THE SOUTHWESTERN NATURALIST 52(2):291–295

JUNE 2007

RAPTOR PREDATION ON ORD’S KANGAROO RATS: EVIDENCE FOR DIURNAL ACTIVITY BY A NOCTURNAL RODENT CLINT W. BOAL*

AND

MATTHEW D. GIOVANNI

U.S. Geological Survey, Texas Cooperative Fish and Wildlife Research Unit, Department of Range, Wildlife, and Fisheries Management, Texas Tech University, Lubbock, TX 79409-2120 (CWB) Department of Range, Wildlife and Fisheries Management, Texas Tech University, Lubbock, TX 79409 (MDG) Present address of MDG: School of Natural Resources, University of Nebraska, Lincoln, NE 68583 *Correspondent: [email protected] ABSTRACT—As a group, kangaroo rats (Dipodomys) are considered almost exclusively nocturnal. Diurnal activity by kangaroo rats has occasionally been noted, but the anecdotal nature of these accounts suggests it is a rare occurrence. While using video cameras and recorders to collect data on food habits at nests of ferruginous hawk (Buteo regalis) and Swainson’s hawk (B. swainsoni) on and near the Rita Blanca National Grassland in 2003 and 2004, we detected numerous D. ordii. We subsequently explored possible relationships between the frequency of capture of D. ordii by these raptors and the time of day and moon phase. We recorded 54 deliveries of D. ordii to 8 of 12 ferruginous hawk nests and 8 of 14 Swainson’s hawk nests. D. ordii was captured throughout the course of the day; there were no relationships between hour block of day or moon phase with the number of deliveries. Our review of other diet studies for ferruginous hawk, Swainson’s hawk, Harris’ hawk (Parabuteo unicinctus), and prairie falcon (Falco mexicanus) revealed kangaroo rats accounting for 1.1 to 6.6% of prey used. In contrast to the prevailing view that kangaroo rats are almost exclusively nocturnal, our data and those from other diet studies of some diurnal raptors suggest kangaroo rats might be more active during diurnal periods than previously thought. Diurnal activity by kangaroo rats might be related to local forage conditions. A more complete understanding of diurnal activity patterns by kangaroo rats will require more focused examination of their behavior in relation to vegetation and climate conditions. RESUMEN—Como grupo, las ratas canguro (Dipodomys) son consideradas casi exclusivamente de ha´bitos nocturnos. Actividades diurnas en las ratas canguro ocasionalmente han sido reportadas, pero la naturaleza anecdo´tica de estos registros sugiere que estas actividades son muy raras. Mientras usa´bamos ca´maras de video y grabadoras para recolectar datos sobre los ha´bitos alimenticios en los nidos del gavila´n herrumbroso (Buteo regalis) y del gavila´n langostero (B. swainsoni) en o cerca de Rita Blanca Nacional Grassland en 2003 y 2004, encontramos numerosas D. ordii. Subsecuentemente exploramos la posible relacio´n entre la frecuencia de capture de D. ordii por estas aves rapaces y la hora del dı´a y la fase lunar. Registramos 54 entregas de D. ordii a 8 de 12 nidos del gavila´n herrumbroso y a 8 de 14 nidos del gavila´n langostero. La captura de D. ordii fue durante el transcurso del dı´a. No hubo relacio´n entre la hora del dı´a ni la fase lunar con el nu´mero de entregas. Nuestra revisio´n de otros estudios sobre dietas del gavila´n herrumbroso, gavila´n langostero, halco´n Harris (Parabuteo unicinctus) y halco´n de la pradera (Falco mexicanus) revelo´ que las ratas canguro contabilizan entre el 1.1 y 6.6% de la presa usada. Contrario a la idea actual de que las ratas canguro son de ha´bitos casi exclusivamente nocturnos, nuestros datos y otros datos de estudios de dietas de algunas aves rapaces diurnas sugieren que las ratas canguro tal vez son ma´s activas durante periodos diurnos de lo que antes se pensaba. La actividad diurna de las ratas canguro puede estar relacionada con condiciones locales de forrajeo. Para entender mejor los patrones de actividad diurna de las ratas canguro se necesitara´ examinar con mayor detalle su comportamiento con relacio´n a las condiciones climatolo´gicas y vegetales.

