한국환경생태학회지 29(2) : 193~200, 2015 pISSN 1229-3857 eISSN 2288-131X Korean J. Environ. Ecol. 29(2): 192-199, April 2015 http://dx.doi.org/10.13047/KJEE.2015.29.2.192

Movement and Home Range of the Red-Tongued Viper ( ussuriensis) Inhabiting Gapado1a

Byoung-Soo Kim2, Hong-Shik Oh3*

ABSTRACT

This study was conducted to investigate the movement and home range of the red-tongued viper snake (Gloydius ussuriensis) from June 2006 to June 2009. This snake inhabits an islet on Jeju Island, Gapado. A total of 132 individual were marked during the study. Among the marked individuals, the number of snakes recaptured more than once was 22 (16.8 %) and the number of individuals recaptured more than twice was eight (6.1 %), indicating a relatively low recapture rate. The durations from capture to recapture varied from 1 to 710 days. However, the capture points were not much different, indicating that the moved distance of snakes and the interval between capture-recapture were not correlated. The home ranges of the Red-tongued viper snakes calculated from data of the snakes which were captured more than three times using the MCP(minimum convex polygon) method were 8∼167 ㎡(64.0±57.0 ㎡), suggesting that this snake is relatively sedentary. Home range size differences between female (Mean=62.0 ㎡) and male (Mean=66.0 ㎡) snakes were not significant. In the red-tongued viper population of Gapado, there was no statistically significant relationship between body size and home range size although it was positively correlated (r=0.675). Our results provide valuable data to understand life patterns of the red-tongued viper snakes and will be useful when conducting further ecological studies on other snake species.

KEY WORDS: BEHAVIORAL ECOLOGY, POPULATION, , REPTILIA, JEJU ISLAND

INTRODUCTION either (1) difficult to be predicted because of random movement, or (2) sedentary whether they are concentrated In regards to the life history of , moving from or not concentrated in one area (Seigel et al., 2001). one habitat to another habitat is related to the securing Recently, it was revealed that snakes' moving distance and of resources including food and water, mating, sunbathing, home range differ by species, time slot, gender and hibernation, nest, hideout, survival and reproduction. Most seasonal factors (Shine and Lambeck, 1985; Shine, 1987; animals move only when it is necessary, because their Fitzgerald et al., 2002; Brito, 2003; Shine et al., 2003; movement is associated with energy loss, unfavorable Whitaker and Shine, 2003). In particular, the red-tongued environmental conditions and exposure to predators. Also, viper snakes, which belong to the Gloydius, are very how often and how far animals move differ greatly from sedentary like that inhabit an islet one species to another (Pough et al., 2004). in the northeast of (Shine et al., 2003). On the other In case of the movement patterns of snakes, they are hand, some snakes like Crotalus adamanteus, a type of

1 Received 06 August 2014; Revised (1st: 27 January 2015, 2nd: 07 March 2015); Accepted 08 March 2015 2 Shinseong Girl’s Middle School, Jeju 690-140, ([email protected]) 3 Dept. of Science Education, College of Education, Jeju National Univ., Jeju (690-756), Korea a This paper is to complement the lead author of doctor’s thesis was written. This work supported by the Jeju Green Environment center under the Research development program(2013). * Corresponding author: Tel: +82-64-754-3283, Fax: +82-64-725-4902, E-mail: [email protected] Movement and Home Range of the Red-Tongued Viper Snake (Gloydius ussuriensis) Inhabiting Gapado 193

rattlesnake that inhabits the United States, move relatively movement and home range. long distances (Waldron et al., 2006). Red-tongued viper Accordingly, this study was carried out to identify the snakes belong to Reptilia, , Serpentes, Crotalinae moving distance and home range of the red-tongued viper and Gloydius (O’Shea, 2005); they are distributed in snakes that inhabit Gapado, an islet of Jeju Island, by using Korea, northeast of China and far east of (Zhao the mark-recapture method, and to provide data necessary and Adler, 1993). Until now, in regards to the ecology for ecological studies on other types of snakes that inhabit of red-tongued viper snakes, studies have been conducted Korea. to identify their reproductive cycle and litter size (Kim and Oh, 2014a), sexual size dimorphism (Kim and Oh, MATERIALS AND METHODS 2014a) and prey use (Kim and Oh, 2014c). Also, a study on capture-recapture was conducted for resource This study was carried out in Gapado, the annexed management in national parks by using a PIT tag (Lee island located in the southwest of Jeju Island. The island's et al, 2009). However, there is no information on snakes' area is about 0.87 ㎢ or 1/2100 of Jeju Island. It belongs

