was evidently similar in color to about a dozen female gray myotis we have seen taken from a maternity colony in Adair County, Kentucky, on 2 June 1958. Indiana specimens 162 Order / Chiroptera • Family / Vespertilionidae females, were taken to the Joseph Moore Museum on the Earlham College campus where they were identified as Myotis sodalis and entered in the Museum collection. . . . These bats represent the first reported maternity colony ol Myotis sodalis. During the summers of 1972 and 1973, Cope and his co-workers captured 31 Indiana myotis along Nolands Fork River, but no maternity colony was located. In the summer of 1974, the investigators were successful in finding the colony (Humphrey et al., 1977): The nursery roost was located under the loose bark of a dead bitternut hickory (Carya cordiformes ...), that was alive in 1969. The bats used spaces behind several different patches ol bark, some interconnected and others evidently isolated. The two places most consistently occupied by the largest numbers of animals had ESE and SSW exposures and consequently received much solar heat on their outer surfaces. During the winter of 19741975 much of the bark used for roosting the previous summer fell off the tree, while new slabs curled away from the trunk to form new potential roosting spots. It was found that part of this population also used an alternate roost under the loose bark of a living shagbark hickory about 30 meters away. Another alternate roosting site was suspected, but not found. The resident population in 1974 consisted of 25 females and 23 young that were reared to the volant stage. In 1975, there were at least 23 young weaned by 28 females. The first bats arrived at the nursery site in 1975 on 4 May, and substantial numbers appeared beginning 18 May. Available data suggest that each female bore a single young. Parturition in 1974 took place from 25 June to 4 July. Young were volant at 25 to 37 days of age. In 1975, young reached the volant stage 16 days earlier than they did in 1974. Indiana myotis were present at the roost until 10 October 1974 and until 19 September 1975. Some observations were made on parental care, although the colony was in a difficult site to observe. One evening, about half the adults of the colony each carried their single young to new locations, from one patch of bark to another. The young were easily visible and clung to a teat while being carried. These young appeared to be from 7 to 10 days old and were about one-quarter to onethird the size of their mothers. From 21 July to 19 August 1974, when young were beginning to fly and forage, the observers noted "pairs of M. sodalis flying in tandem." Such flights were characterized by the following bat staying within 1 meter of the leader, and it was thought that such pairs were composed of an adult and young. Such pairs occasionally returned to the nursery tree and landed together. One pair flew 290 meters from the roost to a foraging site. The male embryo removed from a gravid Myotis sodalis shot on 18 June 1959 in Lagrange County measured 39 mm in total length. The teeth were not completely through the gums, but the tips were quite evident. There were few hairs present on the head, but some hairs were located posterior to the nostrils, on the chin and muzzle. No hairs were present between the nostrils. There were fringes of hair along the upper and lower lips. The chin region was heavily haired over an extensive area. One hair was present at the angle of the jaw on either side of the head. There appeared to be a chin gland present. On the dorsum, short hairs were over the rump area down to below the level of the knees and along the legs to the base of the feet. Sparse hairs were present on the toes. The only hair on the venter was as described above and was confined to die head. Five immature males taken in late July and early August had testes ranging from 1.5 by 2.5 mm to 2.5 by 4 mm. Males widi the largest testes (9 mm long) were collected in mid-August; specimens taken in September had testes that averaged smaller in size than August specimens, but our sample (19 animals) is rather small. PARASITES AND DISEASES. None of 34 Indiana myotis tested for rabies by Whitaker and Miller (1974) was infected with this disease. Whitaker (1973) examined 43 Indiana myotis for external parasites and found 15 (44.1 percent) to be infested. The mite Macronyssus crosbyi, a nonnal parasite of this species, was the only parasite regularly found (70 taken). Spinturnix globosus is a peculiar large spinturnicid mite found in the anus of M. sodalis. It was originally described from M. sodalis from Ray's Cave, in Greene County, Indiana (Rudnick, 1960). Six individuals of this species were found. Also two Lasionycteris noctivagans • Silver-haired Bat 163 myobiid mites were taken. They were listed by Whitaker (1973) as Acanthopthirius sp., but have since been described as A. lucifugus (Fain and Whitaker, 1976). Wilson (1961) reported two fleas, Myodopsylla insignis, on this species. A nonvolant young M. sodalis examined by Humphrey et al. (1977) was host to 7 bat bedbugs (Cimex sp.) and about 125 unidentified mites. DECIMATING FACTORS. DeBlase and Cope (1967) reported the discovery of a dead M. sodalis impaled on a barbed-wire fence. The remains of numerous individuals have been found in various caves, but cause of death is generally unknown for these specimens. Cope reported about 1,000 Indiana myotis destroyed by vandals in one cave. Some years ago, the floor of a low passage in Big Wyandotte Cave was littered with bat wings. The tour guide told us that a domestic cat had been capturing and eating the bats (discarding the wings). Although we could make no positive identification from only the wings, the bats were all Myotis and undoubtedly some were sodalis. Others have reported that cats have fed on M. sodalis in this cave. Humphrey et al. (1977) reported the unsuccessful attempt of a screech owl to catch a foraging M. sodalis near the nursery colony. TAXONOMY. Myotis sodalis Miller and Allen was described in 1928 from specimens taken in Big Wyandotte Cave, Crawford County, Indiana. No subspecies are recognized. SELECTED REFERENCES. Cope and Humphrey, 1977; Cope, Richter, and Mills, 1974; Easterla and Watkins, 1969; Hall, 1962; Humphrey, 1978; Humphrey and Cope, 1977; Humphrey, Richter, and Cope, 1977; Mumford and Calvert, 1960; Mumford and Cope, 1958; Richter, Seerley, Cope, and Keith, 1978. Silver-haired Bat Lasionycteris noctivagans (LeConte) Lasionycteris noctivagans: Evermann and Butler, 1894b DESCRIPTION. The silver-haired bat is a medium sized, blackish bat with whitish hair tips (frosting) both above and below. The amount of silver tipping on the dorsum varies considerably among individuals. Three Indiana specimens seen by us exhibit a "chocolate" color phase, in which the blackish color is replaced with brown. One of these specimens has about the same amount of frosting that black-pelaged animals have, but gives the overall appearance of a cocoa-colored individual (Kirkpatrick and Conaway, 1948); the membranes of this specimen are also paler than those of the usual, blackish animals. The brown phase, then, may represent some type of genetic variation (dilution) from the normal color. Lasionycteris is not likely to be confused with any other species of bat occurring in Indiana. Weights and measurements are shown in Table 56. The dental formula is 2^1 p2.f3_ Silver-haired bat. Photo by Anthony Brentlinger, 1 J C J 3 3 ISU AV Center 164 Order / Chiroptera • Family / Vespertilionidae Table 56 Weights and measurements of Lasionycteris noctivagans from Indiana Males Females Total length (mm) n 17 38 X 98.2 100.4 range 87-112 84-113 SD 7.0 7.2 SE 1.7 1.2 Tail length (mm) n 17 38 X 37.9 39.2 range 31-41 27-47 SD 4.0 4.3 SE 1.0 0.7 Hind foot (mm) n 15 34 X 7.6 8.5 range 7-10 7.5-11.0 SD 0.9 1.0 SE 0.2 0.2 Tragus (mm) n 16 26 X 5.6 6.0 range 4-7 5-8 SD 1.0 1.0 SE 0.2 0.2 Forearm (mm) n 16 29 X 41.4 41.5 range 39-44 39-43 SD 1.5 1.3 SE 0.4 0.2 Weight (grams) n 11 28 X 10.6 10.4 range 7.4-16.4 7.0-15.7 SD 2.5 2.0 SE 0.7 0.4 STATUS AND DISTRIBUTION. The silverhaired bat is evidently an uncommon spring and fall migrant and a rare winter resident; its exact status is poorly known. It evidently occurs throughout Indiana (Map 19). Spring specimens were taken from 18 April to 28 May and fall specimens from 29 August to 6 November. Hibernating individuals have been found on occasion in caves and buildings. Most of the winter records and specimens are from Tunnel Cave, Clifty Falls State Park (Jefferson County). In the collection of the Joseph Moore Museum are three specimens from Tunnel Cave, a male dated 10 December 1960, and two females dated 16 February 1964. An unsexed specimen taken from a building on the Earlham College campus, Richmond, on 16 January 1956 is also in this collection. It is possible that March and November records of the silver-haired bat from Indiana represent migrants, rather than wintering individuals. In spring and fall, one may occasionally observe four or five silver-haired bats foraging about the edges of a woodlot at dusk, but more often singles or two or three bats are present. On the evening of 23 April 1954, four silver-haired bats were shot by James B. Cope as they foraged at one site on the Clark State Forest (Clark County) near Henryville. We have practically no data concerning the numbers of this bat in Indiana during various seasons of the year. There is no indication Map 19. The Silver-haired Bat, Lasionycteris noctivagans, in Indiana Lasionycteris noctivagans • Silver-haired Bat 165 that it is at all common in winter; in fact, it is quite rare and has only been seen in three caves. We have no reports of large numbers during migration. HABITAT. Most Indiana specimens of Lasionycteris were shot flying about the borders of woodlots, over ponds, or along streams. One was shot on 3 May as it foraged about the edge of a wooded slope, bordering a clover field. Two, shot on 18 April, were feeding over a road and along a mowed power line, both through a mature, deciduous woods. Most of those taken in Tippecanoe County have been shot about the borders of two mature, deciduous woodlots surrounded by pasture or cultivated fields. One silver-haired bat was observed (date not recorded) beneath the loose bark of a tree (Porter County) by Raymond Grow; A. H. Westing and W. C. Bramble caught two beneath the loose bark of a dead tree (Perry County) on 6 November. Dan Bloodgood caught one beneath the loose bark of a shagbark hickory on 14 September. One was shot over a small pond surrounded by pasture, but near a wooded area. In Newton County, one was shot at dusk as it foraged around large, scattered oak trees on a sand ridge. Several silver-haired bats have been found in caves, and a few have been captured in or on the sides of houses. One was hanging on the side of a house on 6 September, at 11:30 A.M. At 6:45 on the morning of 3 September 1959, a silver-haired bat was noted flying about a small concrete lighthouse 100 yards from shore at the end of a breakwater extending out into Lake Michigan, at Michigan City. The bat lit on the side of the structure, then crawled into a narrow crevice between the building and the end of a steel catwalk, where it was easily captured by hand. A second Lasionycteris was then noted flying weakly about and it was easily knocked down by hand and caught. The morning was clear and cool (55 to 58°F) and there was a 7to 8-mile-an-hour wind; the bats flew mostly on the lee side of the structure and gave the appearance of being quite tired. R. D. Kirkpatriek found a female hanging 8 feet above the ground on a cliff face, just outside a cave entrance (Jennings County) on 3 January. ASSOCIATED SPECIES. While shooting bats at dusk, we have collected the following species along with Lasionycteris noctivagans: Lasiurus borealis, Nycticeius humeralis, Eptesicus fuscus, Myotis sodalis, Myotis lucifugus, and Pipistrellus subflavus. The red bat has been the most common associate at these times, but this may be because it, too, emerges early in the evening and is more easily shot than species that may emerge to forage later. Two silver-haired bats found in Donaldson's Cave on 6 March shared the room in which they roosted that day with little brown myotis, southeastern myotis, big brown bats, and eastern pipistrelles. Silverhaired bats share Tunnel Cave with little brown myotis, Keen's myotis, big brown bats, and pipistrelles. HABITS. Silver-haired bats are usually observed foraging rather early in the evening—about the same time that red bats, pipistrelles, and evening bats are flying about. At some collecting sites, the silverhaired bat was the first species noted at dusk. We should point out, however, that most of the silver-haired bats collected by shooting in Indiana have been taken where pipistrelles (a noted early flyer) are absent or very rare. Where Pipistrellus subflavus is common, as in southern Indiana, it is normally the first species to become active at dusk. Lasionycteris usually flies fairly low, hawking insects from 15 to 30 feet above the ground, along woods borders. On 15 May, however, several were feeding above the crowns of trees in a woodlot at Lafayette; these bats were feeding 50 to / 5 feet from the ground when over the canopy. They occasionally made forays away from the trees out over the adjacent pasture, where they tended to fly nearer the ground. Two silver-haired bats shot on 18 April were feeding along a mowed strip beneath a power line, parallel to an adjacent woods; the bats flew rather slowly back and forth along this open strip, feeding not more than 20 feet from the ground. Most silver-haired bats seen in Indiana are migrants, but we have no idea where these