Among prey animals, times of activity will normally be times of increased mortality risk (Halle, 2000). The timing of prey activity, therefore, has a substantial potential to affect

the extent of predation risk (Lima and Dill, 1990). Nocturnal behavior and moonlight avoidance might be behavioral adaptations to reduce predation risk. As a group, kangaroo rats

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The Southwestern Naturalist

(Dipodomys) are widely considered as exclusively nocturnal (Kenagy, 1976; Lockard, 1978; Garrison and Best, 1990; Reichman and Price, 1993; Daly et al., 2000), even reducing activities on clear or moonlit nights (Lockard and Owings, 1974; O’Farrell, 1974; Kaufman and Kaufman, 1982). However, diurnal activity has occasionally been noted (Moore, 1929; Lockard, 1978; Daly et al., 2000), but the anecdotal nature of these accounts suggests it is a rare occurrence. Ord’s kangaroo rat (D. ordii) is described as active only at night (Jorgensen and Hayward, 1965; Kenagy, 1976). Langford (1983) reported activity of D. ordii was greatest during the first hour after sunset and then, to a lesser extent, several hours before dawn. Kaufman and Kaufman (1982) found D. ordii activity was low during the first hour following sunset and greatly reduced on moonlit nights. Furthermore, a review of predators of D. ordii identified only species that were nocturnal (e.g., Bubo virginianus, Asio otus) or primarily nocturnal (e.g., Vulpes macrotis) in foraging behavior (Garrison and Best, 1990). This would suggest that D. ordii is indeed almost exclusively nocturnal and subject to nocturnal predators. It was therefore intriguing that we detected numerous D. ordii among the prey used by diurnally active ferruginous hawks (Buteo regalis) and Swainson’s hawks (B. swainsonii) in our study of prey use by sympatric grassland raptors (Giovanni and Boal, unpubl. data). This led us to investigate the frequency of occurrence of D. ordii in the diets of raptors in our study. Additionally, assuming delivery time corresponded to time of capture, we examined possible temporal patterns of diurnal activity by D. ordii. Finally, we review other food habit studies of diurnal raptors to assess the uniqueness of our findings, and discuss our results in context of reported and assumed nocturnal behavior of kangaroo rats in general. METHODS—We used video cameras and time-lapse recorders to collect food habits data at ferruginous hawk and Swainson’s hawk nests (Giovanni, 2005). Video data were collected during the hawk breeding seasons (i.e., May to July) of 2003 and 2004. We recorded data daily from 0530 h to 2100 h. A time stamp was included on the video data so that the times of all prey deliveries were known. All nests under video surveillance were located on rangelands, grasslands, and pasturelands in Dallam County, Texas; Cimarron County, Oklahoma; and Union County, New Mexico. Full descriptions of the study area and the methods of prey identification are presented in Giovanni (2005).

vol. 52, no. 2

FIG. 1—Number of Ord’s kangaroo rats (Dipodomys ordii; n 5 54) observed delivered during 1-h blocks of time at 8 Swainson’s hawk (Buteo swainsoni) and 8 ferruginous hawk (B. regalis) nests in Union County, New Mexico; Cimarron County, Oklahoma; and Dallam County, Texas, 2003 and 2004. To supplement our data, we reviewed published food habits data for diurnal raptors in western North America (Bednarz, 1988; Snyder and Snyder, 1991; Olendorff, 1993; Steenhof, 1998; Catron et al., 2004). Due to different methods in collection of data on food habits, the quality and resolution of prey use information varied among studies. However, by compiling the data, we were able to synthesize a broader understanding of diurnal raptor use of kangaroo rats.

RESULTS—We recorded 54 deliveries of D. ordii to ferruginous hawk and Swainson’s hawk nests. More of these occurred in 2003 (n 5 35) than 2004 (n 5 19). The majority of captures in 2003 were by Swainson’s hawks (77%), but ferruginous hawks captured slightly more in 2004 (58%) than did Swainson’s hawks (42%). In 2003, we detected D. ordii among the prey used by 3 of 6 (50%) ferruginous hawk pairs and 5 of 6 (83%) Swainson’s hawk pairs. Although the number of D. ordii captured in 2004 was less than in 2003, they were more common among ferruginous hawk pairs (i.e., 5 of 6; 83%) but less common among Swainson’s hawk pairs (3 of 8; 38%). Ord’s kangaroo rats seemed to be captured throughout the course of the day (Fig. 1), and there was no correlation between hour block of day and number of deliveries (r 5 20.173, P 5 0.537). When examining the moon phase during each of the 54 D. ordii deliveries, we found 12 (22%) captures occurred during each of the first and second quarters, 14 (26%) during the third quarter, and 16 (30%) during the fourth quarter. Thus, it seemed there was no relation-

