ISSN 0327-9383 Mastozoología Neotropical; 1(2): 103-112 SAREM, 1994 THE IMPACT OF AN ERUPTION OF VOLCAN HUDSON ON SMALL MAMMALS IN ARGENTINE PATAGONIA Oliver P. Pearson Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, U.S.A. ABSTRACT: Sixteen months alter an explosion of Volcan Hudson had covered parts of Santa Cruz Province in Argentina with 5 to 15 cm of ash that had killed thousands of sheep, the native mice were present and thriving. Eligmodontia, Phyllotis, and Reithrodon were present in greater-than-expected numbers, and Abrothrix xanthorhinus was present in normal numbers. All were reproducing vigorously and appeared to be healthy. Molar teeth of Eligmodontia and Phyllotis were not abraded noticeably more than in other regions. RESUMEN: Impacto de una erupción del volcán Hudson sobre pequeños mamíferos en la Patagonia argentina. Dieciséis meses después de que una explosión del volcán Hudson cubrió con una capa de 5 a 15 cm de ceniza parte de la provincia de Santa Cruz (Argentina) y produjo la muerte de cientos de ovejas, los ratones nativos estaban presentes y en expansión. Eligmodontia, Phyllotis y Reithodon fueron encontrados en mayor número al esperado, y Abrothix xanthorhinus fue encontrado en su nivel de abundancia normal. Todos los ejemplares estaban activos reproductivamente y en condiciones saludables. Los molares de Eligmodontia y Phyllotis no presentaron mayor desgaste que en otras regiones. Key words: Argentina, volcano, mouse populations, Volcan Hudson INTRODUCTION Volcanic eruptions can destroy all lifenearby and can affect plant and animal populations hundreds of kilometers distant, but the impact of volcanic activity on small mammals has rarely been studied. One of the most pertinent reports traced the recovery of small-mammal populations following a period of activity of Volcan Parícutin in 1943 in Mexico (Burt, 1961). Most vertebrates were exterminated out to the zone where less than about 20 cm of ash fell. Even 17 months after the initial eruption, animal life was practically absent where die ash was 15 cm or more deep. Another study (Andersen and Mac Mahon, 1985; Mac Mahon et al., 1989) documented the recovery of smallRecibido 22 Febrero 1994. Aceptado 25 Junio 1994. mammal populations for seven years following the brief but spectacular eruption of Mount St. Helens in the United States in 1980. Mammals were trapped in many different habitats that had suffered different kinds and degrees of volcanic mayhem. The sites at Mount St. Helens most useful for comparison with the South American eruption described below were those subjected to relatively gentle ashfall. Pumice and ash 7-30 cm deep near Mount St. Helens caused substantial mortality to shrubs and forbs but not trees; herbaceous vegetation recovered quickly where tephrafall was less than 4.5 cm. Three species of small rodents and one species of shrew persisted in a high tephrafall area four months after the eruption and, in general, popu- 104 lations of those species persisted over the next seven years while the vegetation was recovering. On 12-15 August, 1991, a volcano in southern Chile, Volcan Hudson, erupted for the first time in 20 years and sent a blanket of ash and pumice across Argentina to the Atlantic Ocean. By 20 August the ash was detected in Australia (Smithsonian, 1991: N° 7), and within a month injection of Hudson aerosols finto the stratosphere was contributing to a substantial ozone loss over Antarctica (Hofmann et al., 1992). Satellite photographs on 13 and 14 August show dramatically the clearly defined plume that dropped more than 2 km3 of ash on about 100,000 km 2 of Santa Cruz Province, Argentina, and contributed to the death of thousands of sheep (López, ms). In Argentina 5 to 15 cm of ash fell along the southern shore of Lago Buenos Aires near the towns of Los Antiguos and Perito Moreno (Smithsonian, 1991, N° 8; Corbella et al., 1991; López, ms). Los Antiguos is 130 km SE of Volcan Hudson, so the impact of the eruption would have been more severe closer to the volcano in Chile. Much of the ash was re-distributed during the week of violent winds that followed the