spring and fall flights originate or terminate. Only a few bats of this species have been banded in Indiana (most of these were captured in caves), and band recoveries have 166 Order / Chiroptera • Family / Vespertilionidae shed no light on migratory movements. J. B. Cope recaptured one a year after banding in a cave. It is possible that in winter this bat is more abundant in Indiana than is now known; it may hibernate in the crevices of rock outcrops, in caves, buildings, or in tree cavities. Spring migration seems to occur in a relatively short period from about mid-April to the end of May, but fall migration is more prolonged, extending at least from late August to early November. Six of eight silver-haired bats shot in April and fifteen of seventeen shot in May were females. Eight of fourteen shot from August to October were females. Adult males are evidently not common in Indiana or have habits that reduce their vulnerability to being shot. Relatively few silver-haired bats have been observed in their roosts in Indiana. On 6 March, a male and a female were extracted from a narrow fissure in Donaldson's Cave. The bats were wedged together at the back of this crevice, about 18 inches from its opening; the crack had been formed when a piece of limestone had pulled away from the cave wall at the point where the wall met the ceiling, about 10 feet from the cave floor. Both bats were very torpid. Their roosting site was at least 120 feet from the cave entrance, in a rather large, dry room. The male and female found 6 November in Perry County were beneath the bark of a 14-inch dead black oak tree; the bats were 7 feet from the ground. In the fall of 1952, Raymond Grow captured and released a silver-haired bat he found beneath the loose bark of a tree in Lake County. In September 1954, Raymond Grow saw one clinging to the outside of a screen door at the Northern Indiana Public Service Plant, Michigan City; this plant is located on the shore of Lake Michigan. The two silverhaired bats captured at the lighthouse were near this plant. When we caught these bats (3 September) we noted that they appeared weak; they were easily captured, although a few moments before they had been flying about. We believe the bats had just reached land after migrating from over Lake Michigan. Perhaps bats that fly over a portion of this lake are exhausted when they reach land and take the first opportunity to hang on some available structure, such as the screen door mentioned above. (We will have more to say on this subject under the discussion of Lasiurus borealis, another migratory species.) Most silver-haired bats collected in Indiana in the fall were quite fat. FOOD. Whitaker examined two Indiana silver-haired bats with food in their stomachs. One contained only Trichoptera and the other 90 percent Trichoptera and 10 percent Scarabaeidae. Gould (1955) reported a stable fly (Stomoxys calcitrans) in the mouth of a silver-haired bat from Massachusetts. Novakowski (1956) recorded the feeding of this bat on dipterous larvae, presumably obtained from a tree hole in which the bats were living. REPRODUCTION. Single females collected 26 April and 12 May contained no visible embryos. A female collected 9 May contained 2 embryos weighing 0.2 grams each. Another female taken 12 May had at least one 5-mm embryo; part of her reproductive tract was destroyed by shot. The label notation on a specimen shot on 14 May mentions that embryos were present, but gives no other details. One of two females taken on 20 May (Vigo County) contained two embryos measuring 2.4 mm, but the other showed no visible embryos or placental scars. We have relatively little information on testis size in Lasionycteris noctivagans. A male shot on 4 May had testes only 3 mm long and represents the only spring-taken specimen for which we have data. Two males taken on 3 September each had testes 7 mm long. Single males had testis lengths of 5, 3, 5, 5, and 3 mm on 6 September, 20 September, 7 October, 8 October, and 24 October respectively. PARASITES AND DISEASES. One batbug (Cimex adjunctus) and one myobiid mite (Acanthopthirius sp.) were found on 8 silver-haired bats examined by Whitaker (1973). Of 11 bats of this species examined for internal parasites, 4 harbored a total of 6 cestodes and 8 harbored 63 trematodes. These parasites have not been identified to species. A male silver-haired bat captured in a house in Montgomery County on 12 September 1960 tested positive for rabies. The bat had bitten a woman on the finger during the night as she was sleeping. Mumford idenPipistrellus subflavus • Eastern Pipistrelle 167 tified the bat for the Public Health Service. None of 7 Indiana silver-haired bats examined by Whitaker and Miller (1974) were rabid. DECIMATING FACTORS. We have no records of predation or of enemies of this species. TAXONOMY. No subspecies of L. noctivagans is recognized. SELECTED REFERENCES. Barbour and Davis, 1969; Miller, 1897. Eastern Pipistrelle Pipistrellus sabflavus (F. Cuvier) Georgian Bat, Northern Pipistrelle Vespertilio georgrenus: Quick and Langdon, 1882 Vesperugo carolinensis: Evermann and Butler, 1894b Pipistrellus subflavus: McAtee, 1907 DESCRIPTION. The eastern pipistrelle is the smallest species of bat found in Indiana. The upper parts range from pale yellowish brown or grayish brown to dark reddish brown. Immatures tend to be the most grayish. There is considerable variation in the pelage color of this bat. Most of the hairs on the dorsum are tricolored, having a broad, blackish basal band, then a narrower, yellowish band, and dusky tips. The venter is much the same color as the dorsum, but usually has less of a grayish cast because the belly hairs are not dark-tipped. The forearm is usually reddish and contrasts sharply with the blackish wing membranes, especially in live animals. Four specimens showing varying amounts of white in the pelage have been taken; in one, whitish blotches are present on the wing membranes, as well. Weights and measurements are shown in Table 57. The dental formula is I _~.C — P—M— = 34. 3 12 3 STATUS AND DISTRIBUTION. This little bat is a common permanent resident in southern Indiana, and an uncommon summer resident in northern Indiana (Map 20). There are few records for the northern half of the state. The northern extent of its known range is Tippecanoe County and Wells County, in western and eastern Indiana respectively. It winEastern pipistrelle. Photo by Richard LaVal ters as far north as Parke County (in Turkey Run State Park). Pipistrellus subflavus is one of the most abundant bats in summer throughout the southern one-third of Indiana. Winter populations there are quite small in comparison, indicating that most pipistrelles leave the state in the fall. The largest wintering population found in one cave was 112, in Wilson's Cave (Jefferson County). Only one breeding area has been discovered (Cope, Baker, and Confer, 1961), in Jackson County, where maternity colonies have been found for several years. Almost every cave examined during the 168 Order / Chiroptera • Family / Vespertilionidae Table 57 Weights and measurements of Pipistrellus subflavus from Indiana Males Females Total length (mm) n 28 24 X 83.5 82.5 range 74-91 76-90 SD 5.0 4.46 SE 0.94 0.91 Tail length (mm) n 28 24 X 34.3 36.4 range 31-39 31-42 SD 2.75 3.14 SE 0.52 0.64 Hind foot (mm) n 28 24 X 8.6 7.9 range 7-11 6-9 SD 1.49 0.97 SE 0.28 0.19 Forearm (mm) n 19 9 X 34.3 34.2 range 32-41 33-36 SD 2.06 1.3 SE 0.47 0.43 Tragus (mm) n 15 3 X 5.4 5.0 range 3.5-6.5 4-6 SD 0.90 1 SE 0.23 0.57 Weight (grams) n 26 23 X 6.0 5.6 range 2.8-9.2 2.6-71 SD 1.58 1.35 SE 0.31 0.28 winter has contained at least one pipistrelle; some of these caves are quite small and no other species of bat hibernates in them. The number of pipistrelles seen per visit to 72 Indiana caves ranged from 1 to 112. In 49 of these caverns, 1 to 10 were present; 11 to 24 were found in 10 caves; 25 to 50 were in 10 caves. In only 3 caves did we find more than 50 pipistrelles: in Dillon Cave (Orange County), Coon's Cave (Monroe County) and Wilson's Cave (Jefferson County). There are few data for making comparisons Map 20. The Eastern Pipistrelle, Pipistrellus subflavus, in Indiana of pipistrelle populations for different years. Walter L. Hahn observed "not more than 50" eastern pipistrelles in Coon s Cave on 29 March 1908. We have records of 31, 80, and 40 there on 23 February 1952, 13 December 1952, and 22 February 1954 respectively. Hahn saw "about 10" pipistrelles in Eller's Cave (Monroe County) on 29 March 1908. Our records show 18 there on 26 March 1952 and 25 on 3 January 1953. Banta (1907: worked in Mayfield's Cave (also Monroe County) from 1903 to 1905 and noted that P. subflavus was "fairly abundant," but gave no figures. Hahn observed 17 pipistrelles there on 11 January 1907; only 4 were found on 26 January 1952. We can conclude little concerning population trends from these comparisons until we learn more about normal, annual fluctuations in winter populations. We also know little about summer populations of Pipistrellus subflavus, and very few breeding colonies have been located. But judging from the number of pipistrelles obPipistrellus subflavus • Eastern Pipistrelle 169 served and shot throughout southern Indiana during the summer and early fall, the summering population must be many times greater than the wintering population. For example, on the evening of 8 August 1957, Mumford observed at least 75 pipistrelles along a 3-mile stretch of road in Martin County. At one spot there were 30 to 40 feeding over a small area; at another site at least 25 were foraging over an area 100 yards square. Smaller congregations of pipistrelles have been noted in several parts of southern Indiana during the summer. HABITAT. Pipistrellus subflavus appears to be most abundant in well-wooded regions which contain streams and ponds. It probably roosts by day in trees (there are no observations to confirm this), for at dusk it emerges about woodland borders. Observed day roosts are caves, mines, and (rarely) buildings. On 6 May 1961, there were 5 pipistrelles hanging beneath the roof of a small, open-sided picnic shelter. Two females were hanging from the ceiling of a room in an abandoned house on 5 May 1954. On 27 April 1952, a male was captured while hanging under a small porch of a building. All of these roosting sites were well lighted. Maternity colonies discovered in Indiana have been associated with buildings. Eight pregnant pipistrelles were taken from one maternity colony in a barn 3 miles south of Riley (Vigo County) 1 June 1964. ASSOCIATED SPECIES. In southern Indiana, red bats and pipistrelles are most frequently seen foraging together in spring, summer, and fall. Both species emerge early in the evening and perhaps our data are biased somewhat by this fact. For example, the little brown myotis has seldom been shot while feeding with pipistrelles at dusk, but Myotis lucifugus appears to emerge from its daytime roost considerably later than Pipistrellus subflavus. In fact, relatively few little brown myotis have been collected in Indiana by shooting in the evening, and observations at their maternity colonies suggest that they do not emerge to feed until deep dusk, often too late for the bat hunter to collect them. Other bats collected with eastern pipistrelles by shooting were big brown bats, evening bats, hoary bats, red bats, silver-haired bats, and Indiana myotis. During the winter, eastern pipistrelles hibernate in caves with all other species of bats that use caves at this season. It is not unusual, however, to find only P. subflavus hibernating in small caves. Pipistrelles do not hang in clusters with other species of bats during the winter. HABITS. When hibernating, Pipistrellus subflavus is normally solitary; individuals are found hanging singly throughout the caves. We have only once found even two hanging together: on 3 September 1963, two pipistrelles hanging on the wall in Ray s Cave appeared to be copulating; the bats were captured and both were males, but one had an erect penis. Most pipistrelles hang on the walls of the caves during hibernation; relatively few roost on the ceiling, but some enter dome-like depressions in the ceiling or hang inside hollow stalactites. Where there are breakdowns, consisting of large chunks of limestone which have fallen from the walls or ceilings, pipistrelles frequently hibernate on the underside of these chunks, sometimes not more than two feet from the cave floor. They also hang on the underside of ledges along the cave wall, again frequently near the cave floor or the surface of the cave stream. On rare occasions, a single bat of this species may be found in a small crevice in the cave ceiling. Once the wintering site is selected, an individual bat may hang there for long periods. Two pipistrelles occupied the same spots from at least 1 March to 5 May in a small cave in Washington County. Hahn (1908) found that the eastern pipistrelle moved about inside the cave during the winter less than other species he studied in what is now Spring Mill State Park. He reported that an individual remained in the same spot an average of two weeks, and one hung at the identical spot for 44 days. We have the general impression that others may actually use the roosting site for even longer periods, when not unduly disturbed. In fact, Banta (1907) recorded one that hung at the same spot from 20 November to 5 April; he implied that others spent long periods at other sites. The fur of many hibernating eastern pipistrelles is covered with fine droplets of condensed moisture, possibly indicating that these bats have spent long inactive periods at one spot. We have little information on the buildup 170 Order / Chiroptera • Family / Vespertilionidae