Hair-snare survey to assess distribution of margay (Leopardus wiedii) inhabiting El Cielo Biosphere Reserve, Tamaulipas, Mexico, PJ Downey

Tags: Mexico, El Cielo Biosphere Reserve, cloud forest, gray fox, National Wildlife Federation Science, Bobcat Research Conference, gray foxes, Journal of Wildlife Management, Chamberlain, margay, station, Margay Research Expeditions, stations, ocelot, John Weaver, Oklahoma State University, margays, Wildlife Conservation Society, hair samples, Norberto Castro, Cleveland Zoological Society, deciduous forest, SAS Institute Inc., Wildlife Genetics International Inc., the Faculty of the Graduate College, statistical analysis, Ronald A. Van Den Bussche Dr, Mike Chamberlain, Cleveland Metroparks Zoo, M. Chamberlain, J. Weaver, tropical deciduous forest, Schmidly, relative abundance, GDFR, Wildlife Genetics International, direction, Ronald A. Van Den Bussche, Texas, F. Sunquist, University of Chicago, North America, National Wildlife Federation, Tamaulipas Mexico, Tamaulipas, Mexico, SAS Institute, Wildlife Conservation, Pie Plate, endangered species, bobcat, Oklahoma City Zoological Society, Graduate College, Dr. John Weaver, LEOPARDUS WIEDII, A. Gordon Emslie Dean, Eric C. York, Eric C. Hellgren Thesis Adviser, Oklahoma City Zoo, Eric C. Hellgren, James H. Shaw
Content: HAIR-SNARE SURVEY TO ASSESS DISTRIBUTION OF MARGAY (LEOPARDUS WIEDII) INHABITING EL CIELO BIOSPHERE RESERVE, TAMAULIPAS, MEXICO By PATRICIA JEAN DOWNEY Bachelor of Science in Geology Oklahoma State University Stillwater, Oklahoma 1994 Submitted to the Faculty of the Graduate College of the Oklahoma State University in partial fulfillment of the requirements for the Degree of MASTER OF SCIENCE December, 2005
HAIR-SNARE SURVEY TO ASSESS DISTRIBUTION OF MARGAY (LEOPARDUS WIEDII) INHABITING EL CIELO BIOSPHERE RESERVE, TAMAULIPAS, MEXICO Thesis Approved: Dr. Eric C. Hellgren Thesis Adviser Dr. James H. Shaw Dr. Ronald A. Van Den Bussche Dr. A. Gordon Emslie Dean of the Graduate College ii
ACKNOWLEDGEMENTS I would like to take this opportunity to express my gratitude to the countless number of people who assisted me in countless ways through my graduate work. I wish to express my sincere appreciation to my major advisor, Dr. Eric C. Hellgren for his advice, guidance and encouragement throughout my studies. A special thanks to Dr. James H. Shaw for not only being a member of my committee, but believing in me enough to let me enter graduate school. My appreciation extends to Dr. Ronald A. Van Den Bussche for his membership on my committee. A thanks to Dr. Larry Claypool for his patient explanations and advice on statistical analysis. My deepest respect and eternal gratitude to one Norberto Castro, Executive Director of the Oklahoma City Zoo, whose unwavering support led this project to completion. I am forever indebted to the Oklahoma City Zoological Park and Botanical Gardens and the Oklahoma City Zoological Society for their financial support of the Margay Research Expeditions and my thesis. A sincere thank you to Bill Savage, Curator of Mammals, who let me change my work schedule as needed to attend classes and spend time in Mexico. My appreciation extends to Dwight Scott and all curators who allowed members of their staff to join the project. A special thanks to all keepers who gave their time and energy as co-leaders of expeditions. My gratitude to the pachyderm staff for their patience with my intermittent absence from the area. My appreciation to Craig Harmon for help and laughs with the ocelot lure trials and Colleen Kovach for iii
hours of data entry. I thank Rhondi Large of WildCare for sharing her gray fox knowledge and access to her animals. I wish to thank the Cleveland Metroparks Zoo and Cleveland Zoological Society's Small Grants Program for funding the genetic analysis of this research. I would like to thank my parents for their financial and Emotional Support through this process. Their help ranged from financing entire trips to Mexico, to worriedly watching the weather channel while I was gone and everything in between. Although my father cannot be here to see the completion of this project, the work is dedicated to him. My appreciation extends to Arturo Caso for his guidance and support. It is an honor to have learned felid ecology and field techniques from such an extraordinary scientist, teacher and friend. I wish to thank Sasha Carvajal for her dependable help with the field work and her friendship. I would like to thank all volunteers who participated in the Margay Research Expeditions. In particular I would like to thank Fred Hanson who joined us on all expeditions. A big thank you to Charlie and Lana Vieh for providing dependable and safe transportation in Mexico and many, many smiles. My gratitude to Nik Gour who has become a wonderful friend and fellow margay researcher. I wish to thank Dr. David Paetkau and Jennifer Weldon of Wildlife Genetics International Inc. for genetic analysis and advice concerning permit issues. My respect and appreciation extends to Dr. John Weaver for his advice and encouragement on this project. I am grateful to Dr. Richard Harrison, Dr. Paul Beier, Dr. Kerri Frangioso, Dr. Lisa Haynes and John Childs not only for their willingness to share their information, but iv
allowing me to cite their studies. My gratitude to Dr. Mike Chamberlain for sharing his knowledge of gray fox. v
Table of Contents Page I. INTRODUCTION ......................................................................................................1 Study Area ...............................................................................................................7 II. METHODS..............................................................................................................10 Hair-snare survey...................................................................................................10 Vegetation sampling ..............................................................................................11 Species testing........................................................................................................12 data analysis ..........................................................................................................12 III. RESULTS ..............................................................................................................14 IV. CONCLUSION......................................................................................................15 Management implications......................................................................................18 V. REFERENCES......................................................................................................21 vi