eruption, and even four months later strong winds were remobilizing enough fine ash to disrupt airport activity 1500 miles 2400 Km) away (Smithsonian, 1991, N° 11). Near Los Antiguos, cliffs of tephra 20 m high remind one that ashfall is not a rare event in this region; the flora and fauna have survived other volcanic eruptions in postglacial times (Smithsonian, 1991, N° 7). The area in Argentina covered by ash from the 1991 eruption is almost en- Oliver P. Pearson tirely semi-arid, bushy steppe devoted to sheep grazing. The ash that fell was not poisonous, although it was notably high in its content of sulfur (3780 mg/ kg) and fluorine (up to 280 ppm in plants) (Corbella et al., 1991; Lopez, ms; Smithsonian, 1991, N° 7). A frequency distribution of particle sizes is bimodal with peaks at 39 and 375 microns but with an appreciable percentage less than 5 microns (Corbella et al., 1991; Smithsonian, 1991, N° 9). News reports commented on irritation of the eyes of humans, livestock, and wild animals such as rheas and guanacos, as well as severe abrasion of vehicles, windmills, sheep shears, and other farm machinery. Contributing to the mortality of sheep was the drying up of small water sources in this semi-arid region by the deposition of tephra, and by the covering of palatable vegetation that was already scarce after several years of drought. The fleece on many sheep became so laden with ash that when it became wet it overburdened the sheep so that they could scarcely walk (Smithsonian, 1991, N° 9). General observations Between 4 and 7 December, 1992, 16 months after the eruption, we visited Los Antiguos and Perito Moreno to assess whether the 5-15 cm of ash had affected the populations of small mammals living there. On windy days the heavy surf on the southeastern shore of Lago Buenos Aires was still carrying appreciable quantities of pumice particles about 1 mm diameter, and windrows of pumice spheres 1 to 2 cm diameter lined the beach. These larger ejecta had probably floated from the Chilean end of the lake more than 60 km away. The vegetation, however, MOUSE POPULATIONS AFTER VOLCAN HUDSON was not covered with ash; most of the ash had blown away in the high winds for which Patagonia is famous. On windy days during our visit in December, 1992, clouds of ash could be seen blowing about in the mountains in Chile to the west of Los Antiguos, as well as local clouds of "dust". The morning and afternoon sky throughout Patagonia still had the brassy color associated with particulate matter in the upper atmosphere. The vegetation did not appear to have been seriously affected. Adequate rainfall had supported new growth. Bunchgras se s had trapped appreciable amounts of ash but were not killed, and ash had formed drifts behind many of the plants. The general aspect, however, was that of a normal, sandy desea habitat with most of the vegetation appearing to be healthy. We saw a number of hares, pichi armadillos, tinamous, and a skunk. On December 5 we counted five fresh hare careases on 20 km of the paved road along Lago Buenos Aires, and on December 7 we counted an additional five recently killed hares on 50 km of the same road between Los Antiguos and Perito Moreno. Tadpoles were noted in a stream 28 km W Perito Moreno. A count of animal tracks in the sandy soil suggested that animal life was abundant. On one morning, after a high wind during the preceding day had erased all tracks, we recorded the number of times that animal footprints crossed a straight line 50 m long, and another straight line 125 m long. This was in bunchgrass steppe at Site 2, described below. The first line was crossed by tracks of 2 mice, 2 hares, birds, and 1 pichi armadillo. The second line was crossed by tracks of 39 105 mice and 4 hares. This reflects abundant populations and a large amount of animal activity during one night. Trapping results Abundante: We sampled the smallmammal fauna with traps of a size appropriate for catching mice and rats (Museum Specials and large Sherman traps). I asume that the percent of traps that captured an animal each night (trap success) reflects the abundante of the various