of hibernating populations in the fall. A few pipistrelles (principally males) may be found in caves throughout the summer. There are evidently some interesting movements involving numbers of pipistrelles into certain caves at times. For example, there were no more than 8 pipistrelles in Ray's Cave at 5:00 P.M. on 2 September 1963; the next morning, 63 (including both sexes) were found in the cave. The new arrivals had obviously entered the cave during the night. We were unable to study these animals to determine how many, if any, remained in the cave during the winter. But our periodic censuses of wintering pipistrelles in Ray's Cave have revealed that more than half a dozen present on a single winter visit is unusual; the more than 25 seen on 7 February 1953 is an exception. Banta's work in Mayfield's Cave also furnished some data on the pattern of seasonal activity of pipistrelles. He found one in the cave on 24 September; the number of pipistrelles had increased considerably by 30 October. By 20 November, he felt all had entered the cave that were to hibernate there. There was very little movement noted after 1 December; the bats became more easily aroused in late winter and early spring. During April there appeared to be some fluctuation in numbers present on Banta's weekly visits. Pipistrelles were "very seldom" seen in the cave later in the spring and during the summer. We have noticed that at least from May to July pipistrelles hang near the cave entrance; in winter, they are normally found farther inside the cave. We have records for Pipistrellus subflavus in Indiana caves for each month of the year, but relatively few are present from mid-May to mid-August. From our meager information, we cannot determine whether most of the pipistrelles found in caves in summer are males; both sexes have been taken in June and July. Further research may shed some light on this problem. That females are found in caves during the season of the year when young are being reared suggests that all females do not bear young each year; perhaps some are not bred or some immatures do not breed. We have no evidence that this species forms maternity colonies in caves. We know practically nothing about the daytime roosting habits of P. subflacu.s during the nonhibernating season. At dusk this bat appears about the borders of wooded areas, along streams lined with trees, and about ponds or lakes near forested areas. We assume that the normal daytime roost is among the foliage of or in hollow trees, but we have never actually seen one in such a site. The pipistrelle emerges early in the evening, and the earliest individuals to appear often forage quite high, about the treetops or above them. As dusk deepens, the bats usually feed closer to the ground, coming down to hawk insects and to drink. On 3 August 1948, about 20 pipistrelles were feeding over and along Sugar Creek, in Turkey Run State Park. The first bat appeared at 7:05 P.M., then the group centered its activity over a gravel bar and swift riffle in the stream; perhaps some type of insect was emerging from the water or most of this activity was concerned with drinking. The bats fed actively for about 30 minutes then dispersed. The following evening, at the same site, a pipistrelle appeared at 6:57, it fed, drank, and flew about for at least 10 minutes before other pipistrelles were seen. During this period, it drank only in the quiet water of an eddy beneath an overhanging rock or just downstream from a large rock in the water. Its foraging area was 30 to 40 yards long and 10 yards wide. Within the next 45 minutes, 25 to 30 pipistrelles were present. As it became darker, these bats left the creek and began feeding among the treetops along it; in this case, it appeared that the early part of the evening activity was centered around drinking. On 14 August 1948, at a small farm pond about 2 miles from the Sugar Creek site, at least 20 pipistrelles were feeding and drinking at 7:15 P.M. By 7:30 only 2 or 3 were present; the bats had dispersed rather quickly, evidently after drinking. On 27 July 1961, along Coal Creek (Fountain County), the first pipistrelles seen at dusk were foraging about 75 feet in the air about the crowns of tall trees bordering the stream. The same behavior was observed along a small stream in Jackson County, on 8 July 1952. On 11 August 1959, in Monroe County, Mumford was watching for bats at dusk near a small pond adjacent to a low, flat woods composed mostly of pin oak. A few red bats were foraging at 8:05; at 8:10, about 20 pipistrelles suddenly appeared in the air and fed over the woods, the pond, a soybean field. Pipistrellus subflavus • Eastern Pipistrelle 171 and a nearby weed field. There were no buildings or caves in the vicinity and the bats had evidently emerged from the trees. From 15 to 20 pipistrelles were feeding at dusk in Vigo County on 5 June 1958 over a fallow field bordering a woods. The bats foraged in small circles low over the field and along the woods border at elevations below the treetops. On 25 July 1954, in Bartholomew County, at least 15 pipistrelles and 2 red bats were feeding over a wheat stubble near the edge of a woods, and 15 red bats and 30 pipistrelles were observed along the road within a few miles. In Martin County, on 8 August 1957, approximately 40 pipistrelles were foraging over a weed field along a small creek near the edge of a forested area. Pipistrelles are among the first bats to emerge from their daytime roosts at dusk. In southern Indiana, they can frequently be seen feeding with red bats and chimney swifts while there is still considerable light. As has been noted several times with red bats, a pipistrelle occasionally flies toward a swift. Whether the bat and the bird are after the same insect we do not know, but this is a possibility. Sometimes pipistrelles appear at dusk before red bats are on the wing; on other occasions, red bats are seen earlier than pipistrelles. Both of these species have also been seen at dawn, flying about after the sky is quite light. On 28 July 1954, at 5:32 A.M., at least three pipistrelles were watched with binoculars as they flew about just above the treetops. They appeared to fly into the foliage of a large oak and disappeared from sight at 5:40. The morning was clear and calm and the temperature was about 64°F. One pipistrelle was shot at 2:00 on a bright, sunny afternoon in either October or November in Posey County. In our study of the emergence of bats from Donnehue's Cave (Lawrence County) we found that pipistrelles were the first bats to leave the cave on eight of eleven nights (May to October) that we netted this species (Mumford and Whitaker, 1975). We captured 4 before sunset in a net set across the mouth of the cave. Although P. subflavus left Donnehue's Cave relatively early in the evening, the species remained active over a considerable period of time on certain nights. Some were netted five hours after sunset, and this species was probably active throughout the night on occasion (we did not tend the nets all night to determine this). Since we caught the greatest number (114 of 143) of pipistrelles during August and September, most of them were undoubtedly in migration. The activity patterns of migrants may differ from that of more sedentary animals; this requires further study. The pattern of bats, especially transients, emerging from a cave may differ from that of bats emerging from noncave roosts at other, or the same, seasons. Other bats using the cave during the period of our study were the little brown myotis, Keen's myotis, the southeastern myotis, the Indiana myotis, and the big brown bat. We have been interested in determining how long individual bats are active at night and whether they return periodically to their daytime roost after dark. Since practically all summer daytime roosts of P. subflavus are probably in trees, this type of information is lacking. Bats are efficient feeders, and they can fill their stomachs with food in a period of minutes at dusk when insects are abundant. But what does a bat do the rest of the night after it has fed at dusk? And how often does a bat feed during the night? While netting the cave entrances, we have usually banded and released each bat shortly after it was netted. A pipistrelle banded and released at 8:30 P.M. on 18 May at Big Wyandotte Cave was again netted as it entered the cave at 10:25 P.M. Some clue as to nightly activity was obtained the night of 22-23 August, when we netted Big Wyandotte Cave throughout the night. We captured pipistrelles from 7:20 P.M.. to 5:00 A.M., netting 1 to 6 (average 3.3) in each half-hour interval of the period. In attempting to capture pipistrelles by the use of fine wires stretched just above the water of pools where bats drink, Mumford noted that bats knocked into the water by striking the wires would frequently fly directly from the water's surface. Others would swim to the edge of the pool and could be easily captured by hand. Perhaps those that got their pelage excessively wet when they struck the water were then unable to fly as easily as those that did not; also, some may have been stunned by striking the wires. SEX RATIOS. We have obtained some sex ratio data from netting cave entrances and 172 Order / Chiroptera • Family / Vespertilionidae some from banding or observations in caves. During the winters of 1951-52 and 1952-53, a total of 137 eastern pipistrelles were banded in Coon's Cave (Monroe County); of these, 105 were males. Males outnumbered females in smaller samples from thirteen other caves. In Beck's Mill Cave (Washington County) females outnumbered males 9 to 2 on 22 December 1952, and in Upper Twin Cave (Lawrence County) 5 males and 7 females were present on 23 November 1951. The overall percentage of males banded in fourteen caves during the winter was 75.4. Sex ratio data from the netting of eastern pipistrelles at cave entrances during the fall swarming period are available from four caves. At Donnehue's Cave on three nights in August and one in September, we captured 98 males, 12 females, and 4 for which sex was unrecorded. On the night of 22-23 August 1974, we netted Big Wyandotte Cave from dusk to dawn, catching 50 males, 9 females, and 7 for which sex was unrecorded. A net set across the mouth of Ray's Cave from 6:30 to 9:00 P.M. on 3 September 1964 captured 30 male and 9 female pipistrelles. At Boone's Cave (Owen County), 15 males and no females were netted on 24 August 1962. Most of the relatively few pipistrelles seen in caves in June and July were males. Representative observations follow: Salt Petre Cave (Crawford County), 10 June 1954, a single male; Branson's Cave (Lawrence County), 11 July 1949, 1 male; May's Cave (Monroe County), 7 June 1954, 5 males; Buckner's Cave (Monroe County), 7 June 1954, 7 males. On three nights in June and July 1971, at Donnehue's Cave, 16 males and 2 females were netted at the cave entrance. LONGEVITY. The relatively few banding returns of hibernating pipistrelles have shed little light on longevity. Of 314 males and 104 females banded, 19 males and 8 females have been recaptured. Only 3 of these animals had been banded more than two years previously; these were recaptured after 26, 36, and 57 months. The average length of time between banding and the last recapture of these 27 pipistrelles was about 18 months. MIGRATION. We know practically nothing about the migration of P. subflavus in Indiana, where the bat lives near the northern borders of its geographic range. A male shot 26 May in Tippecanoe County, north of any known wintering sites, was undoubtedly a migrant. Circumstantial evidence for the movements of large numbers of animals is the fact that summer populations in southern Indiana are high, but hibernating numbers are quite low. HOMING. A male pipistrelle taken from Sullivan's Cave on 1 March 1955 was banded and then released the next day in Donnehue's Cave. On 1 February 1958, the bat was again captured in Sullivan's Cave. The caves are about 8 miles apart. We have carried out no homing experiments with this bat in Indiana, since so few maternity colonies have been found and they were all small. FOOD. Whitaker (1972b) examined the stomachs of 23 pipistrelles from Indiana (Table 58). Homopterans of the family Cicadellidae (leafhoppers) were the most commonlyeaten food, followed by ground beetles (Carabidae). Other foods represented at higher volumes were Diptera, Coleoptera, Lepidoptera, and delphacids (Homoptera). Ross (1967) found the stomach of one pipistrelle from Indiana to contain 80 percent Cicadellidae and 20 percent Formicidae. Since the pipistrelle is the smallest Indiana bat, most of the foods eaten were small. We were surprised that seven eastern pipistrelles netted at the entrance of Donnehue's Cave on the evening of 20 August between 8:31 and 10:00 had empty stomachs. Five of these were captured between 9:01 and 10:00 and seemingly had had ample time to feed, for it was dark shortly after 8:30 that evening. REPRODUCTION. The usual number of young at birth is probably two. We have records of twelve gravid females with twin embryos and two females each with a single embryo. In addition, each of sixteen gravid females examined (but not collected) appeared to contain two embryos (Cope and Humphrey). Walter L. Hahn (1908) took a female on 6 June that contained three small (2-mm) embryos, and Lindsay (1958) examined a female taken on 21 May that had four embryos measuring 3 mm. Perhaps some embryos are resorbed before parturition in this species. Parturition in pipistrelles in Indiana evidently occurs in late June and earlyJuly. We have no actual dates, although two females confined to cages gave birth (perhaps prematurely) to young on 21 June and 30 June. Another caged female, injured