LIST OF TABLES
Table
Page
1. Microhabitat characteristics of hair-snare sites
on El Cielo Biosphere Reserve.........................................................27
2. Summary of results from published and unpublished
hair-snare studies targeting felids ....................................................28
vii
LIST OF FIGURES
Figure
Page
1. Geographic range of Leopardus wiedii...................................................29
2. Diagram depicting layout of hair-snare station
and vegetation plots.....................................................................31
3. Frequency of gray fox hits by season in each habitat
at El Cielo Biosphere Reserve, Tamaulipas, Mexico...............................33
4. Felid hair-snare study sites plotted against gray fox
(Urocyon cinereoargenteus) range (IUCN/SSC 2004).............................35
5. North American range of coyote (Canis latrans; IUCN/SSC 2004)................37
6. North American range of bobcat (Lynx rufus; Cat Survival Trust 1996)...........39
viii
INTRODUCTION The Felidae consists of 37 species evolved from a common ancestor 12 to 2 million years ago. Felids recognized today are a result of 4 radiations from this ancestor (Collier and O'Brien 1985). The first radiation occurred 12 million years ago and led to the ocelot lineage consisting of the ocelot (Leopardus pardalis), margay (L. wiedii), tigrina (L. tigrinis), Geoffrey's cat (Oncifelis geoffroyi), kodkod (O. guigna), pampas cat (O. concolor) and Andean mountain cat (Oreailurus jacobita). The second branch occurred 8­10 million years ago and includes close relatives of the domestic cat such as the jungle cat (Felis chaus), sand cat (F. margarita), and Pallas cat (Otocolobus maul). The third branch radiated several times 4­6 million years ago giving rise to the pantherine lineage, e.g., jaguar (Panthera onca), lion (P. leo), leopard (P. pardus): and again 2 million years ago leading to a split between the pantherines and the lynx (Lynx canadensis: Collier and O'Brien 1985). The Neotropical Region has been inhabited by 10 extant species of felids since the Pleistocene, 1­5 million years ago (Eizirik et al. 1998). These species appeared in South America 2-5 million years ago during the formation of the Panamanian Isthmus (Johnson et al. 1998). Seven of the 10 species comprise the ocelot lineage (Johnson et al. 1998), of which the ocelot and margay are considered sister species (Eizirik et al. 1998). These species are considered sympatric throughout most of their range from the Southern United States (ocelot) or northern Mexico (margay) to South America (Eizirik et al. 1998 1
Today, most felids are listed as threatened or endangered by the Convention on International Trade of endangered species (CITES 2001: Collier and O'Brien 1985). Margays were listed on Appendix 1 of CITES in 1989 (Nowell and Jackson 1996) and are classified as insufficiently known by the IUCN (de Oliveira 1998). The margay was one of the most heavily exploited Latin American cats in the 1970s and 1980s. Demand for pelts increased and coincided with exploitation of other spotted cats, namely the ocelot and jaguar. An average annual trade of 14,000 margay pelts was reported to CITES from 1976 to 1984. Hunting is prohibited throughout most of its range with the exception of Ecuador, Guyana and El Salvador (Nowell and Jackson 1996). Today, international trade has virtually ceased, but margay numbers are still in decline due to habitat destruction (Sunquist and Sunquist 2002). Schinz named the margay Felis wiedii in 1821 in honor of the Prince of Wied who discovered this felid while on expedition to South America (Guggisberg 1975). The genus name has since been changed to Leopardus (Pocock 1941). Before 1852, the margay held status as a member of the fauna of the United States as verified by 1 specimen recorded at Eagle Pass, TX. This individual is believed to have been a vagrant and is the sole record of the subspecies L. w. cooperi (Tewes and Schmidly 1987). In Mexico, the margay occurs along both coasts from Tamaulipas in the east to Sinaloa in the west. Their range then extends southward to the Isthmus of Tehuantepec and finally through the state of Chiapas and the Yucatan Peninsula. This species inhabits all of Central America and the South American countries of Colombia, Venezuela, Guyana, Brazil, Uruguay, Argentina, Paraguay, Bolivia, Peru and Ecuador (Tewes and Schmidly 1987). 2
Today, 10 subspecies of margay are recognized (de Oliveira 1998). All subspecies have been distinguished on the basis of coat color and pattern and/or skull measurements (Pocock 1941). The names and ranges of the subspecies (Figure 1) are as follows: L.w. amazonica - Amazon, Brazil; L.w. boliviae ­ Brazil, Bolivia; L.w. cooperi ­ Nuevo Leon; L.w. glauculus - Sinaloa through north Oaxaca; L.w. nicaraguae - Honduras to Costa Rica; L.w. oaxacensis -Tamaulipas to Oazaca; L.w. pirrensis - Panama to northern Peru; L.w. salvinia ­ Chiapas, Guatemala, El Salvador; L.w. vigens - Orinoco to the Amazon basin and L.w. yucatanicus Chiapas to northern Guatemala and Yucatan (de Oliveira 1998). The margay closely resembles and is often mistaken for the ocelot. In some areas of South America the margay is even referred to as "little ocelot" (Guggisberg 1975). Although not identical, the spotting pattern of their coats is similar and both felids share a darker head, tail and back and white belly. Despite the similarities in coat pattern, the margay is smaller and more slightly built than the ocelot, and proportionately a longer tail and back legs (Guggisberg 1975). Margays are arboreal and physically well-designed to exploit this ecological niche. The most prominent adaptive feature is a highly flexible ankle joint that can supinate through 180° (Alderton 1993). This feature allows the margay to travel head first down a tree trunk and the ability to grasp a branch equally well with fore or hind paws (Tewes and Schmidly 1987). Margays also have been known to move from limb to limb in forest canopies often suspending by one leg. Other adaptations to an arboreal lifestyle include broad feet with long claws, highly mobile toes for gripping and stability 3