target species. Trapping results were as follows: SITE 1: Lago Buenos Aires, 30 km W Perito Moreno. On the night of 45 December, 73 traps were set in bushy steppe habitat with an estimated ground cover of 30 percent. Plants, listed in approximate sequence of decreasing coverage of the ground were: Senecio, duraznillo (Colliguaya), bunchgrasses, a very spiney, low cushion plant, and bushes of molle (Schinus marchandii). In spite of heavy rain during the night, which sprang a number of traps, 10 mice were captured, all Eligmodontia, and 1 Gecko. Trap success for small mammals was 14%. All Eligmodontia in this study are assumed to belong to the species morgani, in spite of the fact that their tails average 20 mm longer than those of the type and nine topotypes from the nearby type locality (Alíen, 1905). SITE 2: Estancia La Aurora, 4.4 km E Los Antiguos. This site is 21 km west of Site 1. On the night of 5-6 December 128 traps were set in two kinds of habitat, bushy steppe and bunch-grass steppe, and on the night of 6-7 December 24 traps were set in bushy steppe. This gives a total of 152 trap-nights. Overall trap success was 27% (Table 1). The bushy steppe had 30 — Spring A Autumn P 25 — E E A R A P E 0 SITE 2 Spring, 1992 II CHILE CHICO Spring, 1986 R III RIO NEGRO Springs P R IV RIO NEGRO Autumns R V CHILE CHICO Autumn, 1986 VI CHILE CHICO Autumn, 1987 Fig. 1: Springtime trap success in the ashfall zone near Los Antiguos (Site 2) 15 months after the eruption (I) compared with springtime trap success nearby at Chile Chico before the eruption (II), with springtime and autumn trap success in bushy steppe and bunchgrass steppe habitats in Rio Negro Province before the eruption (III and IV), and with trap success at Chile Chico in two autumns before the eruption (V and VI). A= Abrothrix, E= Eligmodontia, P= Phyllotis, R= Reithrodon. uosirad 'd Jamo A MOUSE POPULATIONS AFTER VOLCAN HUDSON about 35% ground cover; the dominant plants were duraznillo (Colliguaya) up to 4 m tall, bunchgrasses, neneo (Mulinum spinosum), paramela (Adesmia cf. boronioides), and molle (Schinus marchandii). The bunchgrass steppe appeared to have been burned within the past few years, leaving it with fewer bushes. The dominant plants were bunchgrasses, neneo, duraznillo, and calafate (Berberis). There had been no grazing by livestock at either of these two sites for many months. It may be seen • in Table 1 that success in capturing Eligmodontia at Site 2, near Los Antiguos, was greater than would be expected in the spring at the beginning of the season of reproduction. The capture of numerous Phyllotis xanthopygus was a surprise, since they are usually caught in cliffs and rocky places. No such habitat was nearby. The Coney Rat, Reithrodon, is notably reluctant to enter Museum Special and Sherman traps, so a trap success of 5% for this species indicates a sizeable population. The number of Abrothrix xanthorhinus captured was close to the "expected". Table 1: Trap success at Site 2; 5-7 December, 1992. Species Eligmodontia morgani Phyllotis xanthopygus Reithrodon auritus Abrothrix xanthorhinus Number Trap caught Success 16 12 7 4 "Expected" Trap Success 11% 8 5 3 2.5% 0.26% 0.43% 2.7% 27 % 5.89% (1) (2) (2) (2) (1)Springtime trap success in bushy steppe in the Province of Rio Negro averaged over several years (Pearson et al., 1987). (2)Springtime trap success in bushy steppe and bunchgrass steppe habitats, averaged over several years, in the Province of Rio Negro (Pearson, unpublished). 107 Through the kindness of Douglas Kelt, comparison can be made with captures he made before the eruption in very similar habitat near Chile Chico, less than 10 km west of Site 2. In the spring of 1986 in 1045 trap nights he had a trap success of 3.6%. His catch was dominated by Abrothrix xanthorhinus, included no Eligmodontia, but included one Abrothrix longipilis. This catch can be compared directly with the 27% trap success at Site 2 in spring of 1992 (Fig. 1).The 14% trap success at Site 1 was not included in this comparison; it is a minimal value because heavy rain during the night sprang many traps. The overall trap