Source: UNIVERSITY OF ILLINOIS submitted to NRP
ECOLOGY OF FOREST SNAKES IN AGRICULTURAL LANDSCAPES
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
0187447
Grant No.
(N/A)
Cumulative Award Amt.
(N/A)
Proposal No.
(N/A)
Multistate No.
(N/A)
Project Start Date
Oct 1, 2000
Project End Date
Sep 30, 2006
Grant Year
(N/A)
Program Code
[(N/A)]- (N/A)
Recipient Organization
UNIVERSITY OF ILLINOIS
2001 S. Lincoln Ave.
URBANA,IL 61801
Performing Department
NATURAL RESOURCES & ENVIRONMENTAL SCIENCES
Non Technical Summary
Habitat loss and fragmentation are the two principal threats to biodiversity. The purpose of this study is to investigate how forest snakes respond to habitat fragmentation and how this affects their ecological role as key nest predators of breeding birds. The second general goal of this research will be to determine the consequences of habitat fragmentation on the genetic structure of snake populations.
Animal Health Component
40%
Research Effort Categories
Basic
60%
Applied
40%
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
13508301070100%
Knowledge Area
135 - Aquatic and Terrestrial Wildlife;

Subject Of Investigation
0830 - Wild animals;

Field Of Science
1070 - Ecology;
Goals / Objectives
I. Habitat Use: I will test the hypotheses that rat snakes use edge habitat, using rat snakes in southern Illinois, where thermal constraints should be much less important to the snakes. In addition to documenting patterns of habitat use, I will test the alternative hypotheses that habitat use patterns of black rat snakes are primarily a consequence of a) thermal demands, b) foraging, or c) predator avoidance. Patterns of habitat use and temperature data for both the snakes and the available habitats will be collected using radio telemetry. Data on prey distribution will be collected by small mammal trapping and by monitoring bird nesting. Habitats will also be sampled to document microhabitat features that might be relevant to snakes for use as refuges and shedding sites. The habitat features and snake locations will be mapped using GPS technology and spatial patterns will be analyzed using GIS. II. Genetic Analyses: A blood sample will be collected from all snakes that are captured. Individual snakes will be permanently identified using passive integrated transponders (PIT tags) implanted subcutaneously. Gravid females will be tracked to nesting sites so that other females using those sites can be captured, and also so that eggs can be collected for incubation. Once eggs hatch the young will be PIT tagged, have a blood sample collected, and then be released at their natural nest site. All snakes sampled will be genetically profiled using a set of six microsatellite markers developed specifically for black rat snakes. These data will be analyzed to determine whether snake populations in isolated woodlots are genetically isolated and if so, whether they show evidence of inbreeding. Paternity analyses will be used to identify the parents of all hatchlings sampled to assess the extent of gene flow via mating. Marked hatchlings later recaptured as juveniles or adults will provide information on the extent of gene flow via natal dispersal.
Project Methods
The work proposed here will require a minimum of five years to complete. The first step will be to identify the woodlots to be used, which will be done in consultation with Dr. Scott Robinson of the University of Illinois. Dr. Robinson's ongoing work on the avian communities in these woodlots will help identify where snakes are abundant and are important nest predators. In the first year of the study, rat snakes will be captured opportunistically, implanted with radio transmitters (Blouin-Demers et al. 2000), and tracked until they enter hibernation. Hibernation sites will be fenced, which will allow many individuals in the population to be captured and sampled as they emerge the following spring (Blouin-Demers et al. 2000). Extensive habitat sampling will be conducted in year one. In year two approximately 25 snakes will be tracked. Particular emphasis will be placed on adult females because their thermal requirements are different when they are gravid (Blouin-Demers and Weatherhead, in press). Thus, how the same females behave when gravid and when not gravid is a powerful tool for determining how thermal demands affect habitat use. Automated telemetry receivers will be installed in year two. These receivers record snake and environmental temperatures 24 h/day, and thus provide far more comprehensive information than non automated approaches (e.g., Brown and Weatherhead 2000). Females will be tracked to nest sites and eggs will be collected for incubation beginning in year two. Intensive radio telemetry will continue through years three and four. If numbers warrant, parallel data collection on additional snake species will begin in year four as the rat snake telemetry is being completed. Genetic analysis will also begin in years three and four. Collection of eggs for incubation will continue in years three and four. In year five the remaining transmitters will be removed from black rat snakes, and the research emphasis in the field will switch to other snake species. However, monitoring of spring emergence from hibernacula to capture rat snakes originally marked as hatchlings will continue in year five and beyond.