(Sunquist and Sunquist 2000), and a proportionately longer tail for balance (Tewes and Schmidly 1987). The margay is strongly associated with wooded habitats, primarily tropical evergreen lowland forests and cloud forest. Its arboreal nature may render it more vulnerable to the effects of deforestation than other felids such as the ocelot (Tewes and Schmidly 1987). However, sightings of margays in cocoa and coffee plantations in South America suggest it can be adaptable if necessary (Mondolfi 1982). Knowledge of margay ecology is extremely limited because little formal in-situ research has been conducted in any part of its range. To date, only 1 published study has collected data on the home range and activity patterns of margay (Konency 1989). This information was gathered for 6 months in Belize in 1985-1986 on a young radio-collared male. The home range of this individual was found to be 10.95 km2. A second male was captured but not radio-collared. Results of this study showed the margay to display a strongly nocturnal activity pattern with peak travel rates from 0100 to 0300 and lowest rates at the 1200 hr. This activity pattern was supported by observations of captive margays. Traveling occurred on the ground and in the forest canopy. During daylight hours this male rested in trees 7-10 m above ground. Ambient moonlight did not appear to affect movement rates (Konency 1989). A second in-situ study conducted from 1990 to 1994 (Carrillo et al. 2000) used track records and arboreal sightings to compare the abundance of mammals, including margays, in Corcovado National Park (CNP) and Golfo Dulce Forest Reserve (GDFR). Both areas are located in Costa Rica, have similar environmental characteristics, but have different hunting regulations. Hunting is prohibited in CNP, but allowed in GDFR. 4
Results of this study showed the abundance of margay on GDFR to be less than CNP and that margay abundance actually increased on CNP during the study (Carrillo et al. 2000). To date, little is known of wild margay reproduction but some information has been derived from captive studies. Females display estrous cycles of approximately 32 days with periods of heat lasting 7­10 days. In captivity, gestation has been measured at 81 days, whereas in the wild it is estimated to be 70 days. Females in captivity give birth to one or 2 kittens per litter (Tewes and Schmidly 1987). It is often difficult to obtain an accurate measure of population size of most carnivores due to their secretive nature and low abundance (Sargeant et al. 1998). However, an estimate of relative abundance is often a useful substitution. The scentstation technique is one such method that allows wildlife biologists to estimate carnivore abundance and monitor trends in these populations (Chamberlain et al. 1999). Originally developed by J. E. Wood in 1959 to determine abundance of gray fox (Urocyon cinereoargenteus) and red fox (Vulpes vulpes) in southeastern United States (Hon 1979), this technique has since been applied to coyote (Canis latrans; Roughton and Sweeney 1982), lynx (McDaniel et al. 2000, Weaver 2002), bobcat (Lynx rufus; Diefenbach et al. 1994), river otter (Lutra canadensis), mink (Mustela vison; Conner et al. 1983) and multiple species surveys (Chamberlain et al. 1999). Recent studies have indicated the scent-station technique is best suited for monitoring trends or fluctuations in carnivore populations over time as opposed to relative abundance as a point in time (Chamberlain et al. 1999). However, limitations to this method exist. For example, the scent-station technique is ill-suited for localized monitoring of wide-ranging carnivores and species that are rarely detected (Sargeant et 5
al. 1998). Recommendations to refine the scent-survey method include conducting multiple surveys each year, having surveys contain as many stations as possible, and placing stations as far apart as possible (Diefenbach et al. 1994). The design of the traditional scent-station is a 1.0-m diameter circle cleared of vegetation and sifted with fine soil to permit track identification. The desired attractant is presented in the center of the circle by a variety of means including drops applied to a clump of grass (Hon 1979), a saturated cotton ball (Brady 1979; Conner 1983), encapsulated (Knowlton and Tzilkowski 1979) or on a plaster of Paris disc (Chamberlain et al. 1999). Scent-stations are typically established at equal intervals along a transect and checked daily for visitation. Harrison (1997) examined the efficiency of scent stations of Central American felids. In this study, captive margays showed highest investigation times with bobcat urine. Scent-station visitation by wild felids including margays was too infrequent to test specific attractants for individual species (Harrison 1997). The hair-snare station differs from the scent-station in that it depends on microscopic or genetic analysis of hair left on a snare to determine species. Weaver expanded the idea and designed a new methodology to detect lynx (Weaver 2002). Lure is placed on a carpet pad between 2 rows of nails. The pad is nailed to a tree at cheek height to the lynx. A lure, Cat CallTM, was formulated to elicit face-rubbing behavior in lynx. A pie plate was hung near the lure site as a visual attractant. In 1996 the protocol was tested with over 100 test snares in the Kootenai forest of northwest Montana. A total of 39 hits were recorded during the trial with lynx detected on 28 snares (Turbak 1998). 6
The objectives of this study were to use hair-snare stations (1) to assess seasonal habitat use of margay and (2) to determine which microhabitat variables are preferred by margays. I predicted a significantly higher wet-season abundance of margays in the cloud forest based on margay habitat and diet preferences. Cloud forest is reported to be the preferred habitat of margays (Tewes and Schmidly 1987) and rodents comprise much of their diet (Konecny 1989). Rodents have been documented to have higher wet-season abundances in cloud forest in the Sierra de Manantlan Biosphere Reserve, western Mexico (Vazquez 2000). Also, as margays are small, arboreal felids, I predicted selection for station sites with higher canopy cover and understory cover. STUDY AREA El Cielo Biosphere Reserve is located approximately 400 km south of Brownsville, TX, in the southwestern corner of the state of Tamaulipas, Mexico. Situated on the Tropic of Cancer, the 1445.3-km2 Reserve occurs between 22 55'30" and 23 25'50" N and 99 05'50" and 99 26'30"W. This area is dominated by the north-south trending mountain ranges of the Sierra de Guatemala, a region of Sierra Madre. The topography is mostly Cretaceous karstic limestone with most slopes exceeding 20% (Peterson 2001). El Cielo claims several distinct climate and vegetation types as a result of the topography and altitudinal gradient, which ranges from 200 to 2200 m. The lowest altitude (200-800 m) is a subtropical deciduous climate zone characterized by a semiwarm humid climate with rains from May to October. natural vegetation is predominately tropical-deciduous and semi-evergreen forests. The canopy is 11-15 m high, and includes a dense understory of trees and shrubs. The most common tree species 7