success for the four species in the spring season at Chile Chico in 1986 (Fig. 1) before the eruption (3.6%) was similar to that in steppe habitat in many spring seasons in Rio Negro Province (5.9%), but after the eruption, trap success (27%) was seven times greater. Eligmodontia, Phyllotis, and Reithrodon were much more prevalent in 1992, whereas Abrothrix was represented with about the same relative frequency in all three samples (Panels I, II, and III in Fig. 1). Kelt also trapped at Chile Chico in the autumn of 1986 and the autumn of 1987. For unknown reasons, mice at Chile Chico were spectacularly abundant in autumn of 1986 (overall trap success 65.7%; Panel V in Fig. 1). Overall trap success at Chile Chico in the more-typical autumn of 1987 (20.6%) was only moderately higher than in the autumn of many seasons in steppe habitat in Rio Negro Province (14.9%; Panels IV and VI, Fig. 1). Since reproduction during spring and summer by the four species represented in Table 1 and Fig. 1 usually increases autumn trap success by a factor of 108 about 1.8 in steppe and bunchgrass habitats (Pearson, unpublished), comparison of the post-eruption, springtime population should be compared only with springtime, pre-eruption populations. Such a comparison suggests that even if small-mammal populations had been seriously reduced by the ashfall, 15 months after the eruption they had recovered to levels that seem to be higher than usual. Reproduction: Reproductive data confirm that the mouse populations were thriving. Of 16 male Eligmodontia captured, 14 were in breeding condition, and the other two were juveniles less than 3 weeks old. Of 10 female Eligmodontia, 2 were lactating and the other 8 were pregnant carrying an average of 6.6 fetuses. The average number of fetuses in 20 pregnant females living in similar habitat in the Province of Rio Negro in earlier years was 5.9 (Pearson et al., 1987). Of the seven Reithrodons captured, three were sexually competent males, one was a juvenile female, and three were pregnant females with 4, 4, and 11 fetuses, respectively. The average number of fetuses in 17 pregnant Reithrodons in Rio Negro Province was 4.53 (Pearson, 1988). Of 11 Phyllotis examined, all 5 males were in breeding condition, 1 female was young and entering her first estrus, 1 female was parous, 2 were lactating, and 2 were carrying 9 and 8 fetuses, respectively. These are remarkably large litters for Phyllotis (Pearson, 1975). Of the four Abrothrix xanthorhinus captured, two were breeding males and two were pregnant females, each with six fetuses. Health: Pyke (1984) noted the con- Oliver P. Pearson dition of two kinds of mice (Peromyscus and Microtus) living in meadowsteppe habitat before, during, and after the eruption of Mount St. Helens. His study area, 400 km NE of the volcano, received a 2.2 cm layer of ash with particle sizes from 0.11 to 116 microns diameter, which is somewhat finer than the ash at Los Antiguos. No abnormalities of the eyes were noted in mice trapped before the eruption, but up to 10 days after the ash deposition numerous trapped Peromyscus showed loss of fur around the eyes and two had swollen upper eyelids. The eyes of Microtus specimens appear to have been unaffected. Histopathologic examination of affected specimens revealed no silicosis-like involvement of lungs or gastrointestinal tissue. When caged on a substrate of ash, Microtus and Peromyscus both developed severe swelling of the eyelids. Burt (1961) and Andersen and MacMahon (1985) did not note eye pathologies, but they were trapping closer to the source of the tephra so presumably the particles were coarser. All of the mice captured in Argentina appeared to be in good health. Irritation of the eyes, which had been reported in larger animals shortly after the eruption (Corbella et al., 1991), was not detected in our specimens. The lungs of one or more individuals of each species of mouse captured near Los Antiguos were examined under low magnification without detecting any pathology. Longevity: Skulls of all mice were saved for examination of the teeth. Unusually great wear of the molars wouid