by shot Pipistrellus subflavus • Eastern Pipistrelle 173 Table 58 Foods eaten by Pipistrellus subflavus (n = 23) from Indiana (from Whitaker, 1972b) Percent Percent Food Item Volume Frequency Cicadellidae (leafhoppers) 21.7 52.2 Carabidae (ground beetles) 18.1 34.8 Diptera 10.7 26.1 Coleoptera 7.8 34.8 Lepidoptera 7.3 21.7 Cercopidae (froghoppers) 7.2 13.0 Muscoid flies 4.6 8.7 Formicidae (ants) 4.6 8.7 Reduviidae (assassin bugs) 3.7 13.0 Chrysomelidae (leaf beetles) 3.7 13.0 Chrysopidae (lacewings) 2.8 4.3 Tipulidae (craneflies) 2.2 4.3 Hemiptera (true bugs) 2.2 4.3 Ichneumonidae (ichneumon-flies) 1.1 8.7 Grass stems 0.9 4.3 Insects (unidentified) 0.7 8.7 Culicidae (mosquitos) 0.7 4.3 Chironomidae (midges) 0.2 4.3 100.2 on 7 July, gave birth to twins on 9 July. Mumford and Handley (1956) published an account of the behavior of one of these families just after birth. The combined weight of two embryos removed from a female on 22 June was 2.3 grams. Males with the largest testes have been collected from July to September. Four males taken in July had testes averaging 4.8 mm in length; 35 males taken in August had testes averaging 4.8 mm in length; 3 males collected in September had testes ranging from 3.5 to 6 mm (averaging 4 mm) in length. From our data, testis size is smallest in midwinter (one male with 1-mm testes on 24 February), then increases to a maximum in summer, with a rather abrupt decrease in fall (3 males with 2-mm testes on 1 October). We suspect that a considerable amount of copulation takes place in August. PARASITES AND DISEASES. The single internal parasite found in 55 individuals of this bat was one nematode in one bat. Only the chigger Euschoengastia pipistrelli (54 individuals) was found on 13 pipistrelles examined for external parasites. The only previous parasite records for this species are those of Wilson (1961), who found two specimens of the bat flea (Nycteridopsylla chapini), normally found on Eptesicus fuscus. Of twenty-four pipistrelles from Indiana examined for rabies, 3 (12.5 percent) were rabid (Whitaker and Miller, 1974). DECIMATING FACTORS. We occasionally find a dead pipistrelle hanging in a cave, but we have no idea of the cause of death of these individuals. Some must meet with accidents, for we have examined several with part of the wing membrane or uropatagium missing. A maternity colony in Jackson County was discouraged from using its roost of the previous year when heavy grease was smeared on the rafters where the bats hung. A pipistrelle shot while flying about on a sunny day had part of its body covered with what appeared to be red paint. No doubt a few of these bats are killed by vandalism in caves. TAXONOMY. The subspecies in Indiana is Pipistrellus subflavus subflavus (F. Cuvier). There is considerable color variation in Indiana specimens. SELECTED REFERENCES. Barbour and Davis, 1969; Cope, Baker, and Confer, 1961; Davis and Mumford, 1962; Jones and Suttkus, 1973; Lindsay, 1956b; Mumford and Cope, 1964; Walk:' an^ Jarvis> 1971. 174 Order / Chiroptera • Family / Vespertilionidae Big Brown Bat Eptesicus fuscus (Beauvois) Brown Bat, Large Brown Bat Eptesicus melanops: Rafinesque, 1820 Vespertilio fuscus: Quick and Langdon, 1882; Banta, 1907 Adelonucteris fuscus: Evermann and Butler, 1894b Eptesicus fuscus: Hahn, 1909 DESCRIPTION. Eptesicus fuscus is a rather large (the second largest species of bat in Indiana), brownish bat with blackish membranes and ears. The wings are brown. The body pelage varies in color from tan to chocolate brown and may be reddish brown in some individuals. It is difficult to describe a "typical" specimen other than to say that it is some shade of brown. We have seen several Indiana specimens with white spots and a few scattered, whitish guard hairs. The tragus is broad and blunt-tipped, and the ears are relatively short for so large a bat. The wingspan is about 310 to 355 mm, and the forearm varies from about 42 to 54 mm (average 46 mm). The tail protrudes a few mm beyond the end of the interfemoral membrane. The calcar has a distinct keel. There are prominent odoriferous glands on the upper lips. The only other brownish colored bat in Indiana with a forearm reaching 42 mm is the rare gray myotis, which has a forearm ranging from 42 to 45 mm and a relatively long, narrow, pointed tragus. Myotis grisescens is usually gray, but females may develop a brown pelage during the reproductive season. ' " \ Big brown bat. Photo by Whitaker Weights and measurements are shown in Table 59. The dental formula is I A c — P — M — = 32. 3 12 3 STATUS AND DISTRIBUTION. The big brown bat is a common resident throughout the state (Map 21), especially in spring, summer, and fall. It is probably the most or second most abundant bat in the state. In winter it is much less conspicuous and either hibernates in protected sites or possibly migrates. During the hibernation season, big brown bats are observed most often in the various limestone caves ol south-central Indiana; these bats hibernate also in heated buildings in other parts of the state, but we have relatively few winter records from the northern one-third of Indiana. There is some evidence from banding that the big brown bat is somewhat migratory. Numbers present in Indiana in the summer are far greater than the relatively few found in Indiana caves during the winter. HABITAT. Summer roost sites of Eptesicus fuscus include buildings, bridges, rocky escarpments, and caves, but the species is most common in old buildings. We have seen barns or other buildings that harbor 200 to 400 or more individuals in the summer. Trees and mines are also undoubtedly used, but definite records are lacking. We have one report of a maternity colony in a tree cavity. In the nonhibernating season, E. fuscus has been taken by shooting over ponds, around deciduous woodlots, along streams, and in residential areas. Many big brown bats winter in caves, and others winter in heated buildings; the percentage that uses buildings is unknown, but may be considerably greater than records now indicate. Little is known about the species in hibernation, but 26 big brown bats were found in apparent hibernation on March 26 in a barn north of Terre Haute. It is possible that they came there from some other area after initial arousal from hibernation, although this did not seem likely since the bats were wedged very tightly into tiny crevices, Eptesicus fuscus • Big Brown Bat 175 Table 59 Weights and measurements of Eptesicus fuscus from several counties of Indiana Vigo Jefferson Vanderburgh Northern Indiana Males Females Males Females Males Females Males Females Total length (mm) n 26 61 23 21 13 6 19 11 X 108.4 113.4 113.0 111.9 112.2 108.0 112.9 112.9 range 99-124 106-129 106-121 101-126 105-123 94-113 100-129 102-119 SD 20.85 13.61 4.95 8.14 5.16 7.42 5.65 4.61 SE 4.08 1.74 1.03 1.77 1.43 3.03 1.29 1.39 Tail length (mm) n 30 64 22 21 13 6 19 11 X 39.0 42.3 40.5 42.4 41.2 39.8 42.4 41.3 range 29-47 37-50 33-47 36-50 35-48 35-43 34-46 35-47 SD 4.64 3.81 3.14 4.91 4.59 2.71 3.46 4.00 SE 0.84 0.47 0.65 1.07 1.28 1.10 0.79 1.20 Hind foot (mm) n 30 62 22 19 13 6 18 10 X 10.0 10.0 10.5 10.5 10.2 10.3 10.3 10.5 range 8-12 8-13 9-12 8-13 7-13 8-13 8-12 8-13 SD 1.07 1.07 1.02 1.42 1.87 1.63 1.57 1.50 SE 0.19 0.13 0.21 0.32 0.52 0.66 0.37 0.47 Forearm (mm) n 24 46 23 20 10 6 17 13 X 45.8 46.5 45.8 46.9 46.2 46.5 46.1 47.3 range 42-48 42-50 41-54 42-50 43-49 44-51 44-50 44-50 SD 1.48 1.89 2.90 2.05 1.81 2.88 1.49 2.05 SE 0.30 0.27 0.61 0.46 0.57 1.17 0.36 0.57 Tragus (mm) n 11 29 6 6 2 3 3 6 X 6.5 6.9 6.8 6.2 5.8 6 6.2 6.9 range 6-7 5-8 5-7 6-8 5.5-6.0 6 6.0-6.5 4.5-6.0 SD 0.52 0.91 0.75 0.75 0.35 0 0.28 3.52 SE 0.15 0.16 0.30 0.30 0.25 0 0.16 1.44 Weight (grams) n 29 63 22 19 9 4 19 12 X 15.2 16.7 15.7 16.7 14.5 15.8 14.2 15.2 range 12.2-19.5 12.1-26.1 13.0-19.5 12.8-26.5 12.0-20.6 12.6-17.1 12.0-16.5 12.4-18.6 SD 2.59 2.82 1.85 3.9 2.48 2.12 1.42 1.68 SE 0.48 0.35 0.39 0.89 0.82 1.06 0.32 0.48 both as individuals and in groups. The bats were wedged so tightly that many were torn as they were removed with large forceps, and many myobiid mites attached to their abdomens were squashed. This same barn was visited in midwinter the next year, but no bats were found in these crevices. We suspect that some of the bats found in large numbers in buildings in summer squeeze into narrow crevices in the same buildings for hibernation, while some migrate to caves (in southern Indiana?). However, we have no idea where the majority of big brown bats summering in Indiana spend the winter. Much more work is needed on this. Our winter population data (gathered from 1951 to 1975) for 67 Indiana caves show that 23 caves contained no big brown bats; 33 caves had from 1 to 10; in 7 caves 11 to 24 were present; 1 contained 25 to 50; and 3 Order / Chiroptera • Family / Vespertilionidae 176 Map 21. The Big Brown Bat, Eptesicus fuscus, in Indiana caves contained more than 50. The largest numbers of big brown bats were found in Donaldson's Cave and Donnehue's Cave (both in Lawrence County) and in Tunnel Cave (Jefferson County), where maxima of 74, 58, and 93 were found, respectively. Kirkpatrick and Conaway (1948) reported "about 200" big brown bats in Donaldson's Cave on 29 December 1946. To our knowledge, this is the largest number recorded in an Indiana cave. It is surprising that relatively few big brown bats hibernate in Indiana caves, since the species is so plentiful in summer in the state. It is also interesting that certain caves appear to attract a particular species of bat and other caves do not. This is true for Eptesicus fuscus as it is for most (or all) native species that hibernate in caverns. The three Indiana caves that contain the largest winter populations of E. fuscus may have been favored traditionally by this bat, although data from Donaldson's Cave, gathered in the early 1900s by Hahn, indicate that very few big brown bats used this cave during that period. Donaldson's and Donnehue's caves have streams running throughout their entire length; Tunnel Cave is dry. In 1906 and 1907, Hahn found small numbers of big brown bats in the caves where we have worked. He stated (1908b) that six was the largest number of big brown bats he found in one cave; this was in Mayfield's Cave (Monroe County) on 21 December 1906. Banta (1907) recorded this species only in Mayfield's Cave on four occasions between January 1903 and August 1905. We found none in this cave on 26 January 1952. Hahn studied the caves on the UniversityFarm (now within Spring Mill State Park] from September 1906 to September 1907. He reported E. fuscus as "comparatively rare" and collected only three specimens (Hahn, 1908b). He does not list the species in his paper on the mammalian remains of Donaldson's Cave (Hahn, 1907b). Further evidence of the rarity of big brown bats in caves in Spring Mill State Park and in other southern Indiana caves is Hahn's statement that he found "only ten of this species" in Indiana and Kentucky caves he visited. Certainly, the status of Eptesicus fuscus had drastically changed in Donaldson's Cave by 1946, when Kirkpatrick and Conaway found 200 present on a single visit. We have records of 74 in this cave on 12 February 1955, only 12 on 22 January 1962, and 3 on 13 February 1975. Again, we do not know how much these figures actually reflect population changes. A single visit to a cave per winter does not provide a reliable estimate of the hibernating population of big brown bats (or any other species). We have practically no records of Eptesicus fuscus in caves during June or July, but our visits to caves during this season have not been numerous or systematic. Ten June and July trips to seven caves known to harbor big brown bats in winter resulted in a single observation—two big brown bats in Donnehue's Cave on 22 July. No doubt some caves are used periodically for night roosting by this species in summer, but we need more study of this phenomenon. In August, a few big brown bats begin to roost by day in some caves, and this number gradually increases until the winter population is more or less stable, from about mid-December to midEptesicus fuscus Big Brown Bat 177 February, when, weather permitting, the bats may begin leaving the caves. ASSOCIATED SPECIES. Eptesicus fuSCUS hibernates in caves used by all other species of bats that utilize caves in this season. In a concrete culvert at Lafayette, studied for 20 years, a single Plecotus rafinesquii was the only other bat found in winter with E. fuscus. Possibly this hibernaculum is too cold for less hardy species to winter there successfully. Big brown bats do not mix with other species in the caves. Many hang singly; others are found in small groups; the largest aggregations are invariably found in crevices, and large clusters are not formed on the walls or ceilings. Also, big brown bats tend to hang nearer the cave entrance than do other bats; this is especially true in winter. When feeding, big brown bats have been shot along with red bats, hoary bats, evening bats, pipistrelles, silver-haired bats, Indiana myotis, and little brown myotis. On 3 September, 3 big brown bats and 12 little brown myotis were found behind the window shutters on an occupied house. Our notes do not indicate whether separate shutters were used by each species; there were twelve shutters, all on one side of the house. In the summer of 1948, big brown bats, Indiana myotis, Keen's myotis, and little brown myotis all roosted by day beneath a large concrete bridge in Turkey Run State Park. The species were separated from one another, however. An evening bat was shot at dusk on 8 May as it emerged from the roof of a house; the following evening, a big brown bat was shot as it flew from the same exit hole. HABITS. Eptesicus fuscus