include Bursera simaruba, Brosimum alicastrum, Enterolobium cyclocarpum, Pseudobombax ellipticum, Phoebe tampicensis, Cedrela mexicana and Savia sessiliflora (Gram and Faaborg 1999). The next highest climate zone (800-1400 m) is humid-temperate. Mean annual rainfall is 2,000-2,500 mm with year-round rain. This habitat, known as montane mesophyll forest or "cloud forest", claims El Cielo as its northernmost extension in the Americas (Jones 2000). Canopy height is 20­30 m and epiphytes are common (Gram and Faaborg 1999). The dominant tree species in this habitat are Quercus sartotii, Q. germana, Liquidamber styraciflua, Acer skutchii, Magnolia tamaulipana, Fagus mexicana and Podocarpus reichei. (Gram and Faaborg 1999). Higher elevations (1400-1800 m) are characterized by a temperate sub-humid climate with oak and oak-pine forest predominating. This habitat differs from cloud forest in that it contains fewer understory trees, tall shrubs and epiphytes. Mean canopy height is 20 m. The dominate tree species in this habitat are Quercus affinis, Q. mexicana, Pinus montezumae, P. patula, Liquidambar styraciflua and Acer skutchii. (Gram and Faaborg 1999). It was my observation that oak-pine forest occurred at a lower elevation than specified by Gram and Faaborg (1999). The frequency of oak and pine trees increased, epiphytes became less common, and understory density decreased at approximately 1200 m in elevation. For the purposes of this research, cloud forest will be designated from 800 to 1200 m and oak-pine and pine forest will be designated from 1200 to 1800 m. The climate of El Cielo depends largely upon altitude. The lowlands are typically humid and hot whereas the highlands are comparatively cooler. El Cielo is considered to 8
have 2 seasons, a rainy season and a dry season. The rainy season begins in May or June and lasts through October. The dry season continues from November through April (Peterson, 2001) 9
METHODS HAIR-SNARE SURVEY I conducted 8 hair-snare surveys for margays during the course of the study. Wet- season (May­October) surveys were conducted in September 2003 and May, August and October 2004. Dry-season (November­April) surveys were conducted in November 2002, April and December 2003, and March 2004. Hair-snare stations were established along a transect represented by the 15-km road leading from the village of San Jose at 1400 m in altitude to the north end of Gomez Farias at 200 m. This road traversed all 3 habitat types surveyed in this research -- oakpine forest, cloud forest and subtropical-deciduous forest. Habitats were sampled in a manner equivalent to the relative occurrence along the transect. Approximately 30 hairsnare stations were established at 500-m intervals and alternated between the left-hand and right-hand sides of the road. Each station was placed 50 m perpendicular to the transect. At each station altimeter, northing and easting readings were taken with a Magellan GPS unit. Stations consisted of an 8 X 8 cm carpet pad studded with 2 rows of 2.2-cm nails through the back of the pad. Pads were nailed to trees approximately 0.61 m above ground with the nail rows oriented vertically. This design allowed the lure pad to act as a hair snag. Lure pasted with catnip (Nepetu cataria) was placed between the rows of nails. 10
This method was modeled after the technique employed by the Wildlife Conservation Society (WCS) to survey Canadian lynx in Montana (Weaver 2002). Lure (Weaver's Cat CallTM ) was purchased from John Weaver, a carnivore biologist for the Wildlife Conservation Society. Weaver has tested the lure on many species of felids in captivity including margays. In all species tested, the lure successfully elicited the natural face marking behavior of felids (J. Weaver, personal communication). Surveys were conducted for 8 days. At the end of each survey, all stations were examined for the presence of hair. If hair was present, the lure pad including all nails was sealed in a plastic sandwich bag. Date and station number was recorded twice for each pad, once on the outside of the bag and again on a small piece of paper placed inside the bag. All lure pads were bagged individually. Lure pads not containing hair were thrown away. VEGETATION SAMPLING For each station, I sampled understory cover by the Daubenmire method (Bonham 1989) in a 1-m2 plot at each lure site and in 1-m2 plots 10 m from a lure site in northeast (45°), southeast (135°), southwest (225°) and northwest (315°) directions (Figure 2). Data included cover of forbs, grasses, bare ground, rock, and leaf litter. To sample over story cover, canopy cover measurements were averaged from the 5 plots at each station using a spherical densiometer (Bonham 1989). A margay model was used to measure visual obstruction. The model consisted of white fabric with large black spots cut to the dimensions of a margay profile. The cloth margay was then adhered to the base of a 0.91-m x 1.22-m black cloth. One person stayed seated at the lure site while a second walked backwards in each of the northeast 11
(45°), southeast (135°), southwest (225°) and northwest (315°) directions holding the model at ground height. The point where the margay model was no longer visible from the lure station was determined for that direction. In the event only one person collected the data, the model was nailed at the lure site facing the 4 direction at ground height. The collector walked backwards in each direction. The point where the model could not be seen from a seated position was recorded for that direction. The 4 visual obstruction measurements were averaged at each site. This average was considered the index for understory density. SPECIES TESTING The Wildlife Genetics International (WGI) lab in Nelson, BC, performed all genetic analysis. Samples of blood drawn from margays in ECBR (Carvajal 2005) also were analyzed for reference. Samples were extracted using QIAGEN's DNeasy Tissue kit following the manufacturer's instructions. WGI did not exhaust all of the sample material nor was all the DNA extracted. Leftover sample and DNA will be archived for five years. WGI determined the species of hair samples using a sequence-based analysis of the 16s rRNA mitochondrial gene (Kocher et al. 1989, Johnson and O'Brien 1997). Although specific primers and conditions are proprietary, results can be fully reproduced following procedures in Johnson and O'Brien (1997). DATA ANALYSIS A Chi-square test using the categorical data modeling procedure (PROC CATMOD) of SAS (SAS Institute Inc. 1990) was applied to test the relationship among the frequency of success and failure at hair-snare sites by habitat and season. Success was measured as hit or miss; habitat was measured as oak-pine forest, cloud forest or sub- 12