Figure 1. A map and picture of the survey area at Gapado 194 Byoung-Soo Kim, Hong-Shik Oh Korean J. Environ. Ecol. 29(2) 2015

to Gapa-ri, Daejeong-eup, Seogwipo-si, Jeju Special of the individual no. G001 when it was recaptured. Figure Self-Governing Province in terms of administrative district. 2c shows how to insert the PIT tag under the skin. It is situated at latitude 33° 09′ 35″~ 33° 10′ 30″ and The capturing location was identified using a GPS longitude 126° 15′ 56″~126° 16′ 57″. Gapado is 20 m (eTrex Vista® C, Garmin), and the moving distance and above sea level and its ground is flat. Most of the island home range were calculated using MapSource program is farmland except some areas where there are villages. (ver. 6.8, Garmin). The MCP (Minimum Convex Polygon) Grasslands or simple habitats are developed at some places method was used to determine the size of home range of at the edge of the island, and this study was carried out the individuals that were captured more than three times. in the grasslands, which is located in the southwest of SPSS (ver. 12.0) statistical program was used for a statistical Gapado, where goats are grazed (Figure 1). analysis.  -test was used to analyze sex ratio of the This study used the mark-recapture method to find out captured individuals, and t-test was used to analyze the the movement and home range of red-tongued viper snakes difference in home range between male and female red- from June 2006 to June 2009. Because the number of tongued viper snakes. Also, a correlation between the surveys is determined by whether it is possible to enter moving distance and the interval between capture- recapture the island depending on weather conditions, a survey was and a correlation between the body length and the home carried out 1~4 times a month. In regards to the marking range size were analyzed through Pearson's correlation. of individuals, two methods were used: the method of ventral clipping abdominal scales and the method of RESULTS AND DISCUSSION inserting a PIT tag (Passive Integrated Transponder tag; ID 162A, TROVAN) under the skin (Figure 2). Figure In terms of the number of marked individuals by year, 2a shows the location where ventral clipping is conducted 13 individuals were marked in 2006, 42 individuals in to the individual no. 155, and Figure 2b shows the image 2007, 61 individuals in 2008 and 15 individuals in 2009. Among the 131 marked individuals, except two individuals of which gender marking was omitted, 56 were females and 73 were males. The ratio of males is higher (Males : Females = 1.30 : 1, n = 129), but there was no statistically significant difference (χ2-test, χ2 = 2.240, P> 0.1). (a) However, the ratio of males was higher than the sex ratio of the offspring of the individuals that inhabit Jeju Island (Males : Females = 1.15 : 1). There is no difference in the sex ratio of the offspring of most snakes (Wang et al., 2003; Taylor and Denardo, 2005), but because females face many difficulties due to reproduction (Shine, 1980), their mortality is higher. A good example is Trimeresurus (b) stejegeri stejegeri that inhabits Taiwan: the observed frequency of males was significantly higher than that of females when they become adults (Wang et al., 2003). Among the marked individuals, 22 (16.8 %) were recaptured, and 8 individuals (6.1 %) were recaptured more than twice (Table 1). The recapture rates after inserting the PIT tag were 16.67 % and 8.33 % regarding (c) the surveys that were carried out in Mt. Odae National Park and Mt. Wolak National Park (Lee et al., 2009), Figure 2. Individual marking methods of G. ussuriensis. Ventral clipping method (a) and its actual Accordingly, the recapture rate of this study (16.8 %) was example (b) and PIT tag inserting method (c) similar to the result of the survey conducted in Mt. Odae Movement and Home Range of the Red-Tongued Viper Snake (Gloydius ussuriensis) Inhabiting Gapado 195