June 2007

Boal and Giovanni—Diurnal activity by a nocturnal rodent

ship between phase of the moon and vulnerability of kangaroo rats to predation during diurnal periods. In a review of 20 ferruginous hawk studies, Olendorff (1993) found 412 kangaroo rats among a total of 6,203 prey items, accounting for 6.6% of all reported prey and 0.7% of prey biomass. Ord’s kangaroo rat accounted for 381 of the 412 kangaroo rat prey, but kangaroo rats were not detected in the diets of ferruginous hawks in all 20 studies (Olendorff, 1993). Bednarz (1988) found remains of 53 Dipodomys among 856 Swainson’s hawk prey remains and remains of 25 Dipodomys among 889 prey remains of the sympatric Harris’ hawk (Parabuteo unicinctus) in New Mexico. Steenhof (1998) reported that 68 of 1,671 prey remains recovered from prairie falcon (Falco mexicanus) aeries in Idaho from 1974 to 1980 were Dipodomys. More recently, Catron et al. (2004) found 3 D. ordii among 260 prey remains collected at ferruginous hawk nests in New Mexico. Snyder and Snyder (1991) observed a Swainson’s hawk in Arizona bring a banner-tailed kangaroo rat (D. spectabilis) to its nest in mid-afternoon and found a second banner-tailed kangaroo rat at a different Swainson’s hawk nest. Unfortunately, there are no delivery time data from the majority of these studies with which capture times could be estimated. DISCUSSION—Kangaroo rats are reputed to be almost exclusively nocturnal (Kenagy, 1973; Lockard, 1978; Genoways and Brown, 1993; Daly et al., 2000). In contrast to this prevailing view, R. K. Chesser (pers. comm.) and we have observed kangaroo rats out during daylight hours, albeit rarely. Our data and those from other diet studies of some diurnal raptors (e.g., Olendorff, 1993) suggest some kangaroo rats might be more active during diurnal periods than previously thought. Arguments could be posed that much of the diet data we report here might be derived from kangaroo rats scavenged as roadkills or captured during crepuscular periods when the raptors might also be active. We do not consider either explanation to be sufficient for the frequency of kangaroo rats captured by hawks. First, although scavenging is common among some North American raptors such as bald eagles (Haliaeetus leucocephalus; Buehler, 2000), golden eagles (Aquila chrysaetos; Kochert et al., 2002), and caracaras (Caracara cheriway; Morrison,

293

1996), scavenging by accipiter and buteo hawks seems to be rare. The few accounts of such behavior are primarily anecdotal observations, highlighting the rarity of such behaviors among these species (Chesser, 1979; Stalmaster, 1980; Bacon, 1994; Squires, 1995; Scheffield and Jobe, 1996; Pranty, 2002). There is no mention of Swainson’s hawks scavenging or using carrion in the comprehensive species account by England et al. (1997). Ferruginous hawks are known to occasionally scavenge hunter-killed prairie dogs (Chesser, 1979; Bechard and Schmutz, 1995), but the accounts seem to be primarily associated with wintering birds, which might be related to periods of food shortage or to inexperienced juvenile hawks. Furthermore, the rarity of accipiter and buteo hawks consuming carrion is reflected in the trapping methods used for capturing these species; no experienced raptor researcher attempts to capture these hawks with dead prey (Bub, 1978; Bloom, 1987; Hull and Bloom, 2001). Second, the pattern of crepuscular activity by D. ordii is unclear. For example, Langford (1983) found activity of D. ordii was greatest during the first hour after sunset and then, to a lesser extent, several hours before dawn, and O’Farrell (1974) reported D. ordii did not emerge until 37 min after sunset; in contrast, however, Kaufman and Kaufman (1982) found D. ordii activity was low during the first hour following sunset. Our video data of prey deliveries clearly demonstrated that, assuming delivery time correlates to capture time, kangaroo rats were captured throughout the day. We consider our assumption of delivery time correlating to capture time to be valid. Although some raptors are known to cache prey during periods of food abundance, deliveries of previously cached prey were few in our study (Giovanni, 2005). This probably was due to environmental conditions in our study area (e.g., heat, direct sun, scavenging by insects), which quickly desiccate and degrade cached prey (Giovanni, 2005). These facts also argue against possible scavenging of roadkills. Several researchers have noted moonlight avoidance by kangaroo rats (Lockard and Owings, 1974; O’Farrell, 1974; Kaufman and Kaufman, 1982). Daley et al. (1992) argued that if phase of the moon has little influence on the predation risk of crepuscular activity but a large influence on the predation risk of nocturnal activity, then reduced nocturnal activity at the