appears to be quite hardy and able to withstand more cold than most other species of bats found in Indiana. For example, on 5 March 1954, there were 4 big brown bats in a crevice just outside the mouth of King's Cave (Harrison County). Four feet directly below the bats was a pool of water covered with one-fourth inch of ice. In a somewhat similar situation, 11 females were found in a crevice outside the mouth of May's Cave (Monroe County) on 20 February 1954. Many big brown bats that hibernate in caves choose sites near the cave mouth, where considerable air movement and freezing temperatures may occur. Most other bats of this species are found in winter in the front sections of the caves. In an intensive study of Donnehue's Cave, only 2 big brown bats were observed more than 364 feet from the cave entrance (Mumford, 1958). Since this is a hardy species, it would not be surprising to learn that it also winters in holes in trees. One is drawn to this conclusion by several facts: there is a dramatic decrease in numbers of big brown bats from summer to winter in Indiana; banding has failed to establish that there are extensive migrations out of Indiana in the fall; the known number hibernating in buildings does not account for the summer population; Eptesicus fuscus moves in and out of at least some of its hibernacula throughout the winter. Clearly, caves are not required for successful hibernation; thus, until we obtain more data on the use of buildings by big brown bats during the winter, we feel that trees may be housing a large part of the wintering bats. It seems that E. fuscus is really less of a cave-dweller than are the various native species of Myotis or Pipistrellus subflavus (Hahn, 1909), but perhaps it is in the process of changing its habits in this regard. We have some evidence that big brown bats more readily seek caves in colder weather. There were 12 in Ray's Cave (Greene County) on 30 January 1975, but 18 to 20 there on 13 February 1975; during this period the outside temperature dropped to 10 to 15 degrees below zero Fahrenheit. In the caves, most big brown bats hang singly on the side walls. They often use small crevices formed by flakes of limestone that have become partially separated from the walls. Both vertical and horizontal crevices are used; some bats in a horizontal crack will be lying on their backs. Bats often wedge themselves into narrow places from which they cannot be removed without injury to the bats. Choice crevices normally are occupied annually. There were 40 big brown bats in such a site (Tunnel Cave) on 6 January 1954; the 18 that could be extracted for examination were all males. SEX RATIOS. The sex ratios of wintering Eptesicus fuscus evidently differ from place to place. In the culvert studied in Tippecanoe County from 1951 to 1971, the total known population for the period consisted of 178 Order / Chiroptera • Family / Vespertilionidae 29 males, 86 females, and 2 for which sex was unrecorded. Males outnumbered females here in only two seasons of those studied. This culvert was used by only 3 to 12 big brown bats per winter, so our sample is rather small. Sex ratios of big brown bats banded during hibernation in caves in southern Indiana generally favor males, with some exceptions. In Donnehue's Cave, 40 males, 34 females, and 2 unsexed individuals were banded during one winter season. For three different winters combined, there were 86 males, 40 females, and 15 unsexed big brown bats in Donaldson's Cave, about ten miles away. In May's Cave, on 20 February 1954, we found 1 male, 18 females, and a bat that could not be captured for examination. We have no data on the survival of bats that choose to hibernate in heated buildings. On the Purdue University campus, 29 big brown bats have been captured between 26 November and 14 March during the past 17 years. Some of these animals were found on the ground (some alive, some dead), but most were found flying about in buildings. Of these, 17 were males, 8 were females, and the sex was not recorded for 4. Most of those examined had very little or no fat, even during the middle of the hibernation season. This may indicate that temperatures in the buildings are too high for successful hibernation. We need data on banded individuals to clarify this point. On 10 March, a big brown bat was found on the ground (alive) in the snow at Ft. Wayne (Allen County); its pelage was covered with what appeared to be soot. Perhaps the bat had entered a chimney shortly before being found. A male banded in Kosciusko County on 23 July 1959 was found in that county on 24 December 1960. Nonhibernating big brown bats have been found in and on buildings, in caves, on rock escarpments, beneath concrete bridges, and in covered wooden bridges. About buildings they have been found in attics, hanging on the side of a brick structure, in vines on the side of a building, beneath loose tarpaper on the sides of a house, behind window shutters, in the rafters of small, open-sided structures, and between the stone side of a building and the gutter downspout. MIGRATION. Banding has revealed little about the movements of Eptesicus fuscus. Perhaps the species migrates little, or perhaps the lack of recoveries of banded animals is because most of the population winters in unknown sites. A female banded in Donnehue's Cave on 9 January 1954 was killed near Sullivan, 52 miles westnorthwest, on 16 April 1955. A male banded at the same cave on 9 January 1954 was found alive near Elkton, Kentucky (142 miles south-southwest, on 31 December 1954. A female banded at May's Cave on 20 April 1954 was killed near Hidalgo, Illinois (85 miles west), in April 1962. And a female banded in Donaldson's Cave on 12 February 1955 was captured and released in a cave near Georgia, Indiana (7 miles west), on 29 November 1958. Each year in August we receive telephone calls and letters from homeowners about bats in houses, garages, or breezeways. Most of these animals that we have examined were big brown bats. There is undoubtedly considerable movement in August, when bats probably wander from summer to winter habitats; transients evidently enter houses and other buildings temporarily at this season. Frequently an individual is seen only for a day. Many of the known maternity colonies in Indiana are in unheated barns, from which the bats depart in winter. Although a considerable number of big brown bats has been banded in these summer colonies, few animals have been recaptured at sites other than where banded. Thus, we know the species moves from summer to winter localities but we do not know the distances involved in these movements. Banding has revealed little seasonal or annual changes of cave roosts by E. fuscus. Most marked individuals have been recaptured in the same caves where they were banded. HOMING. Cope et al. (1961) conducted several homing experiments with Eptesicus fuscus in southeastern Indiana. From 70 to 90 percent of 20 big brown bats released 20 miles west of the capture site homed within 4.5 to 9 hours. About 32 to 40 percent of 47 released 40 miles south of another site returned, but none came back before two nights had elapsed. An estimated 8 of 11 returned after three nights after being released 100 miles north of their colony. From four to Eptesicus fuscus Big Brown Bat 179 six nights after release, 31 of 36 homed from 250 miles south. Only 1 of 18 released 250 miles north returned to the colony, after an absence of four nights. The experimental animals were all adults (males and females) and the work was carried out in July and August. We can offer no explanation for the difference in homing performance in these animals. The variation in the percentage that successfully homed from 250 miles south and 250 miles north is interesting. On 30 August 1954, a sample of 41 males and 48 females was taken from a maternity colony in Bedford, banded, and released later that night at Terre Haute (80 miles northwest). The colony was next visited in April 1955, when at least 5 banded males and 9 banded females were present. Since this group was banded with regular aluminum bands only, it is possible that some individuals were not located visually in the colony. Cope and his co-workers had used radioactive bands on their animals and were able to detect unseen bats in the roosts. LONGEVITY. Of 12 males and 11 females banded during hibernation and recovered three or more years later, 1 male and 1 female were at least seven years old, 4 females and 2 males were at least six years old, 2 males and 2 females were at least five years old, 2 males and 1 female were at least four years old and 5 males and 3 females were at least 3 years old. The age of none of these animals was known at the time of banding. Another sample of unknown age, banded during hibernation, contained 1 male and 2 females at least three years old, 1 male and 2 females at least four years old, 1 female at least five years old, 1 female at least six years old, and a female at least nine years old. HABITS. Evidently Eptesicus fuscus wakens and becomes active periodically during the winter. On unusually warm evenings (temperature around 60°F) in January and February, big brown bats may be seen in flight on the Purdue University campus, where they hibernate in buildings. Gwilym S. Jones saw a big brown bat flying slowly over Farmersburg (Sullivan County) in the pre-dusk on 21 February 1976; the day was clear, but a storm was moving in. Winter observations in caves also reveal that a particular bat does not remain long in one spot, but moves about throughout the hibernating period. Some individuals enter and leave the caves all winter long and evidently have alternate (unknown) roosts which they occupy when absent from the caves. We suspect that the prevailing winter temperatures outside the hibernacula are correlated with these movements. The extent of winter movement in and out of Donnehue's Cave was reported by Mumford (1958). Although 109 big brown bats used this cave from August to April, the maximum number observed on any one of 76 visits to the cave was 39 (36.1 percent). More than 30 bats per visit were seen on 3 and 29 December, 7, 12, and 16 January, and 17 February—the three coldest months of the year. Temperature minima for five of these dates ranged from 19°F to 28°F; the minimum for the other date was 33°F. From August 1970 to August 1971 and in February 1973 we netted the entrance to Donnehue's Cave on twenty nights, including each month but January. We captured big brown bats each month but April, again demonstrating winter activity of this species (Mumford and Whitaker, 1975). The big brown bat is a fairly early flyer at dusk, but does not emerge from its daytime roost as early as do red bats or pipistrelles. We have often shot big brown bats in the early evening while the sky was fairly bright. In netting Donnehue's Cave, we caught only one big brown bat within the first half hour and six within the first hour after sunset. In the second, third, fourth, and fifth hours after sunset we netted 16, 14, 13, and 6 big brown bats respectively. Since we did not tend the nets later than 12:20 A.M., we do not know whether this species was active all night. A net at the mouth of Big Wyandotte Cave (Crawford County), tended from 7:20 P.M. to 6:30 A.M. the night of 22-23 August, took E. fuscus during each half-hour period from 8:15 P.M. to 5:10 A.M. Of the 100 captured 66 were taken between 10:30 P.M. and 2:30 A.M.; during this four-hour period, we netted 13, 20, 19, and 14 respectively per hour. The time of emergence from a maternity colony in a building was correlated with the time of sunset and, presumably, therefore with light intensity. As the days became shorter, the bats began emerging earlier in the evening (Table 60). Ilahn (1908a) estimated that E. fuscus flew 180 Order / Chiroptera • Family / Vespertilionidae faster than 10 to 12 mph, but his figures were largely guesswork. FOOD. Hahn (1909) reported that in captivity Eptesicus fuscus ate "any kind of meat." A female shot at dusk on 24 July had a chafer (Cyclocephala borealis) in her mouth. The stomachs of 184 big brown bats were examined for food (Table 61). Ground beetles (Carabidae) formed the greatest volume of food. These were probably taken on the wing. The food second in abundance was Scarabaeidae, and third was the spotted cucumber beetle, Diabrotica undecimpunctata (Chrysomelidae), common in stomachs in late summer. (It is not clear why this particular species was so common in stomachs, but this agricultural pest was seen in numbers at lights.) Thus the three most heavily consumed foods were beetles — indeed, beetles formed 49.1 percent of the entire volume of food in the sample. The fourth most prevalent food taken was stinkbugs, which occurred in nearly a third of the stomachs and constituted nearly 10 percent of the volume of food. Most were a bright green species, possibly Aero sternum hilare (Say). It was not clear why stinkbugs were so often taken. Most of the forms mentioned above (48.0 percent of the total volume) were large, being at least a centimeter in total length. Flying ants were fifth in abundance, sometimes occurring in numbers of 10 to 50 per stomach. Crickets (Gryllidae), although including a few flying forms, constituted a part of the nonflying food, which also included vegetation, Curculionidae, insect larvae, Diplopoda, and mites. The total volume of food considered to be of nonflying forms was 4.3 percent. Four bats had eaten internal organs of larger insects, possibly Scarabaeidae, presumably by splitting the insect open and removing the abdominal contents. The cockroach eaten was apparently Parcoblatta, a flying form. No cellular structure was apparent in the unidentified amorphous material. It had the appearance