tropical deciduous forest; and season was wet or dry. I conducted stepwise logistic regression with backward elimination (PROC LOGISTIC; SAS Institute, Inc. 1997) to evaluate the use of vegetation variables (e.g., understory cover and density, overstory cover) in predicting the presence (hit) or absence (miss) of margays at individual hairsnare sites. An Q = 0.05 was chosen as the level of significance 13
RESULTS A total of 117 hits were recorded during the 8 surveys. Species analysis did not detect margay on any pad. The largest proportion of hits (44.4%; 52/117) was gray fox. Other species identified from the hair samples included dog (Canis familiaris), domestic cat (Felis domesticus), goat (Capra hircus), long-tailed weasel (Mustela frenata ) and horse (Equus caballus). A total of 23 samples failed to produce results after 2 attempts at species identification, 8 samples were of unknown identity (7 were the same identity), and 3 samples were not extracted due to lack of material. Due to the lack of margay data, statistical analysis was conducted using only gray fox data. The frequency of hits by gray fox varied by habitat*season interaction (T2 = 5.90, P = 0.052). Success was uniformly high acRoss Seasons in the oak-pine forest, higher in the cloud forest during the dry season but higher in the wet season in the subtropical-deciduous forest (Figure 3). The frequency of fox successes also varied by habitat (T2 = 8.64, P = 0.013). Hits occurred most frequently in oak-pine forest, followed by cloud forest and then sub-tropical­deciduous forest. Logistic regression analysis indicated no microhabitat variable or combination of variables reliably predicted success or failure of a hair-snare station (Table 1). 14
CONCLUSION The hair-snare technique failed to detect margay although they were known to exist in the area. Carvajal (2005) captured and radio-collared 8 margays (5 males, 3 females) in the cloud forest of El Cielo from June 2001 to August 2004. The males maintained an average home range size of 4.03 km2 and a primarily nocturnal activity pattern. Carvajal (2005) also captured 14 gray foxes in the same study area. I became aware of several recently published and unpublished hair-snare studies targeting felids during my study. Results of these studies revealed clear patterns in the success of hair-snare studies (Table 2). In areas outside the range of gray fox or with low gray fox density, the hair-snare technique has proven successful in detecting felids. Lynx have been detected north of the range of gray fox in Canada (McDaniel et al. 2002, Weaver 2002), Idaho, Montana and Washington (Potter and Plumb 2001). Ocelot have been detected in South Texas, an area of very low gray fox density (Weaver 2002, Shinn 2002; Figure 4). However, surveys within the range of gray fox failed, with the preDominant species detected being gray fox (Table 2; Figure 4). No obvious difference in the lure used in successful and unsuccessful studies was observed. McDaniel et al. (2002) tested the efficacy of 5 different attractants including beaver castoreum and catnip oil, Cat PassionTM , Pacific CallTM, Hawbacker's Cat Lure #1TM, and BB1TM. All 5 lures tested successfully detected lynx (n = 60) with beaver castoreum and catnip oil eliciting the greatest visitation (n = 21). Weaver et al (2003) 15
and Shinn (2002) successfully employed Cat CallTM to detect ocelot in south Texas. All successful studies employed a visual attractant. Hair-snare studies that primarily detected gray fox utilized Cat CallTM, Canine CallTM and beaver castoreum and catnip oil. Visual attractants were employed in only some of these trials. All lures employed in unsuccessful trials are known to be effective in attracting felids. As opposed to hair-snare studies, scent-station surveys have successfully detected and/or monitored bobcat population trends in areas within gray fox range (Brady 1979, Hon 1979, Knowlton and Tzilkowski 1979, Morrison et al. 1979, Chamberlain et al. 1999). The method of species detection is the main difference between the 2 techniques. By virtue of the hair-snare design, visitation at a given station is only recorded by hair left on the pad. The scent-station design utilizes track identification for detection and consequently is not dependent on an animal's face-marking behavior. It is therefore possible to detect species that may have approached a station and turned away. The existence of the above pattern in hair-snare studies warrants investigation of possible interference by gray fox with felid markings. For example, perhaps the skunklike odor emitted from gray fox when marking (personal observation; M. Chamberlain personal communication) renders the station unattractive to felids. If true, cats may approach the station out of curiosity, but choose not to face-rub. This behavior would explain the higher success of scent-station studies in detecting felines approaching the site and also the consistent failure of snares to gather hair in areas of gray fox. Testing this hypothesis with captive animals would be informative. Gray fox are sympatric with coyotes throughout much of their range (Figure 5) and competition for resources such as space and food would be expected. Negative 16