Table 1. The moved distance and home range size of G. ussuriensis on Gapado

Individual classification Capture SVL Movement Home rang size No. Indi. Terms Tag cord Sex Times Date (㎜) distance(m) (㎡) No. (day) 1st 3 June 2006 363 1 G001 00066C6753 F 2nd 17 June 2007 379 364 13 60 3rd 17 June 2008 366 367 9 1st 17 June 2006 330 2 G009 00066C24AE M 2nd 17 June 2007 365 331 3 1st 13 May 2007 361 000669831E 3 G015 M 2nd 17 June 2007 35 365 44 112 (00066AC874) 3rd 26 October 2007 131 376 12 1st 13 May 2007 276 4 G018 00066C16C1 M 2nd 8 April 2008 331 284 3 12 3rd 12 April 2008 4 284 12 1st 23 May 2007 361 5 G023 00066C3095 F 2nd 2 May 2009 710 374 22 1st 23 June 2007 377 6 G032 00066C7D59 M 2nd 8 April 2008 290 377 12 1st 6 July 2007 343 7 G047 968000004228453 M 2nd 4 April 2008 273 350 18 1st 4 April 2008 350 2nd 15 April 2008 11 350 11 8 G056 968000004315834 M 3rd 31 May 2008 46 355 12 95 4th 10 October 2008 132 358 3 5th 19 Arpril 2009 191 358 11 1st 15 April 2008 327 9 G065 968000004317555 F 2nd 26 May 2008 41 334 17 1st 25 April 2008 365 10 G067 968000004312736 M 2nd 16 May 2008 21 371 24 44 3rd 10 June 2008 25 373 3 1st 29 April 2008 390 11 G071 968000004319549 F 2nd 31 May 2008 32 393 20 167 3rd 27 June 2009 392 394 32 1st 29 April 2008 356 2nd 18 May 2008 19 356 6 12 G072 968000004316800 F 12 3rd 31 May 2008 13 361 4 4th 1 June 2008 1 361 3 1st 6 May 2008 344 13 G078 968000004318134 M 2nd 26 May 2008 20 344 7 1st 16 May 2008 324 14 G085 968000004308989 F 2nd 31 May 2008 15 329 5 8 3rd 7 October 2008 129 332 6 1st 16 May 2008 338 15 G087 968000004221296 M 2nd 7 October 2008 144 350 4 1st 18 May 2008 322 16 G088 968000004225112 M 2nd 30 June 2008 43 340 11 1st 18 May 2008 335 17 G089 968000004222305 M 2nd 30 June 2008 43 346 18 1st 22 May 2008 292 18 G091 968000004218634 M 2nd 24 May 2009 367 305 30 1st 30 June 2008 349 19 G104 00066C53AB F 2nd 9 May 2009 313 361 27 1st 30 June 2008 353 20 G105 00066C24C1 F 2nd 7 October 2008 99 356 18 1st 3 October 2008 349 21 G110 0006677783 M 2nd 27 June 2009 267 360 3 1st 31 May 2009 362 22 G128 968000004223661 M 2nd 14 June 2009 14 362 22 196 Byoung-Soo Kim, Hong-Shik Oh Korean J. Environ. Ecol. 29(2) 2015