of milk seen in the stomachs of juveniles, and was found in female bats with young. Vegetation consisted of grass stems in one stomach (about full), and 95 percent wheat seeds in another. Other items taken from stomachs of big brown bats in trace amounts and not included in the table were click beetles (Elateridae), insect larvae, rhysodid beetles, syrphid flies, mites, millipedes, and various other flies (Otitidae, Chironomidae, and Mycetophilidae). Data were available from several sources to determine whether big brown bats feed in winter. The stomachs of 178 bats taken from Table 60 Emergence of Eptesicus fuscus, at 5-minute intervals, from a maternity colony, Bedford, Indiana, 1954 (from Mumford and Whitaker, 1975) Time Periods P.M. 9 Aug. Date and Number Leaving Roost 18 Aug. 29 Aug. 31 Aug. 8 Sept. 7:20 to 7:25 — — — — 4 7:25 to 7:30 — — — — 30 7:30 to 7:35 — — 1 1 82 7:35 to 7:40 — — 9 1 70 7:40 to 7:45 — — 20 7 44 7:45 to 7:50 — 2 65 28 21 7:50 to 7:55 — 17 38 48 12 7:55 to 8:00 4 62 31 44 8:00 to 8:05 11 54 36 34 8:05 to 8:10 47 75 7 3 8:10 to 8:15 60 45 13 8:15 to 8:20 72 23 8:20 to 8:25 83 19 8:25 to 8:30 49 10 8:30 to 8:35 16 8:35 to 8:40 7 Eptesicus fuscus • Big Brown Bat 181 Table 61 Foods eaten by Eptesicus fuscus (n = 184) from Indiana (from Whitaker, 1972b) Percent Percent Food Item volume frequency Carabidae (ground beetles) 14.6 29.8 Scarabaeidae (scarab beetles) 12.4 25.0 Chrysomelidae, Diabrotica undecimpunctata 11.5 21.7 Pentatomidae (stinkbugs) 9.5 29.3 Formicidae (ants) 8.5 16.8 Coleoptera 6.6 15.2 Ichneumonidae (ichneumon flies) 5.0 14.1 Lepidoptera 4.5 7.6 Cicadellidae (leafhoppers) 4.4 20.7 Insects (unidentified) 3.1 11.4 Chrysomelidae (leaf beetles) 3.0 9.8 Gryllidae (crickets) 3.0 9.2 Diptera 1.8 7.1 Reduviidae (assassin bugs) 1.8 7.6 Insects (internal organs) 1.5 2.2 Trichoptera (caddis flies) 1.4 3.3 Vegetation 1.1 1.6 Amorphous material (unidentified) 0.9 1.6 Alleeulidae (comb-clawed bark beetles) 0.8 2.2 Delphacidae (planthoppers) 0.7 3.3 Hymenoptera 0.5 1.6 Muscoid flies 0.5 0.5 Chrysopidae (lacewings) 0.5 2.7 Curculionidae (snout beetles) 0.4 2.7 Blattidae (roaches) 0.3 0.5 Coccinelidae (ladybird beetles) 0.2 0.5 Miridae (leaf bugs) 0.2 2.2 Coreidae (squash bugs) 0.2 0.5 Tipulidae (crane flies) 0.2 1.1 Mydaidae (mydas flies) 0.1 0.5 Calliphoridae (blow flies) 0.1 0.5 Rhagionidae (snipe flies) 0.1 0.5 Nabidae (nabid bugs) 0.1 0.5 Anthieidae (anthicid beetles) 0.1 0.5 Chironomidae (midges) 0.1 0.5 Total 99.7 A few foods found in trace amounts are listed in the text. November through March (not included in Table 61) were examined. All were empty except for one from December, which contained gray amorphous material, apparently not food. Many of these bats were active and apparently had been flying around, since they appeared in and on the outside of buildings. Four bats were caught as they entered Donnehue's Cave in the late fall (one on October 30, two on November 18, and one on December 2) in 1970. The stomachs and intestines of all were completely empty. The digestive tracts of 17 torpid big brown bats taken from a barn in Vigo County on 26 March 1970 were empty, except for small masses of insect material in the posterior end of the intestines of some. This apparently had been present for a long time, judging by the discoloring of the material and by the amount of digestion which had taken place, even of the hard parts. Another group of 11 bats was taken in the third week of October 1965, also at Terre Haute. The intestines of all and the stomachs of 10 were empty. One specimen, 194 Order / Chiroptera • Family / Vespertilionidae Table 65 Foods eaten by Lasiurus borealis (n = 128) from Indiana (from Whitaker, 1972b) Percent Percent Food item volume frequency Lepidoptera 26.2 32.0 Scarabaeidae (scarab beetles) 12.1 21.9 Delphacidae (planthoppers) 11.8 25.8 Formicidae (ants) 6.3 11.7 Cicadellidae (leafhoppers) 5.9 20.3 Coleoptera (unidentified beetles) 5.6 21.9 Carabidae (ground beetles) 5.6 10.9 Reduviidae (assassin bugs) 4.1 14.1 Muscoid flies 2.5 6.3 Hydrophilidae (water scavenger beetles) 1.8 3.9 Diptera 1.8 7.0 Insects (unidentified) 1.6 4.7 Gryllidae (crickets) 1.6 4.7 Miridae (leaf bugs) 1.2 3.1 Chrysomelidae (leaf beetles) 1.1 3.1 Curculionidae (snout beetles) 1.0 3.9 Soft material 0.8 0.8 Dirt 0.8 0.8 Alleculidae (comb-clawed bark beetles) 0.8 0.8 Tipulidae (craneflies) 0.8 3.1 Nabidae (nabid bugs) 0.8 0.8 Fulgoridae (lanternflies) 0.8 0.8 Pentatomidae (stinkbugs) 0.7 2.3 Lygaeidae (chinch bugs) 0.7 0.8 Blattidae, Parcoblatta (roaches) 0.6 0.8 Ichneumonidae (ichneumon flies) 0.6 4.7 Chrysopidae (lacewings) 0.5 1.6 Cbironomidae (midges) 0.5 0.8 Trichoptera (caddisflies) 0.4 1.6 Vegetation 0.4 2.3 Anthomyidae (anthomyid flies) 0.2 0.8 Hymenoptera 0.2 1.6 Syrphidae (hover flies) 0.2 0.8 Flesh (baby bat or placenta?) 0.2 0.8 Chrysomelidae, Diabrotica undecimpunctata 0.1 0.8 Araneae (spider) trace 0.8 Ephemeridae (mayflies) trace 0.8 Drosophilidae (pomaceflies) trace 0.8 100.3 prominent. Jackson (1961) thought that red bats might feed from foliage, or even near the ground, since crickets (Gryllus) were found in their stomachs. REPRODUCTION. We have been unable to obtain definite data regarding copulation in Indiana Lasiurus borealis. While hunting or observing red bats at dusk, we have frequently seen one bat pursue another; several of these "chases" have resulted in the two bats making actual contact in the air. In one case (9 April), the bats came into contact while flying 30 feet above the ground, fell together until about 5 feet from the ground, then separated. We have no way of knowing the sexes of the bats involved or if they were actually copulating. We have recorded chases in April (two), June (two), July (eight), August (nine), September (four), and October (one). On 24 July and 4 September, one of the two Lasiurus borealis • Red Bat 195 participating in a chase was shot; in each case it was a male. Two others, shot as they made contact on 15 August, were both males. Thus, one wonders whether observed chases have anything to do with copulation. Thomas W. Hoekstra watched a red bat chase another on 12 April; the bats came together, then fell to the surface of a road, where Hoekstra captured them. One was a male, the other a female. He was of the opinion that they were copulating. On 5 September, Peter Phendler caught two red bats that were lying on the ground (evidently fighting); both were males, one of which tested positive for rabies. We have found that testis size in red bats is largest in July, August, and September (Whitaker and Mumford, 1972c). If testis size can be taken as an indicator of the mating season, we assume that most copulation takes place during these months. Hamilton (1943) collected female red bats with sperm in their uteri the first week of August. Most young of red bats are evidently born around mid-June in Indiana, but actual dates of parturition are quite difficult to obtain from noncaptive individuals. Females near term may abort their young prematurely in confinement, so one should be cautious in accepting parturition dates for captive (frequently injured) gravid females. We found small (1.8 mm diameter) embryos in a female collected 30 April, and embryos as late as 19 June (10 mm crown-rump length). Since we have seen embryos as large as 22 mm crownrump length, the female taken 19 June was not at term. The 32 gravid red bats examined each contained from 2 to 5 embryos. The average number of young per female was 3.2 and the mode was 3. A summary of the number of embryos per female follows: 2 young (1 female); 3 young (24 females); 4 young (6 females); 5 young (1 female). We have records of 33 adult female red bats (6 June-26 July) with attached young; many of these were found on the ground. Of these females, 14 had 4 young, 12 had 3, 6 had 2, and 1 had 1. It was surprising to note that such a large percentage had 4 young per litter, because from embryo counts 3 young per litter was the most common. Perhaps the fact that a female is carrying 4 young makes her more vulnerable to storms or other disturbances that may dislodge her from her roost and cause her to fall to the ground. Placental scars were observed in 19 females. Seven of these animals exhibited scars in June and 10 in July; none was found in females taken in August. One individual said to have been taken on 3 May had placental scars, as did another female supposedly taken on 18 September. Both of these dates seem unusual, in light of the other data; perhaps there were errors in the dates; both females were among bats found and sent to the Indiana State Board of Health. Five other adult females collected in September had no visible placental scars, nor did six taken in August. It appears that placental scars do not persist for long periods after parturition. A female found with four attached young averaging 74 mm in total length (1 July) contained four faint scars. Another female taken with three attached young averaging 75 mm in total length on 20 June had at least two very faint scars. In the collection of Manchester College is an adult female red bat and five small young of equal size preserved in alcohol. No label accompanies the specimens, but it is possible that they represent a female and her litter. We have examined females in lactation from 14 June to 3 August, but no doubt these dates could both be extended with more data. GROWTH OF YOUNG. There are evidently few weight and size data for newborn red bats. Barbour and Davis (1969) lacked information on this point, and Jackson (1961) must have erred in stating that they weighed only 0.5 grams each at birth. We have not examined known, full-term, newborn red bats, but from a few observations we have made some interpolations. On 17 June a female with two young, still carrying umbilical cords, were found on the ground. Forearm measurements for these young were 12 and 13 mm. The largest embryos we have examined in a gravid red bat measured 22 mm in crownrump length and had total lengths of 44, 44, and 41 mm. Three embryos measuring 20 mm each in crown-rump length had forearms of 10 mm each. We feel that these must have been near term. Another female was found on the ground with 3 attached young on 20 June. These young had forearms measuring 24, 21, 196 Order / Chiroptera • Family / Vespertilionidae and 20 mm and weighed 3.1, 2.9, and 2.5 grams respectively. This family represents the smallest attached young that we have data for and may be an indication of the size of young at, or shortly after, parturition. The largest young we have found with their mother was on 26 July. When disturbed, two of the young flew away and the other glided to the ground and was captured. It weighed 8.4 grams, was 85 mm in total length, and had a forearm length of 38 mm. Three young found with a female on 14 July measured 79, 79, and 81 mm in total length and weighed 6.4, 7.4, and 7.6 grams respectively. An immature male shot in flight on 4 July weighed 6.6 grams, was 81.5 mm in total length, and had a forearm of 36 mm. More information is needed to determine at what size Lasiurus borealis young become independent, but the animals mentioned here may have been near this stage. Stomach analyses of immatures provided some interesting information (Whitaker and Mumford, 1972c). One young weighing only 4.4 grams and having a total length of 64 mm had 60 percent chironomid and 40 percent coleoptera remains in the stomach, but no milk. All of the 29 other immatures found without accompanying females and with total lengths of less than 80 mm and having food in their stomachs contained milk. But 2 of them (forearms 27 and 31 mm) also contained parts of ants and other insects. Two lone immatures, collected 28 July and 11 August, contained only insect remains and thus were evidently independent. They had total lengths of 85 and 86 mm and forearms of 36 and 39 mm respectively. One young, measuring 91 mm in total length and with a 38-mm forearm, had milk in its stomach and intestines. This indicates that some young, though able to feed on insects, continue to nurse, and some become quite large before they are off milk. PARASITES AND DISEASES. There is relatively little information on external parasites. Tipton and Boese (1958) described Steatonyssus furmani from a red bat taken in Jefferson County. A report of Steatonyssus occidentalis on L. borealis by Whitaker and Wilson (1968) was in error; the two individuals found were juvenile S. furmani. Two larval specimens of the tick Dermacentor variabilis were taken from a red bat in Tippecanoe County (Wilson, 1961). Whitaker (1973) reported on the ectoparasites of 234 Indiana red bats. Of the two species of parasites found, 59 were macronyssid mites (Steatonyssus furmani) and 13 were a new species of myobiid mite (Acanthopthirius lasiurus) recently described by Fain and Whitaker (1976). Particularly large numbers of trematodes and a few cestodes were found in the intestines of Indiana red bats. None of these internal parasites has been identified. Rabies was diagnosed in a red bat from Tippecanoe County in September 1960, and 24 of 338 Indiana red bats examined for rabies by Whitaker and Miller (1974) tested positive. In this study the red bat ranked third in incidence of rabies, following the hoary bat and the pipistrelle, both of which were represented by small numbers. DECIMATING FACTORS. Several authors have reported the attacks of blue jays on red bats (Strecker, 1924; McClure, 1942; Allan, 1947; Elwell, 1962; Hoffmeister and Downes, 1964; Mumford, 1973). This type of predation might be of significant importance in the extensive area where the summer ranges of the red bat and the blue jay overlap. Evidently the jay discovers roosting red bats while it is foraging among tree foliage. Other avian