spatial relationships exist between gray fox and coyotes. Coyotes seemed to limit the abundance and distribution of gray fox more as a result of physical dominance and intraguild predation than dietary competition (Fedriani et al. 2000). Gray fox, in turn, alleviate spatial competition with coyotes through behavioral avoidance. Consequently, low dietary competition was documented by Fedriani et al. (2000) in the Santa Monica Mountains of California. The low dietary overlap between the omnivores was explained by predation on different species of small mammals (rodents and lagomorphs) due to differential habitat use. Gray fox are also sympatric with bobcats throughout much of its range (Figure 6). In the Santa Monica Mountains of California, intraguild predation has been documented by bobcats on gray fox as well as a high dietary overlap between the species. However, behavioral avoidance by gray fox to alleviate spatial or dietary competition is not reflected in this relationship. The authors suggested circumstances such as behavioral dominance of gray fox over bobcat or the combined interaction of coyote, gray fox and bobcat as possible explanations (Fuller, personal communication). Bobcat and gray fox have been known to overlap their home ranges, yet their core areas were separate (Chamberlain, personal communication). Fox will be more apt to mark their core areas more frequently than the remainder of their home range. The information presented from the present study and from previous hair-snare studies (Table 2) suggests avoidance by felines. Conversely, there is no evidence of gray fox avoidance of felid markings. The core-area partitioning observed between gray fox and bobcat may be a case of felid avoidance of gray fox. Further study is required to address this question. 17
Results of this study suggest gray fox select macrohabitats as a whole, i.e., oakpine forest, and not individual microhabitat variables. These findings are comparable to previous studies. Chamberlain and Leopold (2000) noted gray fox preferred mature pine stands in nearly all seasons. Casselmen (1989) reported similar findings of gray fox selection of mature pine sawtimber in Louisiana. In Missouri, oak-hickory forests are selected by gray fox for cover and foraging (Haroldson and Fritzell 1984). In light of these studies, it appears hair-snare trials were an effective means to study habitat use of gray fox. MANAGEMENT IMPLICATIONS Felid hair-snare surveys, regardless of design, are not effective or recommended in areas inhabited by gray fox. However, John Weaver's hair-snare protocols utilizing Cat CallTM and a visual aid appear to be sufficient in determining felid presence in areas outside the range of gray fox or in areas of low gray fox density. It is improbable that one can redesign the hair-snare station to prevent gray fox marking. Raising the height of the station above the cheek level of gray fox is problematic due to the species' ability to climb (Neale and Sacks 2001). An alternative would be to change the lure. Fox, however, are motivated by olfactory stimuli (Chamberlain et al. 1999) and will presumably investigate any novel scent in their surroundings. The presence of a visual attractant is not relevant due to the apparent refusal of felids to face-rub sites previously marked by gray fox. I recommend a modified scent-station technique instead of the hair-snare design to determine feline presence or absence in areas inhabited by gray fox. Scent-stations 18
rely on tracks for species identification; therefore, they have the ability to detect species that may have approached the station but turned away. Traditional scent-stations were originally designed to detect canids; therefore the protocols need to be customized to meet the demands of detecting felids. Despite the type of lure station (i.e., scent or hair-snare) a possibility exists that a felid will approach, but turn away if a gray fox has marked it. Chamberlain et al. (1999) augmented the station's ability to detect felids after concern that low visitation was due to bobcats approaching the stations but not being recorded. The addition of 100-m transects consisting of roadside ditches and edges was made adjacent to each scent-station. Visitation to these transects was recorded as well. Another option would be to increase the diameter of the tracking circle. The use of sifted agricultural lime rather than soil to cover the tracking surface is another alternative (Morrison et al. 1981). Although claimed to have increased bobcat visitation in the study, no comment was made as to whether or not higher visitation was directly a function of higher readability potential with lime. Felids hunt with their eyes and ears whereas canids are more olfactory hunters. I strongly advise the use of a visual or audio attractant at each station. Chamberlain et al. (1999) reported highest bobcat visitation rates at stations equipped with a mechanical cottontail rabbit (Sylvilagus floridanus) distress call. Weaver considers hair-snare stations incomplete without the aid of a pie plate for visual attraction (Weaver, personal communication). Another alternative to the hair-snare technique is the use of camera traps. Advantages to this methodology include that it is not subject to observer bias and it has 19
an increased ability to differentiate individuals. Disadvantages include cost and reduced sampling effort. 20
REFERENCES Alderton, D. 1993. Wildcats of the world. Blandford Publishing, United Kingdom. Bonham, C.D. 1989. Measurements for terrestial vegetation. John Wiley and Sons, New York, New York, USA. Brady, J. R. 1979. Preliminary results of bobcat scent-station transects in Florida. Proceedings of the Bobcat Research Conference. National Wildlife Federation Science and Technical Series. 6:8-12. Carrillo, E., G. Wong, and A. D. Cuaron. 2000. Monitoring mammal populations in Costa Rican protected areas under different hunting restrictions. Conservation Biology 14:1580. Carvajal, S. 2005. Ambito hogarereno y patron de actividad del margay Leopardus wiedii (Schinz, 1821) en la reserve de la biosphere "El Cielo" Tamaulipas, Mexico. Thesis, Instituto Tecnologico De Ciudad Victoria, Tamaulipas, Mexico. Casselman, T. A. 1989. Home range, habitat use, and activity of gray foxes in managed southern pine forest. Thesis, Louisiana State University, Baton Rouge, Louisiana, USA. Cat Survival Trust. 1996. http://www.catsurvivaltrust.org/bobcat.htm. Accessed August 13, 2005. Chamberlain, M. J., and B. D. Leopold. 2000. Spatial use patterns, seasonal habitat 21
selection and interactions among adult gray foxes in Mississippi. Journal of Wildlife Management 64:742-751. Chamberlain, M. J., J. W. Mangrum, B. D. Leopold, and E. P. Hill. 1999. A comparison of attractants used for carnivore track surveys. Proceedings of the annual conference of the Southeastern Association of Fish and Wildlife Agencies 53:297-304. CITES. 2001. Appendices I, II and III to convention on international trade in endangered species of wild fauna and flora. US Government Printing Office, Washington, D.C., USA. Collier, G.E., and S. J. O'Brien. 1985. A molecular phylogeny of the Felidae: immunological distance. Evolution 39:473-487. Conner, M. C., R. F. Labisky and D. R. Progulske. 1983. Scent-station indices as measures of population abundance for bobcats, raccoons, gray foxes, and opossums. Wildlife Society Bulletin 11:146-152. Conroy, M. J. 1996. Techniques for estimating abundance and Species richness. Pages 179-191 in D. E. Wilson, F. R. Cole, J. D. Nichols, R. Rasanayagam, and M. S. Foster, editors. Measuring and monitoring biological diversity. Smithsonian Institution Press, Washington, USA and London, UK. de Oliveira, T. G. 1998. Leopardus wiedii. Pages 1-6 in E. Anderson, L. N. Caraway, K. F. Koopman, and D. S. Rogers, editors. Mammalian species. American Society of Mammalogists, New York, N.Y., USA. Diefenbach, D. R., M. J. Conroy, R. J. Warren, W. E. James, L. A. Baker, and T. Hon. 1994. A test of the scent-station survey technique for bobcats. Journal of 22