indicate either unusual longevity or unusually rapid wear resulting from life among the residual and re-distributed MOUSE POPULATIONS AFTER VOLCÁN HUDSON ash, which was notably abrasive (Corbella et al., 1991). Since few mice live more than one year, few if any of the captured animals would have been alive at the time of the eruption. Judging from toothwear, as measured by height of molar cusps (Pearson, 1975; Pearson et al., 1987), populations of Eligmodontia and Phyllotis at Sites 1 and 2 appear to have been composed mostly of old individuals, a c5 — 109 few juveniles only a few weeks old, and few or no individuals of middle -age (Fig. 2). This indicates that the breeding season for both species was just resuming after several months of sexual inactivity during the winter and early spring. The age pyramid for Eligmodontia at Sites 1 and 2 in December is quite similar to a pyramid for a population in springtime in Rio Negro Province. ELIGMODONTIA .30— Cil p. .40 — E.50— .30 1 30 1 1 30 % 1 30 1 1 1 1 1 1 30% December RIO NEGRO December SITES 1&2 d d PHYLLOTIS — .21— 6 .42— . .63— .84— E 1.05 — 1.36 — 30 1 1 q December SITES 2 1 30 % 30 1 1 1 I 1 I December NORTHERN PATAGONIA I 50 % Fig. 2: (Aboye). "Age" pyramid of Eligmodontia caught at Sites 1 and 2 at the beginning of the breeding season compared with that of Eligmodontia caught near the beginning of the breeding season in the Province of Rio Negro in earlier years. The December pyramid for Rio Negro was modified from Figure 6 of Pearson et al., (1987) by increasing the amount of toothwear in November by .05 mm, which corresponda to the rate of abrasion per month shown in Fig. 4 of that report. (Below). "Age" pyramid of Phyllotis at Site 2 at the beginning of the breeding season compared with that of the same subspecies near the beginning of the breeding season in earlier years in the Provinces of Neuquen, Rio Negro, and Chubut in Argentina. Cusp heights of these earlier specimens have been advanced in age an average of 22 days to bring them aIl to December 5, comparable to the pyramid for Site 2 in the fallout crea. The amount of this advance assumed abrasion at a rate of .007 mm per day (estimated from Fig. 2 in Pearson, 1975). 110 The pyramid for Phyllotis at Site 2 is compared with a pyramid of a population of the same species constructed from 21 available specimens collected in the spring (25 October - 1 December) in the Provinces of Neuquen, Rio Negro, and Chubut, Argentina. Toothwear of the Site 2 animals is only slightly greater than that of the sample from farther north, captured before the latest eruption of Volcan Hudson. DISCUSSION Sixteen months after the eruption, three of the species captured were those captured before the eruption during a much greater trapping effort by Kelt. No Abrothrix longipilis, an ominvore, were captured after the eruption, but the habitat is too arid to support many individuals of this species. Kelt had not captured any Eligmodontia before the eruption, whereas this seed and vegetation eater was common (11% trap success) after the eruption. Phyllotis, a seed and vegetation eater, and Reithrodon, an eater of green vegetation, were notably abundant after the eruption. The abundance of Abrothrix xanthorhinus, a seed-eater, was approximately at the level it had been during a springtime sample before the eruption and in similar habitat elsewhere in Argentina. Reproduction of Reithrodon and Phyllotis had ceased during the winter, as is usual, and was proceeding vigorously at the time of our spring collection. Fossorial, herbivorous, pocket gophers (Cratogeornys and Thomomys) survived the eruptions of Parícutin and 1VIount St. Helens in appropriate habitat that received less than 15 cm of tephra (Burt, 1961; Andersen and MacMahon, 1985). A rancher at our study site at Estancia La Aurora in- Oliver P. Pearson formed us that tuco tucos (Ctenomys), a fossorial herbivore similar to pocket gophers, had been present on the Estancia before the eruption in small numbers. We found no evidence of them during our post-eruption visit. Kelt captured specimens of Ctenomys before the eruption near Chile