enemies of red bats are known, including hawks and owls. Mumford found the skull of a L. borealis in the pellet of an unidentified owl from Jackson County. Workers at the Jasper-Pulaski Fish and Wildlife Area watched captive ring-necked pheasants pursue and kill a red bat that got inside the pheasant pen. Automobiles kill some foraging animals; we have eight records of red bats found either dead on roads or on the radiators or front ends of cars. Various accidents have befallen other individuals. One was found alive in a swimming pool, the sides of which were evidently too high and too smooth for the bat to climb out. Another was found caught on a fence. Several have been found in insect light traps (Wilson, 1965), to which they may have been attracted by the insects or by the sound produced by certain types of traps ("black" lights). One was found, dead, imLasiurus cinereus • Hoary Bat 197 paled on the ripe seed head of a burdock plant (Arctium minus). At least nine red bats have been found dead along the beach at the south end of Lake Michigan; four of these were found by Raymond Grow in midSeptember. The implication is that these were migrants caught over the lake by adverse weather conditions and forced into the water. TAXONOMY. Lasiurus borealis borealis (Muller) is the subspecies in Indiana. SELECTED REFERENCES. Barbour and Davis, 1969; Constantine, 1966; Davis and Lidicker, 1956; Kunz, 1968; McClure, 1942; Mumford, 1973; Whitaker, 1972b, 1973. Hoary Bat Lasiurus cinereus (Palisot de Beauvois) Vespertilio pruinosis: Plummer, 1844 Atalpha cinereus: Langdon, 1881b Atalaphacinerea: Evermann and Butler, 1894b Lasiturus cinerus: Engels, 1933 Lasiurus cinereus: McAtee, 1907 DESCRIPTION. Lasiurus cinereus is a large, grayish bat with short, broad, rounded ears bearing black borders. It is the largest species of bat in Indiana. The whitish tips of the body hairs give the bat the appearance of being covered widi hoarfrost, hence its common (vernacular) name. The dorsal surface of the interfemoral membrane is completely furred. The throat is yellowish, and there is a small, irregular yellowish spot at the base of each thumb. Immatures are somewhat paler in color than adults. Individual hairs on the back often have four bands of color—a blackish base, then yellowish, followed by blackish or brown near the whitish tip. Males and females are colored alike, in contrast to the sexually dimorphic coloration of Lasiurus borealis. The skull of L. cinereus is squarish and similar to that of L. borealis, but larger. Weights and measurements are shown in Table 66. An immature male taken 20 July weighed 18.4 grams and had light fat. An adult female taken on 9 August and an imaged male collected 28 August were both very fat, weighing 33.6 and 21.0 grams respectively. The dental formula is I —C — P—M— = 32. 3 12 3 STATUS AND DISTRIBUTION. The hoary bat no doubt occurs throughout Indiana, but is Hoary bat. Photo by Bruce J. Hayward probably uncommon. Specimens are now in museums from 27 counties (Map 24). Reliable sight records or reports of lost and discarded specimens are at hand from additional counties. Findley and Jones (1964) summarized the seasonal distribution of the hoary bat throughout its contiguous range, and Mumford (1969b) supplemented their data for Indiana. Of 83 records of occurrence in the state for which the month is known, 52 are for the months of June (29) and July (23). Most dates fall between 12 April and 5 October; the majority of these are for May to August, evidently the period when hoary bats are most abundant in Indiana. There are 198 0rder / Chiroptera • Family / Vespertilionidae Table 66 Weights and measurements of Lasiurus cinereus from Indiana Males Females Total length (mm) n 20 41 X 124.0 134.8 range 110-138 102-152 SD 8.7 9.6 SE 1.9 1.5 Tail length (mm) n 19 39 X 52.8 56.5 range 44-64 49-65 SD 5.4 5.0 SE 1.2 0.8 Hind foot (mm) n 19 41 X 10.2 10.6 range 8-14 6.0-13.5 SD 1.5 1.9 SE 0.3 0.3 Forearm (mm) n 16 35 X 52.2 54.9 range 47-57 42-59 SD 2.4 3.4 SE 0.6 0.6 Tragus (mm) n 14 37 X 7.8 7.9 range 5-10 5-11 SD 1.5 1.7 SE 0.4 0.3 Weight (grams) n 10 27 X 18.1 28.5 range 14.0-28.2 18.0-38.3 SD 4.1 6.1 SE 1.3 1.2 single records for 20-29 October (Allen County), 21 November and 18 December (both Tippecanoe County), and 31 January (Vigo County). There is a good possibility that some individuals are present in winter. Since the species is presumed to be highly migratory, considerable movements must take place across the state in spring and fall, but we have little indication of the numbers of animals involved. Breeding occurs throughout Indiana, although we have relatively few actual records of females with attached young. Map 24. The Hoary Bat, Lasiurus cinereus, in Indiana Since Lasiurus cinereus is a tree-inhabiting bat, accurate censusing is virtually impossible. At times, several may be seen foraging together or coming to watering areas. On the evening of 29 May 1948, at least three were feeding together along the border of a woodlot (Tippecanoe County); two of them were shot and both were females. Thomas W. Hoekstra and J. Scott Grundy shot five hoary bats at dusk on 27 June 1962, as the bats flew over a small pond near Munster (Lake County); four were females and one a male. Bats were numerous there that evening, but all appeared to be the same species, according to Hoekstra. James B. Cope and his students netted three hoary bats the night of 12 July 1962, near Richmond (Wayne County). HABITAT. The hoary bat is considered to be essentially a tree-inhabiting bat, roosting by day among the foliage. It appears to show a definite tendency to roost in areas where trees are scattered and have openings below their crowns. Such known roosting sites in Indiana include a pasture, residential areas, a Lasiurus cinereus • Hoary Bat 199 wooded lake shore, a wooded campus, and an open stand of mixed woodland, brush, and old fields. Two individuals were taken in buildings; another was found clinging to the side of a building. There are a few records of skulls and other remains of hoary bats in Indiana caves (Hahn, 1907b; Mumford, 1953b), but neither a live nor an intact animal has been discovered in caverns in this state. It would be interesting to know at what seasons and under what conditions hoary bats enter caves. Most Indiana specimens of L. cinereus have been shot or netted about woodlot borders, along small streams through pasture or cropland, or over ponds (mostly in unwooded areas). Eight females were shot at Lafayette around the borders of a pastured, deciduous woodlot surrounded on three sides by pasture and on the fourth by a cultivated field. Judging by the frequency with which hoary bats are submitted to the Indiana State Board of Health, residential areas may be used by this species to a considerable extent. ASSOCIATED SPECIES. Other bats shot along with hoary bats at dusk include big brown bats, Indiana myotis, pipistrelles, silver-haired bats, red bats, and evening bats. Red bats are a common associate in Indiana, possibly indicating that both species of Lasiurus find certain habitats to their liking. A man tearing down a small building on his residential lawn (Lafayette) on 21 November found a hoary bat and three smaller bats in the building. He destroyed the smaller bats before giving the L. cinereus to Mumford. HABITS. The hoary bat is primarily a solitary species, except at favored feeding and drinking areas, where three or more may be seen together. Family groups consisting of a female and her two young are also encountered. It appears to us that the hoary bat emerges to forage a bit later on the average than does the red bat, although some hoary bats have been observed in the evening when it was still relatively bright. Many of the individuals that have been noted early in the evening were flying quite high (often out of shotgun range). Some of these passed overhead in rather straight flight and may have been animals moving from their daytime roosts to feeding or drinking areas some distance away. As dusk deepens, the bats seem to forage nearer the ground. This may partially explain why fewer hoary bats have been collected by shooting. The flight of the hoary bat is usually strong, fast, and direct. The wingbeats seem regular, rather deliberate, and somewhat birdlike. Amazingly erratic flight may be exhibited by foraging individuals, especially on windy evenings. On calm evenings, hoary bats have been watched lazily circling about 200 feet in the air for periods of up to twelve minutes. One watched (and later shot) on a windy evening seemed to feed mostly into the wind, then it climbed slightly and "hung" motionless for a few moments, before going into a downward glide on set wings. We think that L. cinereus glides more than do other Indiana species of bats when feeding. The size of the foraging area has not been precisely determined, but our impression is that an individual bat ranges rather widely. On one occasion, however, a single hoary bat was kept under observation for ten minutes, as it fed near the ground and over an area probably not more than 50 feet in diameter. As many as three hoary bats have been noticed feeding together (two of them were collected) about the border of a woodlot; the bats fed mostly from 12 to 25 feet from the ground and roughly 15 yards from the edge of the woods, over the edge of a pasture. On six evenings from 14 June to 12 July, a hoary bat (presumably the same each night) left its daytime roost (not discovered) in a large tree in a residential area between 8:16 and 8:35. Its behavior was about the same each evening; it appeared to emerge from an area about 40 feet above the ground, circle briefly, then fly directly away until out of sight. The bat appeared each evening at about the same time after sundown. Only a few hoary bats have been found in their day roosts; four roost sites were in trees. On 15 June an adult female with two attached young was hanging 25 feet from the ground in a thornless honey locust. A male was captured in a honey locust at an unspecified height above the ground on 29 August. On 26 September, Fred Miller found one hanging 6 feet from the ground on a green fruit of a black walnut tree in a pasture. The bat hung in full sunlight. A female was caught in an unidentified tree on 6 May and a female with two young was roosting in a box elder tree on 14 June. A male was captured in a small building on 21 November; it hung 200 0rder / Chiroptera • Family / Vespertilionidae with three smaller bats (which we were unable to obtain for examination). Another male was found clinging to the side of a brick building on 31 January (Whitaker, 1967b). An adult female was captured while clinging to the roof inside a ring-necked pheasant pen constructed of poultry netting supported by poles. When the bat Miller found was stroked with a finger, it snapped its jaws and gave the usual chittering noise characteristic of agitated hoary bats. Most observations and collections of hoary bats have been made at dusk. The activity patterns of the species are largely unknown. An adult female was netted at 12:20 A.M. on 29 July. Adult male hoary bats are evidently quite rare in Indiana. One was taken 31 May in Jennings County. Males collected 21 November (Tippecanoe County) and 31 January (Vigo County) could not be definitely aged as young of the year or fully adult. In contrast, we have some 40 records of adult females from the state. We have about 40 additional records of hoary bats collected in Indiana from July to November, but accurate age data are not available for most of them. Little is known about the winter habits of this species, but most authors indicate that hoary bats migrate south to spend the winter. The evidence for this is somewhat scanty and not based on recoveries of marked animals. FOOD. We have accumulated relatively little information regarding the food of Lasiurus cinereus in Indiana. Three specimens had insects in their stomachs. Two contained 100 percent Lepidoptera and the third contained 65 percent Lepidoptera, 25 percent Chrysomelidae, and 10 percent muscoid flies. One individual taken in July had only vegetation in its stomach. Details of collection were not available, but vegetation is probably not a normal food. Possibly the bat had been hurt and confined to the ground, where it consumed plant material as an emergency food. A hoary bat taken 31 January (Vigo County) after a warm period of four to five days, had a "plug" of well digested grass stems in the posterior end of the intestine. This material appeared to have been there for a considerable period. Fresh grass stems and some shed snake skin were in the stomach (Whitaker, 1967b). Ross (1967) examined the digestive tracts of 139 hoary bats from New Mexico. Of these, 136 contained Lepidoptera (Phalaenidae, Geometridae, Gelechiidae) in numbers up to 25 per bat. Other insects present were Formicidae (78 individuals), Acrididae (1), Isoptera (7), Chrysopidae (1), Coleoptera (9), Scarabaeidae (4), Chrysomelidae (2), and Cerambycidae (1). Poole (1932) found a pentatomid and a culicid in a stomach of L. cinereus from Pennsylvania. Bishop (194/) found a hoary bat feeding on Pipistrellus subflavus in New York, and Orr (1950) observed one pursuing a small bat in California. Low numbers of coleopterans have been found in stomachs of this species, and Brisbin (1966) reported L. cinereus as reluctant to take beetles. Overall, this species is primarily a moth feeder. REPRODUCTION. Each of eleven gravid hoary bats collected in Indiana from 5 May to 8 June contained two embryos. Parturition evidently occurs mostly in late May and early June. A female captured on 28 May (Vigo County) gave birth (perhaps prematurely) to two young the same day. Three adult females with attached young