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vol. 52, no. 2

The Southwestern Naturalist

full moon should be at least partially compensated by elevated crepuscular activity. Thus, if crepuscular or diurnal activity occurs, it might coincide with full moon periods. For example, Daley et al. (2000) noted predation of kangaroo rats by the diurnal loggerhead shrike (Lanius ludovicianus; n 5 4) occurred predominantly during the full moon phase. In contrast, we found no pattern of association between predation events and moon phase. We found D. ordii was not uncommon prey among both ferruginous hawks and Swainson’s hawks. Similarly, Olendorff (1993) considered kangaroo rats as an important prey item for ferruginous hawks (6.6%), Bednarz (1988) found they accounted for 2.8% and 6.2% of the prey of Harris’ hawks and Swainson’s hawks, and Steenhof (1998) found they accounted for 4.1% of prairie falcon diets. Although it seems that it is not uncommon for some diurnal raptor species to capture kangaroo rats, it has possibly been overlooked due to the tendency for researchers to focus within the taxonomic group they are studying. The question remains, however, as to why primarily nocturnal kangaroo rats are susceptible to predation by diurnal raptors. We suspect it may be due to local forage conditions. For example, reported activity patterns of D. spectabilis are similar to those of D. ordii: they are almost exclusively nocturnal and avoid foraging on moonlit nights (Lockard and Owings, 1974). However, Lockard (1978) reported D. spectabilis began foraging during the full moon and even in daylight during a period of drought and, hence, seed shortage. Such a situation might have led to D. ordii in our study to become more diurnally active and, therefore, susceptible to diurnal avian predation. Our study area had been under a prolonged period of drought, and range conditions were considered poor, which might have resulted in some kangaroo rats engaging in diurnal foraging. Ultimately, however, an understanding of diurnal activity patterns by kangaroo rats will require more focused examination of their behavior in relation to vegetation and climate conditions. We thank C. Taylor and A. Pruett for helping with various aspects of this study. These data were collected during a study funded by the Texas Parks and Wildlife Department; the U.S. Department of Agriculture Forest Service, Rita Blanca and Kiowa National Grass-

lands; and the U.S. Geological Survey, Texas Cooperative Fish and Wildlife Research Unit. We thank D. Garcia, N. Walls, and the numerous private landowners that allowed us access to hawk nest sites, and R. Chesser for discussion and insights on kangaroo rats. T. Estabrook, N. McIntyre, and 2 anonymous referees provided constructive comments on the manuscript, and C. Ortega provided the Spanish translation of the abstract.

LITERATURE CITED BACON, B. 1994. Red-tailed hawk gets venison lunch. Passenger Pigeon 56:83. BECHARD, M. J., AND J. K. SCHMUTZ. 1995. Ferruginous hawk (Buteo regalis). In: A. Poole and F. Gill, editors. The birds of North America, number 172. Birds of North America, Inc., Philadelphia, Pennsylvania. BEDNARZ, J. C. 1988. A comparative study of the breeding ecology of Harris’ and Swainson’s hawks in southeastern New Mexico. Condor 90:311–323. BLOOM, P. H. 1987. Capturing and handling raptors. In: B. A. Giron Pendleton, B. A. Milsap, K. W. Cline, and D. M. Bird, editors. Raptor management techniques manual. National Wildlife Federation, Washington, D.C. Pages 99–122. BUB, H. 1978. Bird trapping and bird banding, translated by F. Hamerstrom and K. Wuertz-Schaefer (1991). Cornell University Press, Ithaca, New York. BUEHLER, D. A. 2000. Bald eagle (Haliaeetus leucocephalus). In: A. Poole and F. Gill, editors. The birds of North America, number 506. Birds of North America, Inc., Philadelphia, Pennsylvania. CATRON, J. L. E., P. J. POLECHLA, JR., AND R. R. COOK. 2004. Prey of nesting ferruginous hawks in New Mexico. Southwestern Naturalist 49:270–276. CHESSER, R. K. 1979. Opportunistic feeding of mankilled prey by ferruginous hawks. Wilson Bulletin 91:330–331. DALY, M., P. R. BEHRENDS, AND M. I. WILSON. 2000. Activity patterns of kangaroo rats – granivores in a desert habitat. In: S. Halle and N. C. Stenseth, editors. Activity patterns of small mammals: an ecological approach. Springer-Verlag, Berlin, Germany. Pages 145–158. DALY, M., P. R. BEHRENDS, M. I. WILSON, AND L. F. JACOBS. 1992. Behavioural modulation of predation risk: moonlight avoidance and crepuscular compensation in a nocturnal desert rodent, Dipodomys merriami. Animal Behaviour 44:1–9. ENGLAND, A. S., M. J. BECHARD, AND C. S. HOUSTON. 1997. Swainson’s hawk (Buteo swainsonii). In: A. Poole and F. Gill, editors. The birds of North America, number 265. Birds of North America, Inc., Philadelphia, Pennsylvania. GARRISON, T. E., AND T. L. BEST. 1990. Dipodomys ordii. Mammalian Species, number 353.