Chico, which is only a few kilometers west of Estancia La Aurora. Since the habitat at both of our study sites appeared to be suitable for tuco tucos, and since their digging activity makes their presence easily detectable, it is possible that the 5 to 15 cm of ashfall reduced the number of tuco tucos. Unfortunately we lack post-eruption data from Chile Chico and quantitative pre-eruption data from our sites. With the available data it is not possible to determine with certainty whether life in the abrasive ash from Volcan Hudson caused unusually rapid wear of the teeth, as it did in certain Mexican species after the eruption of Volcan Parícutin (Burt, 1961). Most of the Eligmodontia and Phyllotis at Sites 1 and 2 had greatly worn teeth, but they were obviously over-winter survivors of considerable age, and comparable populations from other regions also contained a high percentage of individuals with worn molars (Fig. 2). It seems unlikely that abrasion by ash from Volcan Hudson advanced molar wear in adult mice by more than a month or two. No pathologies attributable to tephra were detected. We conclude that despite possible immediate effects of the eruption and ashfall, 16 months later all four of these species of mice living in the ash zone were abundant, healthy, and reproducing vigorously. Eligmodontia, Phyllotis, and Reithrodon were more abun- MOUSE POPULATIONS AFTER VOLCAN HUDSON dant than expected. Possibly, their predators had not recovered, and possibly the amount and quality of food for mice had improved after the eruption because of reduced competition from sheep and other herbivores such as guanacos. ACKNOWLEDGEMENTS I am grateful to Douglas Kelt for generously sharing his trapping data from Chile, and to Javier Bellati and Carlos Lopez of INTA for providing important bibliographic sources. LITERATURE CITED ALLEN, J.A. 1905. Reports of the Princeton University Expeditions to Patagonia, 1896-1899. Vol. 3, Part 1. Princeton, 210 p. ANDERSEN, D.C., and J.A. MAC MAHON. 1985. The effects of catastrophic ecosystem disturbance: the residual mammals at Mount St. Helens. Journal of Mammalogy, 66:581-589. BURT, W.H. 1961. Some effects of Volcán Parícutin on vertebrates. Occasional Papers of the Museum of Zoology, University of Michigan, 620:1-24. CORBELLA, H., R. SCASSO, M. LUCERO, M. PALACIOS, P. TIBERI, P. RIAL, D. PEREZ, L. BARLETTA, R. RUIZ, C. PAZ, C. CHEPPI, P. ESPINA, J. RUBIO, L. CASTANON, H. MIGLIORA, E. QUARGNOLO, M. LAMOUREAUX, M. BREGLIANI, D. TORRES, A. CIBILS, J. GIACOMINI and D. PALACIOS. 1991. Erupción del Volcán Hudson - Agosto 1991 - Efectos sobre el territorio 111 de la Provincia de Santa Cruz. Waxen, N° 4, Publicación Científica, Universidad Federal de Patagonia Austral, Rio Gallegos. 29 p. HOFMANN, D.J., S.J. OLTMANS, J.M. HARRIS, S. SOLOMON, and T. DESHLER. 1992. Observation and possible causes of new ozone depletion in Antarctica in 1991. Nature, 359:283-287. LOPEZ, C.R. (manuscript). Una rápida evaluación de los efectos de la erupción del volcán Hudson en la Patagonia Argentina. EEA INTA Bariloche, Argentina. MAC MAHON, J.A., R.R. PARMENTER, K.A. JOHNSON, and C.M. CRISAFULLI 1989. Small mammal recolonization on the Mount St. Helens Volcano: 1980-1987. American Midlánd Naturalist, 122:365-387. PEARSON, O.P. 1975. An outbreak of mice in the coastal desert of Perú. Mammalia, 39:375-386. PEARSON, O.P. 1988. Biology and feeding dynamics of a South American herbivorous rodent, Reithrodon. Studies on Neotropical Fauna and Environment, 23:25-39. PEARSON, O.P., S. MARTIN, and J. BELLATI. 1987. Demography and reproduction of the silky desert mouse (Eligmodontia) in Argentina. P. 433-446 in Studies in Neotropical Mammalogy: Essays in honor of Philip Hershkovitz. B.D. Patterson and R.M. Timm (eds.), Fieldiana: Zoology, n.s., N° 39. 112 PYKE, D.A. 1984. Initial effects of volcanic ash from Mount St. Helens on Peromyscus mani-culatus and Microtus montanus. Journal of Mammalogy, 65:678-680. Oliver P. Pearson SMITHSONIAN INSTITUTION 1991. Bulletin of Global Volcanism Network, 16: Nos. 7:2-5; 8:1415; 9:2-3; 11:11-12.
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