have been recorded on 14 June (Wayne County), 15 June (Tippecanoe County), and 19 June (Vigo County). Two partially grown young were obtained (without a female) on 12 June (Clay County) and a small young was collected on 30 June (Wayne County). Four laetating females were collected between 28 May and 14 July. A female taken 5 May contained embryos only 6 mm in crown-rump length. The largest embryos examined by us (1 June) had forearm lengths of 16.5 mm each and were probably near term. These two young weighed 3.9 and 4.0 grams. Two young born to a captive female on 28 May had forearm lengths of 15 and 17 mm and total lengths of 55 and 57 mm respectively. Two embryos, each with 16-mm forearms, examined 29 May, weighed 3.3 and 3.5 grams, while two others on 3 June had 15-mm forearms and weighed 3.1 grams each. From these data, it appears that newborn hoary bats have forearms of about 16 mm, total lengths of from 55 to 57 mm, and weights of about 3.5 to 4.0 grams. Our measurements are nearly identical with those of Bogan (1972), although he presented no weights for newborn young. Barbour and Davis (1969) stated that Lasiurus cinereus • Hoary Bat 201 Table 67 External parasites of several species of bats from Indiana, listed by host, in order of decreasing abundance Parasites Hosts parasitized Species examined Total Average Total Percent Southeastern Myotis 5 Mites (Acarina) other than chiggers 60.0 Olabidocarpus whitakeri 52 10.40 3 60.0 Chigger mites (Trombiculidae) 20.0 Euschoengastia pipistrelli 1 0.20 1 20.0 Indiana Myotis 43 Mites (Acarina) other than chiggers 32.6 Macronyssus crosbyi 70 1.63 14 32.6 Paraspinturnix globosus 6 0.14 4 9.3 Acanthophthirius lucifugus 2 0.05 1 2.3 Silver-haired Bat 8 Batbugs (Cimicidae) 12.5 Cimex adjunctus 1 0.13 1 12.5 Mites (Acarina) other than chiggers 62.5 Macronyssus macrodactylus 19 2.38 5 62.5 Acanthophthirius sp. 1 0.13 1 12.5 Eastern Pipistrelle 56 Chigger Mites (Trombiculidae) 23.2 Euschoengastia pipistrelli 54 0.96 13 23.2 Evening Bat 12 Mites (Acarina) other than chiggers 41.7 Steatonyssus ceratognathus 12 1.00 5 41.7 Red Bat 234 Mites (Acarina) other than chiggers 11.1 Steatonyssus furmani 59 0.25 26 11.1 Acanthophthirius lasiurus 13 0.06 7 3.0 Hoary Bat 25 Mites (Acarina) other than chiggers 8.0 Pteracarus completus 2 0.08 2 8.0 Acanthophthirius lasiurus? 13 0.52 1 4.0 newborn young have forearm lengths of 18 to 19 mm. We have few data on growth rates of young hoary bats from Indiana. Relatively few young have been examined and all kept in confinement died within seven or eight days, thus weight and measurement data for them are probably not meaningful. An immature female was shot in flight on 18 July; it had a forearm length of 53 mm and a total length of 114 mm. It was flying slowly and steadily in a straight line, with rather deliberate wingbeats. It exhibited none of the driving, erratic flight characteristics of adults and probably had not been independent very long. A young male, shot in flight on 19 July, had a 53-mm forearm and was 135 mm in total length. An immature male collected 6 July weighed 17 grams, had a forearm of 51 mm and a total length of 124 mm; the specimen label does not indicate whether this animal was capable of flight. Data for adult males are lacking. Testis size was not recorded for the only adult male we examined. A male taken 31 January had testes 2.4 mm long. Testis lengths for nine subadults varied from 3 to 6 mm and averaged 4.1 mm. PARASITES AND DISEASES. Whitaker (1973) examined 25 Indiana hoary bat specimens, but the only external parasites found were 13 unidentified individuals of Acanthopthirius and 2 other myobiid mites originally listed as Pteracarus chalinobus but now reidentified as Pteracarus completus. Table 6/ summarizes data on external parasites from several species of bats in Indiana. 202 Order / Chiroptera • Family / Vespertilionidae Nixon A. Wilson found the cestode Cycloskrjabinia taborensis in a hoary bat from Indiana (Stunkard, 1962). Whitaker and Miller (1974) found 10 of 51 Indiana hoary bats positive for rabies. DECIMATING FACTORS. One hoary bat specimen obtained had flown into a guy wire. Boys with a BB gun shot a female carrying two young from her daytime roost in a tree. Another family was found on the ground after the wind blew its roost tree down. McAtee recorded a female and two large young that were chased from their daytime roost in a tree by a robin. On 18 December 1976, the wings and most of the skin of a hoary bat were found beneath some pines being used as a roosting site by long-eared owls (Tippecanoe County). The bat remains were quite fresh, and circumstantial evidence suggests the owls captured the bat. TAXONOMY. The subspecies in Indiana is Lasiurus cinereus cinereus (Beauvois). SELECTED REFERENCES. Barbour and Davis, 1969; Poole, 1932; Provost and Kirkpatrick, 1952; Mumford, 1969b: Whitaker, 1967b. Rafinesque's Big-eared Bat Plecotus rafinesquii Lesson Big-eared Bat Corynorhinus macrotis: Butler, 1895 Corynorhinus megalotis: Allen, 1916 Corynorhinus rafinesquii: Miller, 1924 Corynorhinus townsendii: Hall and Kelson, 1959 (probably) Plecotus rafinesquii: Handley, 1959 DESCRIPTION. This medium sized bat is brownish above, whitish below, and has extremely long ears (more than one inch long) unlike those of any other species of bat in Indiana. The hairs on the dorsum are dark brown or blackish on the basal two-thirds, then pale reddish or yellowish brown on the distal one-third. The ventral hairs are black or blackish at their bases, with white or whitish tips. The throat and chin may be about the color of the back in some individuals. As the pelage becomes worn, the overall color of the dorsum becomes darker. Weights and measurements are shown in Table 68. The dental formula is I —C — P—M — = 36. 3 13 3 STATUS AND DISTRIBUTION. PleCOtUS rafinesquii is rare in Indiana and seemingly restricted to the south-central portion of the state. Most of our records are from Lawrence and Washington counties (Map 25); three are from two small caves about 2 miles northeast of Saltillo and eight are from the caves on Spring Mill State Park, east of Mitchell. The last observation known to us from Spring Mill was made in November 1907. Two individuals of Plecotus (presumably this species) were seen and one collected near Greencastle (Putnam County) on 26 December 1894. A big-eared bat was collected (Ohio State Museum 1741) from a small cave near Kossuth (Washington County) on 25 October 1955; another was captured in a cave near Smedley (Washington County) on 21 December 1962. The presence of a Plecotus rafinesquii at West Lafayette (Tippecanoe County) on 13 December 1959 was a surprise; this specimen represents the northern-most known record of this species in the United States (Wilson, 1960). HABITAT. All records except one of Plecotus rafinesquii for Indiana are from caves; a single specimen was taken from a long, concrete culvert. The caves where the big-eared bat has been found in Lawrence and Washington counties are located in areas of deciduous woodlands. The Spring Mill State Park caves where Walter L. Hahn made his observations of this species are located in a solid, near-virgin tract of large trees. The Washington County sites are not so extensively wooded, but have cultivated and wooded areas intermixed. We do not know what the appearance of the Putnam Countysite was in 1894, when Plecotus was found Plecotus rafinesquii • Rafinesque's Big-eared Bat 203 Rafinesque's big-eared bat. Photo by Philip C. Shelton there, but today it is sparsely wooded and mostly cultivated land. The culvert mentioned above is located beneath a welltimbered slope near the Wabash River; this forested area has probably been relatively undisturbed for many years. It would appear, then, that P. rafinesquii is a forest-inhabiting bat. In other parts of its range it has frequently been found in buildings. Some of the caves housing big-eared bats in Indiana have a stream flowing throughout their length, but some are without streams. The culvert where a big-eared bat was collected frequently has the floor covered with water, as it was the day the specimen was taken there. ASSOCIATED SPECIES. The caves at Spring Mill were also used by little brown myotis, Keen's myotis, Indiana myotis, southeastern myotis, gray myotis, pipistrelles, and big brown bats when Hahn made his observations there on the big-eared bat. The Plecotus rafinesquii found near Kossuth on 25 October 1953 was the only bat noted in the cave. Two big-eared bats captured on 28 December 1954 shared the cave with six pipistrelles and one little brown myotis. Big brown bats also used the culvert where a Plecotus was taken at West Lafayette; this big-eared bat represents the only other species of bat observed in this culvert during periodic checks between December 1946 and the winter of 1972-73. Table 68 Weights and measurements of Plecotus rafinesquii from Indiana Character n x Range SD SE Total length (mm) 6 96.5 84-104 8.0 3.3 Tail length (mm) 6 45.8 43-50 2.8 1.1 Hind foot (mm) 5 10.0 9-11 0.7 0.3 Ear (mm) 6 32.1 27-35 1.1 2.7 Tragus (mm) 5 13.6 13-15 0.9 0.4 Forearm (mm) 6 43.8 43-45 0.4 1.0 Wing expanse (mm) 3 276.3 225-305 25.7 44.6 Weight (grams) 3 10.6 10.2-11.2 0.5 0.3 204 Order / Chiroptera • Family / Vespertilionidae Map 25. Rafinesque's Big-eared Bat, Plecotus rafinesquii, in Indiana HABITS. Most of the 17 big-eared bats observed in Indiana were single animals, but two were found in a cave on three occasions. Two found on 28 December were hanging about 2 inches apart on a cave wall, according to R. D. Kirkpatrick, who first observed them. Hahn observed six big-eared bats in the Spring Mill caves during the winter of 1906-07 and another during May 1907. He was making almost daily visits to these caves over that time period. The big-eared bat appears to prefer to roost, even in winter, in the twilight zone near cave entrances. Here, the temperature is often quite cool and air movement is sometimes considerable. The only other bat occurring in Indiana that chooses similar winter roosting sites is the big brown bat. Evidently both species are equally hardy. The Plecotus rafinesquii that Wilson captured in Tippecanoe County had entered the culvert between 6 and 13 December. Minimum daily temperatures at a nearby weather station during this period ranged from 22 to 37°F, and maximum temperatures ranged from 30 to 49°F. At the site where the bat hung the temperature was 48°F. Hahn (1909) reported two big-eared bats that "flew out into the cold air" from a cave on 22 February, but he did not indicate the air temperature at the time. Hahn (1908b) found that big-eared bats at Spring Mill in 1906-07 were all "not far within the cave." Other observers have found big-eared bats 25 feet within a small cave, two others about 60 feet from the cave entrance, and one 137 feet from a culvert entrance. Most were hanging from the cave wall, but the one in the culvert was suspended from the ceiling. Two individuals were hanging separately about 6 feet from the cave floor, and two were hanging near each other 12 feet from the floor of a cavern. Evidently this bat is easily aroused from hibernation or is more alert than most other bats during the winter. Hahn was able to capture only two of the seven he observed. A few minutes after Kirkpatrick found two hanging near each other, one had moved to a new location in the cave. When the bats are sleeping, their long ears are usually curled along either side of the neck, and the end of the forearm of the bat overlaps the ear. As the bats awaken, the ears are uncurled slowly until they are straight. A male found in deep torpor on 28 December had the ears laid along the neck and shoulders, but the ears were uncurled. Despite Hahn's intensive work in the Spring Mill caves and the fact that he observed more big-eared bats than any other person in Indiana, he did not see the species flying about at dusk and decided that it was a late flyer, probably not emerging until the light was too dim for observation. Bats of this species that Hahn kept in captivity were swift and steady flyers. Like Hahn, we have never seen a Plecotus in flight outside a cave, and none have been collected by shooting bats at dusk in Indiana. FOOD. We have no data on the food habits of this species in Indiana, but we assume that it feeds heavily on moths. Captive animals kept by Hahn refused to eat and soon died. REPRODUCTION. No Plecotus has been recorded from Indiana during the breeding season. A male taken 13 December had testes Plecotus rafinesquii • Rafinesque's Big-eared Bat 205 measuring 2 by 4 mm, but single males collected 28 December and 12 January had scrotal testes measuring 4 by 7 mm. PARASITES AND DISEASES. We have no data from Indiana specimens. DECIMATING FACTORS. No data are available from Indiana. TAXONOMY. The subspecies in Indiana is currently known as Plecotus rafinesquii rafinesquii Lesson. Other names that have been applied to Indiana big-eared bats appear in the synonymy at the beginning of this account. On the assumption that the two Plecotus mentioned by Butler (1895) from Putnam County were P. townsendii, some authors have indicated that P. townsendii occurs in Indiana. The specimen that Butler examined (now lost) was never seen by these authors, but Hall and Kelson (1959) mapped the range of P. townsendii across Indiana. Since all extant specimens of Plecotus from the state are rafinesquii, we think P. townsendii should be eliminated from the Indiana list. We are aware that both P. townsendii and P. rafinesquii breed in Kentucky. Clearly, further research on this genus is needed in our region. SELECTED REFERENCES. Barbour and Davis, 1969; Hall, 1963; Handley, 1959; Hoffmeister and Goodpaster, 1963; Jones and Suttkus, 1975; Wilson, 1960.