Progress 10/01/00 to 09/30/06

Outputs
The central goal of this study was to test the hypothesis that snakes select habitat to meet thermoregulatory needs, so that habitat use will vary seasonally, by sex, and by latitude. Understanding this relationship helps understand how snakes are affected by habitat fragmentation in agricultural landscapes, and in turn how their use of habitat affects their broader ecology (e.g., predator-prey relationships). This hypothesis was tested by comparing habitat use by ratsnakes and racers in Illinois. Illinois ratsnakes used upland forest more and forest edges less than ratsnakes at their northern range limit, although female ratsnakes in Illinois used edges less than males, regardless of their reproductive status. As predicted, relative to Illinois ratsnakes, racers preferred forest edges and avoided forest interior. Also, female racers used edge more than males, especially while gravid. Most seasonal differences in habitat use among and between species were consistent with the thermal ecology hypothesis. Thus, both latitudinal and species differences in habitat use were broadly consistent with differences in thermal ecologies. Forest fragmentation is likely to be detrimental to ratsnakes in Illinois but beneficial to racers. Separate from this work, a collaborative study on the genetic structure of northern ratsnake populations was completed. The thermal ecology study will form the basis for Ph.D. student Gerardo Carfagno's dissertation (currently being written). This work has produced three publications thus far with several more in preparation

Impacts
Agricultural landscapes are characterized by both decreases in the area of original habitat and fragmentation of the original habitat that remains. Preserving biodiversity in natural habitats within agricultural ecosystems requires an understanding of how individual species respond to habitat reduction and fragmentation and how those responses affect their interactions with other species. There is abundant evidence documenting the decline of forest birds in fragmented forests, and increased nest predation is often implicated as a proximate cause of the decline. Snakes are the most important avian nest predators in forests in the Midwest, and the snakes' preference for edge habitats may increase their impact on birds nesting in fragmented forests. By determining how black rat snakes respond to habitat fragmentation and how this affects nest predation of breeding birds, it will be possible to assess the impact of potential habitat management practices on the abundance and distribution of snakes, and thus on nesting birds.

Publications

  • Carfagno, G.L.F., Heske E.J. and Weatherhead P.J. 2006. Does mammalian prey abundance explain forest-edge use by snakes? Ecoscience 13:293-297.
  • Gibbs, H.L., Coreyl, S.J., Blouin-Demers, G., Prior, K.A. and Weatherhead, P.J. 2006. Hybridization between mtDNA-defined phylogeographic lineages of ratsnakes (Pantherophis sp.). Mol. Ecol. 15: 3755-3767.
  • Carfagno, G.L.F. and Weatherhead, P.J. 2007. Intraspecific and interspecific variation in use of forest-edge habitat by snakes. Can. J. Zool. (In Press).


Progress 01/01/05 to 12/31/05

Outputs
The fieldwork begun three years ago in the Cache River State Natural Area in southern Illinois is being completed this year, led by Ph.D student Gerardo Carfagno. The goal this year was to complete tracking of black ratsnakes and blue racers that still carried transmitters, with particular emphasis on fine-scale activity patterns. Toward the end of the active season, snakes were captured and brought back to campus so transmitters could be removed. As transmitters were removed, information on the snakes' health was recorded for use in a publication on health effects of transmitters. At present, only 4 of the initial 14 snakes remain to be recaptured for transmitter removal. These snakes will be captured by the end of October. Fine-scale tracking was necessary to assess the frequency with which snakes moved between habitats, and thus encountered different microclimatic conditions. Ratsnakes proved to be very sedentary whereas racers make numerous movements over the course of a day. These data will be used when quantifying patterns of thermoregulation. With the completion of fieldwork, attention will be focused on data analysis and writing.