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GENOWAYS, H. H., AND J. H. BROWN. 1993. Biology of the heteromyidae. American Society of Mammalogists Special Publication 10, Lawrence, Kansas. GIOVANNI, M. D. 2005. Prey partitioning between sympatric grassland raptors. Unpublished M.S. thesis, Texas Tech University, Lubbock. HALLE, S. 2000. Ecological relevance of daily activity patterns. In: S. Halle and N. C. Stenseth, editors. Activity patterns of small mammals: an ecological approach. Springer-Verlag, Berlin, Germany. Pages 67–90. HULL, B., AND P. BLOOM. 2001. The North American bander’s manual for raptor banding techniques. North American Banding Council, Point Reyes Station, California. JORGENSEN, C. D., AND C. L. HAYWARD. 1965. Mammals of the Nevada Test Site. Brigham Young University Science Bulletin, Biological Series 6:1–81. KAUFMAN, D. W., AND G. A. KAUFMAN. 1982. Effect of moonlight on activity and microhabitat use by Ord’s kangaroo rat (Dipodomys ordii). Journal of Mammalogy 63:309–312. KENAGY, G. J. 1976. The periodicity of daily activity and its seasonal changes in free-ranging and captive kangaroo rats. Oecologia 24:105–140. KOCHERT, M. N., K. STEENHOF, C. L. MCINTYRE, AND E. H. CRAIG. 2002. Golden eagle (Aquila chrysaetos). In: A. Poole and F. Gill, editors. The birds of North America, number 684. Birds of North America, Inc., Philadelphia, Pennsylvania. LANGFORD, A. 1983. Pattern of nocturnal activity of male Dipodomys ordii (Heteromyidae). Southwestern Naturalist 28:341–346. LIMA, S. L., AND L. M. DILL. 1990. Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology 68:619– 640. LOCKARD, R. B. 1978. Seasonal change in the activity pattern of Dipodomys spectabilis. Journal of Mammalogy 59:563–568.

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LOCKARD, R. B., AND D. H. OWINGS. 1974. Seasonal variation in moonlight avoidance by bannertail kangaroo rats. Journal of Mammalogy 55:189– 193. MOORE, A. W. 1929. Some notes upon Utah mammals. Journal of Mammalogy 10:259–260. O’FARRELL, M. J. 1974. Seasonal activity patterns of rodents in a sagebrush community. Journal of Mammalogy 55:809–823. OLENDORFF, R. R. 1993. Status, biology, and management of ferruginous hawks: a review. Raptor Research and Technical Assistance Center, Special Report. U.S. Department of the Interior, Bureau of Land Management, Boise, Idaho. PRANTY, B. 2002. Red-shouldered hawk feeds on carrion. Journal of Raptor Research 36:152–153. REICHMAN, O. J., AND M. V. PRICE. 1993. Ecological aspects of heteromyid foraging. In: H. H. Genoways and J. H. Brown, editors. Biology of the heteromyidae. American Society of Mammalogists Special Publication 10, Lawrence, Kansas. Pages 539–574. SHEFFIELD, S. R., AND N. JOBE. 1996. Winter carrion feeding of red-tailed hawks in Oklahoma. Journal of Raptor Research 30:43–44. SNYDER, N. F. R., AND H. A. SNYDER. 1991. Birds of prey: natural history and conservation of North American raptors. Voyageur Press, Inc., Stillwater, Minnesota. SQUIRES, J. R. 1995. Carrion use by northern goshawks. Journal of Raptor Research 29:283. STALMASTER, M. V. 1980. Salmon carrion as a winter food source for red-tailed hawks. Murrelet 61:43–44. STEENHOF, K. 1998. Prairie falcon (Falco mexicanus). In: A. Poole and F. Gill, editors. The birds of North America, number 346. Birds of North America, Inc., Philadelphia, Pennsylvania.

Submitted 12 September 2005. Accepted 22 September 2006. Associate Editor was Philip D. Sudman.

raptor predation on ord's kangaroo rats: evidence for diurnal activity by ...

habits, the quality and resolution of prey use in- formation varied among studies. However, by compil- ing the data, we were able to synthesize a broader understanding of diurnal raptor use of kangaroo rats. RESULTS—We recorded 54 deliveries of D. ordii to ferruginous hawk and Swainson's hawk nests. More of these ...

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