National Park but higher than that of Mt. Wolak National flasgellum, a snake that moves around to capture preys Park. Compared to the recapture rates of Vipera berus 45 % (Secor and Nagy, 1994). Also, it was reported that the (females 54 %, males 33 %), which inhabits Switzerland, moving distance of other snakes that belong to the and Drymarchon coupri 43 % (Stevenson et al., 2009), Viperidae, the snakes that capture preys by ambush, is which inhabits Georgia, the United States, recapture rates short: Vipera latastei. which inhabits northern Portugal, of red-tongued viper snakes were very low. The durations moves an average 5.4 m per day (Brito, 2003), and from capture to recapture varied from 1 to 710 days, but Gloydius shedaoensis, which inhabits a small island there was no correlation between the moving distance and situated at the northeast of China, moves an average 2 the interval between capture-recapture (n = 33, r = 0.173, m per day (Shine et al., 2003). P> 0.1). In case of G015, it moved 44 m during 35 days The results of calculating the home ranges of eight and 22 m during 14 days, but G001, G009, G018, G023, individuals that were captured more than three times using G072, G091 and G104 moved only 3-32 m during more the MCP (Minimum Convex Polygon) method show that than 300 days. Also, the interval between capture- the home ranges of the red-tongued viper snakes in recapture of other individuals varied but there was no Gapado were 8∼167 ㎡ (64.0㎡ ± 57.0 ㎡), considerably difference in their locations (Table 1). smaller than that of Vipera latastei in northern Portugal, The movement of snakes is associated with the securing which is 2,400 ㎡ (Table 1). Their home range is usually of resources including water and food, mating, sunbathing, identified by attaching a signal transmitter on their body. hibernation, nest, hideout and various factors required for Home range sizes vary among snakes (Pough et al, 2004), reproduction (Pough et al., 2004), and snakes' mortality and it is known to be affected by their body size, seasonal may go up by getting exposed to many risks such as factors, gender, reproduction and ecological characteristics energy loss and encountering with natural enemies. In (Shine, 1979; Shine and Lambeck, 1985; Fitzgerald et al., general, there may be a difference in the mortality by age 2002; Brito, 2003; Shine et al., 2003; Whitaker and Shine, due to the moving distance: young snakes primarily die 2003; Waldron et al., 2006). when dispersing after hatching out, and the mortality of Some snakes like G. shedaoensis, which inhabits an islet juvenile snakes that stay in one place is low. Also, male situated at the northeast of China (area 0.73 ㎢) and has snakes may die during the mating season because they average home ranges of 2,429 ㎡ and 2,851 ㎡ (< 0.3ha) need to move a long distance to find their mate, and for females and males respectively, are very sedentary and oviparous adult females may die while moving to lay eggs. move less than 2 m a day. On the other hand, the male Also, the mortality of ambush foragers, animals which are Hoplocephalus stephensii (Stephen's banded snake), which sedentary and capture their prey by ambush, is lower than belongs to the Elapidae and inhabits the forests formed that of active foragers, animals that hunt preys while in the east coast of Australia, has a large home range of moving a long distance (Bonnet et al., 1999). an average 202,000 ㎡ (Fitzgerald et al., 2002). The male As it is shown in the mark-recapture results, the Crotalus adamanteus (Eastern diamondback rattlesnake) red-tongued viper snakes living in Gapado tend to stay and the male C. horridus (Canebrake rattlesnake), the in one place and barely move long distances. This can kinds of rattlesnakes that inhabit same places in South have a positive effect in decreasing their mortality. Also, Carolina, the United States, also have very large home because most of the red-tongued viper snakes that inhabit ranges of 848,200 ㎡(169,100~3,104,800 ㎡) and 483,800 Gapado prey on small lizards like Scincella vandenburghi ㎡(86,000~1,075,000 ㎡) respectively (Waldron et al., and large centipedes such as Scolopendra subspinipes 2006). The red-tongued viper snakes that inhabit Gapado mutilans, they can minimize the loss in energy used while seem to be staying in one place, and their home range moving to other places where the availability of food was smaller than that of G. shedaoensis, which is very resources is low (Kim, 2011). In fact, there is a study that sedentary. The island where G. shedaoensis inhabit is reported that the energy consumption of Crotalus cerastes, isolated, but because it is situated at the migratory route the rattlesnake that inhabits the American Continent and of birds, many birds pass through this island in spring and captures preys by ambush, is lower than that of Masticophis autumn. Because G. shedaoensis preys on migratory birds Movement and Home Range of the Red-Tongued Viper Snake (Gloydius ussuriensis) Inhabiting Gapado 197