Wildlife Management 58:10-34. Eizirik, E., S. L. Bonatto, W. E. Johnson, P. G. Crawshaw Jr., J. C. Vie, and D. M. Brousset. 1998. Mitochondrial DNA control region in two Neotropical cats (Mammalia, Felidae). Journal of Molecular Evolution 47:613-624. Fedriani, J. M., T. K. Fuller, R. M. Sauvajot, and Eric C. York. 2000. Competition and intraguild predation among three sympatric carnivores. Oecologia 125:258-270. Gram, W. K., and J. Faaborg. 1999. The distribution of Neotropical migrant birds wintering in the El Cielo Biosphere Reserve, Tamaulipas, Mexico. The Condor 99:658-670. Guggisberg, C. A. W. 1975. Wildcats of the world. Taplinger Publishing, New York, New York, USA. Haroldson, K. J., and E. K. Fritzell. 1984. Home ranges, activity, and habitat use by gray foxes in oak-hickory forest. Journal of Wildlife Management 48:222-227. Harrison R. L. 1997. Chemical attractants for Central American felids. Wildlife Society Bulletin 25:93-97. Hon, T. 1979. Relative abundance of bobcats in Georgia: survey techniques and preliminary results. Proceedings of the Bobcat Research Conference. National Wildlife Federation Science and Technical Series. 6:8-12. IUCN/SSC Canid Specialist Group. 2005. http://www.canids. org/species/ Canis_latrans.htm. Accessed August 13, 2005. IUCN/SSC Canid Specialist Group. 2005. http://www.canids.org/species/ Urocyon_cinereoargenteus.htm. Accessed July 20, 2005. Johnson, W. E., M. Culver, J.A. Iriate, E. Eizirik, K.L. Seymore and S. J. O'Brien. 1998. 23
Tracking the evolution of the elusive Andean mountain cat (Oreailurus jacobita) from mitochondrial DNA. Journal of Heredity 89:227-232. Johnson, W.E. and O'Brien, S. J. 1997. Phylogenetic reconstruction of the Felidae using 16s rRNA and NADH-5 mitochondrial genes. Journal of Molecular Evolution 44: (Supplement 1) S98-Sll6. Jones, F. A., and D. L. Gorchov. 2000. Patterns of abundance and human use of the vulnerable understory palm, Chamaedorea radicalis (Arecaceae), in a montane cloud forest, Tamaulipas, Mexico. The Southwestern Naturalist 45:421-430. Kocher, T., W. Thomas, A. Meyer, S. Edwards, S. Paabo, E. Villablanca, and A. Wilson. 1989. Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of National Academy of Sciences 86:6 196- 6200. Knowlton, F. F. and W. M. Tzilkowski. 1979. Trends in bobcat visitations in scent survey lines in western United States. Proceedings of the Bobcat Research Conference. National Wildlife Federation Science and Technical Series. 6:8-12. Konecny, M. J. 1989. Movement patterns and food habits of four sympatric carnivore species in Belize, Central America. Pages 243-264 in K. H. Redford and J. F. Eisenberg, editors. Advances in neotropical mammology. Sandhill Crane Press. Gainsville, Florida, USA. McDaniel, G. W., K. S. McKelvey, J. R. Squires, and L. F. Ruggiero. 2000. Efficacy of lures and hair-snares to detect lynx. Wildlife Society Bulletin 28:119-213. Mondolfi, E. 1982. Notes on the biology and status of the small cats in Venezuela. Pages 125-146 in S. D. Miller and D. D. Everett, editors. Cats of the world: biology, 24
conservation and management. National Wildlife Federation. Washington, D.C., USA. Morrison, D. W., R. M. Edmunds, G. Linscombe, and J. W. Goertz. 1981. Evaluation of scent-station variables in northcentral Louisiana. Proceedings of the Bobcat Research Conference. National Wildlife Federation Science and Technical Series. 35:281-291. Neale, J. C. C. and B. N. Sacks. 2001. Food habits and space use of gray foxes in relation to sympatric coyotes and bobcats. Canadian Journal of Zoology 79:1794-1800. Nowell, K., and P. Jackson. 1996. Wild cats: status survey and conservation action plan. IUCN/SSC Cat Specialist Group, International Union for the Conservation of Nature and natural resources (IUCN), Gland, Switzerland. Peterson, M. B. 2001. Resource use and livelihood strategies of two communities in the El Cielo Biosphere Reserve, Mexico. Thesis, Miami University, Oxford, Ohio, USA. Pocock, R. I. 1941. The races of the ocelot and the margay. Osgood Testimonial Volume 319-369. Potter, T., and G. Plumb. 2001 May 9. Lynx workshop Missoula March 8-9, 2001. http://www.forestry.umt.edu/research/cesu/NEWCESU/Assets/ partner%20Activities/ Lynx%20Notes.pdf. Accessed August 22, 2005. Roughton, D. R., and M. W. Sweeny. 1982. Refinements in scent-station methodology for assessing trends in carnivore populations. Journal of Wildlife Management 46:217-229. Sargeant, G. A., D. H. Johnson, and W. E. Berg. 1998. Interpreting carnivore scent- 25
station surveys. Journal of Wildlife Management 62:1235-1245. SAS Institute. 1990. SAS/STAT users guide, version 6. SAS Institute, Cary, North Carolina, USA. Shinn, K. J. 2002. Ocelot distribution in the lower Rio Grande Valley National Wildlife Refuge. Thesis. University of Texas-Pan American. Edinburg, Texas, USA. Sunquist, M., and F. Sunquist. 2002. Wildcats of the world. University of Chicago Press,London, UK and Chicago, Illinois, USA. Tewes, M. E., and D. J. Schmidly. 1987. The neotropical felids: jaguar, ocelot, margay, and jaguarundi. Pages 696-711 in M. Novak, J. A. Baker, M. E. Obbard and B. Malloch, editors. Wild furbearer management and conservation in North America. Ministry of Natural Resources, Ontario, Canada. Turback, G. 1998. Seeking the missing lynx. http://www.nwf.org/national wildlife/article.cfm?articleId=114&issueId=15. Accessed September 22, 2005. Vazquez, L. B., R. A. Medellin and G. A. Cameron. 2000. Population and community ecology of small rodents in montane forest of western Mexico. Journal of Mammalogy 81:77-85. Weaver, J. 2002. Favorite beastie. Wildlife Conservation (April):10-11. Weaver, J., P. Wood, and D. Paetkau. 2003. A new non-invasive technique to survey ocelots. http://www.wcs.org/media/file/Ocelot0002.pdf. Accessed June 23, 2005. 26
Table 1. Microhabitat characteristics of hair-snare sites on the El Cielo Biosphere Reserve. Sites include those marked by gray fox (hit) or all other sites (not hit).