Impacts
Agricultural landscapes are characterized by both decreases in the area of original habitat and fragmentation of the original habitat that remains. Preserving biodiversity in natural habitats within agricultural ecosystems requires an understanding of how individual species respond to habitat reduction and fragmentation and how those responses affect their interactions with other species. There is abundant evidence documenting the decline of forest birds in fragmented forests, and increased nest predation is often implicated as a proximate cause of the decline. Snakes are the most important avian nest predators in forests in the Midwest, and the snakes' preference for edge habitats may increase their impact on birds nesting in fragmented forests. By determining how black rat snakes respond to habitat fragmentation and how this affects nest predation of breeding birds, it will be possible to assess the impact of potential habitat management practices on the abundance and distribution of snakes, and thus on nesting birds.

Publications

  • Carfagno, G.L.F., Heske E.J. and Weatherhead P.J. 2006. Does mammalian prey abundance explain forest-edge use by snakes? Ecoscience (In Press).


Progress 01/01/04 to 12/30/04

Outputs
The fieldwork begun two years ago in the Cache River State Natural Area in southern Illinois was continued this year, led by the Ph.D student Gerardo Carfagno. The field work will be completed this fall. The goals this year were to continue tracking black ratsnakes and blue racers beginning when they emerged from hibernation to obtain data on snake body temperatures and to continue quantifying the habitat used by the snakes. Substantial progress toward meeting all these goals was realized. We captured 26 new racers and 32 new ratsnakes. Transmitters were implanted into 6 ratsnakes and 4 racers (in addition to the 14 individuals implanted in 2003). Snakes were located every 48 hours through the spring and summer, and twice a week beginning at the end of September. Six snakes were lost during the summer, two due to transmitter failure and the rest to predation. The remaining snakes will be tracked until they go underground, which occurs between October and mid-November. Tracking in 2004 yielded approximately 700 new snake locations and approximately 60,000 body temperature measurements. Habitat variables were quantified at every second new snake location (approximately 300 thus far), and at randomly selected locations (approximately 150 thus far). Two female ratsnakes and two female racers were captured while gravid and laid eggs in the lab. Those eggs were incubated and blood samples collected before releasing the hatchlings. Overall, therefore, substantial progress was made on all major objectives.

Impacts
Agricultural landscapes are characterized by both decreases in the area of original habitat and fragmentation of the original habitat that remains. Preserving biodiversity in natural habitats within agricultural ecosystems requires an understanding of how individual species respond to habitat reduction and fragmentation and how those responses affect their interactions with other species. There is abundant evidence documenting the decline of forest birds in fragmented forests, and increased nest predation is often implicated as a proximate cause of the decline. Snakes are the most important avian nest predators in forests in the Midwest, and the snakes' preference for edge habitats may increase their impact on birds nesting in fragmented forests. By determining how black rat snakes respond to habitat fragmentation and how this affects nest predation of breeding birds, it will be possible to assess the impact of potential habitat management practices on the abundance and distribution of snakes, and thus on nesting birds.

Publications

  • Weatherhead, P.J. and Blouin-Demers, G. 2004. Understanding avian nest predation: Why ornithologists should study snakes. J. Avian Biol. 35:1-6.


Progress 01/01/03 to 12/31/03

Outputs
The fieldwork begun last year in the Cache River State Natural Area in southern Illinois was continued this year, led by the Ph.D student Gerardo Carfagno. The goals this year were to capture black ratsnakes and blue racers as they emerged from hibernation sites located last fall, implant a subset of these with radio transmitters, radio track the snakes through the active season, obtain data on snake body temperatures, and to continue quantifying the habitat used by the snakes. Substantial progress toward meeting all these goals was realized. We captured 99 racers and 54 ratsnakes. Transmitters were implanted into 7 ratsnakes (in addition to the 11 individuals implanted in 2002) and 12 racers. Snakes were located every 48 hours through the spring and summer, and twice a week beginning at the end of September. Eight snakes were lost during the summer, two due to transmitter failure and the rest to predation. The remaining snakes were tracked until they went underground, which occurred between October and mid-November. Tracking in 2003 yielded approximately 950 snake locations and approximately 75,000 body temperature measurements. Habitat variables were quantified at every second new snake location (approximately 420 thus far), and at randomly selected locations (approximately 200 thus far). Two female ratsnakes and three female racers were captured while gravid and laid eggs in the lab. Those eggs were incubated and blood samples collected before releasing the hatchlings. Overall, therefore, substantial progress was made on all major objectives.