by ambushing them from trees or the ground (Shine et Gapado are highly sedentary and their habitats are limited, al., 2002a, 2002b, 2002c), it does not move a long distance a fact that contradicts their very low recapture rates. This to capture preys. Many red-tongued viper snakes that may be so because the size of red-tongued viper snakes inhabit Gapado were also observed to be hiding behind that inhabit Gapado is smaller than those living in Jeju stone walls or nearby them to ambush preys as shown in Island (Kim and Oh, 2014b). The red-tongued viper snakes Figure 3. They chose stone walls as their ambush sites that inhabit Jeju Island are also easily found as they usually because they can hide right away when in danger. Also, hide behind stone walls. the size of the home range of red-tongued viper snakes Home ranges of snakes also vary according to their that inhabit Gapado is much smaller than that of G. gender. While there is a small difference between the shedaoensis, which inhabits in a similar environment, home ranges of male and female G. shedaoensis, a very because there may be a difference in the size of available sedentary snake which inhabits an islet in the northeast habitats. G. shedaoensis can use the whole island as its of China (Shine et al., 2003), the average home range of habitat, but as for red-tongued viper snakes, except the female Epicrates inornatus, a snake that belongs to the village and farmland, they can only use the southwestern Boidae and inhabits Puerto Rico, 7,890 ㎡, is larger than part of Gapado, the survey site, and the grasslands formed that of its male (5,000 ㎡) counterparts (Puente-Rolón and at the edge of Gapado as their habitats. Bird-Picó, 2004). Also, there is a significant difference As such, the red-tongued viper snakes that inhabit between the home ranges of female and male Pseudonaja

(a) (b)

(c) (d) Figure 3. The habitat scene of G. ussuriensis on Gapa-do. It is consisted of several compartments by a stone wall line (a and b). G. ussuriensis prefer the space under the stone wall (c) or the surroundings f the stone wall (d) to the grassland center for ambush site 198 Byoung-Soo Kim, Hong-Shik Oh Korean J. Environ. Ecol. 29(2) 2015

textilis (Brown snake) and Hoplocephalus stephensii, very in Australia (Whitaker and Shine, 2003), but sometimes venomous snakes that belong to the Boidae and inhabit there is no correlation between them as in the case of Australia. The home ranges of male and female P. textilis, Gloydius shedaoensis which inhabits an islet in the which inhabit agricultural areas in the northeast part of northeast of China (Shine et al., 2003). The size of the Australia, are 117,900 ㎡ and 14,600 ㎡, males showing home range and the body length of the red-tongued viper eight times larger home ranges than that of females snakes living in Gapado did have a correlation but it was (Whitaker and Shine, 2003). Also, the home ranges of not statistically significant (Figure 4). male and female H. stephensii, which live in the forests Home ranges of snakes also vary according to seasonal in the east coast of Australia, are 202,000 ㎡ and 54,000 factors, ecological characteristics and the pregnancy status ㎡, males showing a four times larger home range than for females. In other words, the size of the home range that of females (Fitzgerald et al., 2002). Also, the home of the male Vipera latastei, a type of European Viperidae, range of male C. adamanteus, a kind of rattlesnake, is tends to get larger during September (Brito, 2003); the three times broader than that of nonpregnant females home range of G. shedaoensis, a type of Viperidae which (males: 842,000 ㎡, nonpregnant females: 286,300 ㎡) inhabits an islet in the northeast of China and captures (Waldron et al., 2006). According to the study result of preys through ambush, is very small (Shine et al., 2003). calculating the home range of the serpent Elaphe Home range sizes also vary depending on the pregnancy schrenckii, which inhabits Korea, using the MCP method of females: the home range of pregnant snakes is smaller based on the radio tracking result, the home range of males than that of nonpregnant ones (Whitaker and Shine, 2003; (389,600 ㎡) was two times larger than that of females Waldron et al., 2006). Accordingly, because there are (162,500 ㎡) (Lee et al., 2009). As such, the difference various factors that affect the home ranges of snakes, more between the home ranges of females and the males are studies need to be carried out to verify the details of the more evident in snakes with large home ranges. On the ecological characteristics of snakes. other hand, because the red-tongued viper snakes in This study was conducted in a limited area, but can Gapado mostly stay in one place, their home range was serve as precious data in understanding the Viperidae very small: there was no significant difference between which play an important role in maintaining biodiversity. the home ranges of females, 62.0 ㎡ (n = 4), and males There aren't any ecological studies on snakes that inhabit 66.0 ㎡ (n = 4). (t=0.092, df=6, P>0.05). Korea, and data from this study can serve as models for Sometimes there is a correlation between the size of conducting ecological studies on other snakes that are the home range and the size of a snake such as Pseudonaja necessary in improving the diversity. textilis, a big snake with a large home range which lives REFERENCES

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