Herbaceous
Hit
Not Hit
Variable
N
xY
SE
N
xY
SE
Canopy Cover (%)
52
31.2
3.2
111
29.9
1.9
Grass (%)
46
1.4
0.7
112
1.2
0.4
Forb (%)
50
23.4
3.0
114
30.2
2.1
Bare ground (%)
48
14.3
3.0
110
10.8
1.5
Litter (%)
51
49.0
4.1
113
48.5
2.5
Rock (%)
51
19.5
2.7
114
20.0
1.8
Understory 51
12.1
1.3
113
9.7
0.4
density (m)1
1 Distance at which margay model could not be seen from hair-snare site.
27
Table 2. Summary of results from published and unpublished (*) hair-snare studies targeting felids.
Author/Year
Study Area
Mc Daniel et al. Yukon,
2002
Canada
Shinn 2002 Weaver et al 2003
Texas Texas Texas
Type of lure
Visual attractant
5 types
Pie Plate
Cat CallTM Pie Plate Cat CallTM Pie Plate
Cat CallTM Pie Plate
Target species Lynx canadensis Leopardus pardalis L. rufus L. pardalis
Success Yes Yes Yes Yes
Total stations (n) 390 250 250 89
Hits
Hits
by target by
species gray fox
60
0
8
2
29
2
23
0
Success by target species (%) 15.4
Success by gray fox (%) 0
3.2
0.8
11.6
0.8
25.8
0
*Beier 2000
Arizona
Cat CallTM No
L .concolor No
20
0
12
0
60.0
*Harrison 2003 New
Cat CallTM Pie Plate
L. rufus
No
631
1
50
0.16
7.9
Mexico
*Frangioso 2003
California
Cat CallTM Pie Plate
L. concolor L. rufus
No
115
1
52
0.87
45.2
L. concolor
*Childs 2004 Arizona
Canine
No
Panthera
No
?
11
190
?
?
CallTM
onca
*Downey 2005
Tamaulipas, Mexico
Cat CallTM No
L. wiedii
No
250
0
52
0
20.8
* P. Beier, Northern Arizona University; J. Childs, Wildlife Conservation Society; K. Frangioso, Wildlife Conservation Society; R. L. Harrison, University of New Mexico.
28
Figure 1. Geographic range of Leopardus wieidii: 1, L. w. amazonica; 2, L. w. boliviae; 3, L. w. glaucula; 4, L. w. nicaraguae; 5, L. w. oaxacensis; 6, L. w. pirrensis; 7, L. w. salvinia; 8, L. w. vigens; 9, L. w. wiedii; 10, L. w. yucatanica (de Oliveira 1998). 29
30
Figure 2. Diagram depicting layout of hair-snare station (center) and vegetation plots (5 plots/station) arranged 10 m from hair-snare station. 31
32
Figure 3. Frequency of gray fox hits at hair-snare stations by season in each habitat (oakpine, cloud, and subtropical-deciduous) at El Cielo Biosphere Reserve, Tamaulipas, Mexico 33
14
12
10
Frequency
8
Wet
6
Dry
34
4
2
0 Oak-pine
Cloud
Sub TD
Habitat Type
Figure 4. Felid hair-snare study sites plotted against gray fox range (shaded; IUCN/SSC 2004). Red sites successfully detected felids. Blue sites failed and predominantly detected gray fox. 35
36
Figure 5. North American range of coyotes (Canis latrans; IUCN/SSC 2004). 37
38
Figure 6. North American range of bobcat (Lynx rufus; Cat Survival Trust 1996). 39
40
VIDA Master of Science Thesis: HAIR-SNARE SURVEY TO ASSESS DISTRIBUTION OF MARGAY (LEOPARDUS WIEDII) INHABITING EL CIELO BIOSPEHERE RESERVE, TAMAULIPAS, MEXICO Major Field: Zoology Biographical: Education: Graduated from Bishop McGuiness High School, Oklahoma City, Oklahoma, May 1988; received Bachelor of Science degree in Geology from Oklahoma State University, Stillwater, Oklahoma in 1994. Completed the requirements for the Masters of Science degree with major in Zoology at Oklahoma State University in December 2005. Experience: Pachyderm keeper, Oklahoma City Zoo, 1999 through present Professional memberships: American Association of Petroleum Geologists Wildlife Conservation Society
Name: Patricia J. Downey Institution: Oklahoma State University
Date of Degree: December, 2005 Location: Stillwater, Oklahoma
Title of Study: HAIR-SNARE SURVEY TO ASSESS DISTRIBUTION OF
MARGAY (LEOPARDUS WIEDII) INHABITING EL CIELO BIOSPHERE RESERVE,
TAMAULIPAS, MEXICO
Pages in Study: 40
Candidate for the Degree of Master of Science
Major Field: Zoology
Scope and Method of Study: The margay (Leopardus wiedii) is an endangered neotropical felid listed on Appendix 1 of CITES. Knowledge of margay ecology is limited because little in-situ research has been conducted in any part of its range which extends from northern Mexico to South America. The objectives of the research were to (1) investigate seasonal habitat use of the margay and (2) determine which microhabitat variables are preferred by margays inhabiting El Cielo Biosphere Reserve, Tamaulipas Mexico. I conducted 8 hair snare surveys (4 wet-season and 4 dry-season) from November 2002 to October 2004. Each survey consisted of 30 hair-snare stations placed every 500 m along a transect spanning 3 habitats.
Findings and Conclusions: Species analysis did not detect margay on any pad. Gray fox (Urocyon cinereoargenteus) was the predominantly detected species. Statistical analysis was therefore only carried out on gray fox data. Of the 3 habitats surveyed, gray fox detections were greater in oak-pine forest than cloud or tropical-deciduous forests across seasons. Gray fox did not prefer any microhabitat variable or combination of variables. I compared the results of unpublished and published hair-snare surveys and discovered clear patterns in the success (i.e., detection of the target felid) of this technique. The method succeeded only outside the range of gray fox. The technique fails within the range of gray fox and the predominant species detected is consistently gray fox. I do not recommend the hair-snare methodology to survey felids within the range of gray fox. I advise a modified scent-station protocol equipped with a visual or auditory attractant to survey felids within gray fox range.
ADVISER'S APPROVAL: Dr. Eric C. Hellgren

PJ Downey

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