Impacts
Agricultural landscapes are characterized by both decreases in the area of original habitat and fragmentation of the original habitat that remains. Preserving biodiversity in natural habitats within agricultural ecosystems requires an understanding of how individual species respond to habitat reduction and fragmentation and how those responses affect their interactions with other species. There is abundant evidence documenting the decline of forest birds in fragmented forests, and increased nest predation is often implicated as a proximate cause of the decline. Snakes are the most important avian nest predators in forests in the Midwest, and the snakes' preference for edge habitats may increase their impact on birds nesting in fragmented forests. By determining how black rat snakes respond to habitat fragmentation and how this affects nest predation of breeding birds, it will be possible to assess the impact of potential habitat management practices on the abundance and distribution of snakes, and thus on nesting birds.

Publications

  • No publications reported this period


Progress 01/01/02 to 12/31/02

Outputs
Last year the Cache River Natural Area in southern Illinois was selected as the study site for this work, and Gerardo Carfagno was recruited as the Ph.D. student to lead the field work. Data collection for this project was initiated in 2002. The goals this year were to capture black rat snakes and implant radio transmitters in them, radio track the snakes through the active season, and to begin quantifying the habitat used by the snakes. Substantial progress toward meeting all these goals was realized. Eleven snakes were captured that were large enough to carry a transmitter. Transmitters were surgically implanted in each of these snakes between May 15th and September 2nd. Snakes were located every 48 hours through the summer, and twice a week beginning in September. Two snakes were lost during the summer, either due to transmitter failure or because the snakes were taken by predators and removed from the study area. A third snake suffered from a chronic infection following surgery, which eventually required that the transmitter be removed. The remaining 8 individuals will be tracked until they go underground to hibernate, which should occur in October. Tracking has yielded approximately 350 snake locations thus far. Habitat variables have been quantified at every second new snake location (approximately 80 thus far), and at randomly selected locations (approximately 40 thus far). Four females were captured while gravid and laid eggs in the lab. Those eggs were incubated and blood samples collected before releasing the hatchlings.

Impacts
Agricultural landscapes are characterized by both decreases in the area of original habitat and fragmentation of the original habitat that remains. Preserving biodiversity in natural habitats within agricultural ecosystems requires an understanding of how individual species respond to habitat reduction and fragmentation and how those responses affect their interactions with other species. There is abundant evidence documenting the decline of forest birds in fragmented forests, and increased nest predation is often implicated as a proximate cause of the decline. Black rat snakes may be the most important species of avian nest predator in forests in the Midwest, and the snakes' preference for edge habitats may increase their impact on birds nesting in fragmented forests. By determining how black rat snakes respond to habitat fragmentation and how this affects nest predation of breeding birds, it will be possible to assess the impact of potential habitat management practices on the abundance and distribution of black rat snakes, and thus on nesting birds.

Publications

  • No publications reported this period


Progress 01/01/01 to 12/31/01

Outputs
The study site for this project has been selected. After visiting a number of potential sites in southern Illinois, the Cache River drainage was chosen. Past research on nesting birds at this site confirms that rat snakes are abundant and an important nest predator. In addition, the habitat at the site is a mixture of mature upland and bottomland forest, interspersed with both natural and man-made edges. The man-made edges are the boundaries between agricultural fields and forest. Forest restoration is underway in some of those fields. As a consequence, the study site contains a rich variety of the habitat types required for this study. A potential field station at the site has been identified and steps are being taken to make that facility available to support this research. A PhD student (Geraldo Carfagno) has been recruited to undertake this research. He is currently in his first semester at U of I. However, he was given the opportunity to participate in an ongoing snake study last summer, which provided him with valuable experience relevant to the work he will be undertaking. Beginning in March 2002, snakes will be captured at the study site and transmitters will be implanted. At that point data collection will begin.

Impacts
The results from this study will be used to develop guidelines for the management of forests that exist in agricultural landscapes. By knowing how the structure of the boundary between forests and agricultural fields affects the ecological relationship between birds that nest in the forest and their snake predators, potential ways of influencing that relationship can be identified.

Publications

  • No publications reported this period