Oophaga pumilio (strawberry poison dart frog) sex dependent microhabitat selection
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1 Oophaga pumilio (strawberry poison dart frog) sex dependent microhabitat selection University of Manitoba Adam Borton April 07, 2016 (Photo: Kevin McRae) 1
2 1.0 Introduction Widespread population decline of various animal species can often times be attributed to increased human activity. Influences such as agricultural or urban development encroach upon species habitat, causing fragmentation of once continuous environments (Nowakoski et al., 2012). Such influences can be detrimental to populations, as individuals cope with the changing landscapes. Amphibian populations have been declining over the years as essential habitat is lessened by humans. In fact, in comparison to other vertebrate groups, amphibians are the most threatened group of animal species in the world with much of the decline being attributed to human influence and land-use (Murasaki, 2010). In order to prevent further destruction of critical habitat of amphibians, it is important to understand the geographic range in which amphibians live within. An amphibian s home range can be defined as the area which includes the space required for the individual to accomplish its daily movement and activity (Murasaki, 2010). Understanding a species home range is extremely important from both an ecological and conservation standpoint as it will highly influence distribution of the species, population dynamics, resource use, as well as inter and intraspecific interactions (Murasaki, 2010). Through focusing on factors which affect how and why a species occupies a specific site, conservation efforts can begin to be implemented and further loss of habitat avoided. 1.1 Oophaga pumilio Strawberry Poison Dart Frog A species which has begun to be affected by habitat loss is Oophaga pumilio. Oophaga pumilio (O. pumilio), commonly known as the Strawberry Poison Frog, is a diurnal species of frog belonging to the Dendrobitadae family (Murasaki, 2010). The 2
3 species is often found in tropical forested environments in Nicaragua, Costa Rica and Panama (Meuche, Linsenmair & Prohl, 2011). Dendrobatid frogs have evolved a defense mechanism to negate predation, through obtaining chemicals through their diet of alkaloid-containing organisms (Stynoski et al., 2014). After consumption, poisonous chemicals can then be secreted through specific glands found on their skin, enlarging as the organism transitions from juvenile to mature life stages (Stympski et al., 2014). The species typically occupies moist, dense, lowlands or swamps and avoids open, exposed, pasture-like areas in order to avoid dehydration (Nowakowski et al., 2013). Oophaga pumilio is said to be one of the most phenotypically diverse frogs observed, exhibiting a range of colours varying with location. While in Nicaragua and Costa Rica the frog typically appears red-orange with blue-black appendages, it is common to find the frog appearing pure orange or green in other locations (Galeano & Harms, 2016). At maturity, Oophaga pumilio grow to be anywhere from 17.5 to 24 millimetres long (Nowakowski et al., 2013). While current knowledge on amphibian land use is limited in some regards, much research has gone into understanding behavioral and reproductive interactions between O. pumilio. Using such knowledge can be beneficial from a conservation standpoint, as understanding what environments or physiographic features a frog may occupy could benefit protection efforts. Territorial ranges differ between male and female O. pumilio with female frogs maintaining larger territories than males (Meuche, Linsenmair & Prohl, 2011). In general, males compete for dominance against one another in order to defend their home range, attract females and secure reproductive sites (Murasaki, 2010). Males typically occupy smaller home ranges than females as smaller areas are easier to defend 3
4 from other male frogs (Murasaki, 2010). Male O. pumilio territory typically includes locations which are conducive to attracting or courting females, namely oviposition and courting sites located on dry leaves in the organic litter of the forest floor (Meuche, Linsenmair & Prohl, 2011). Male O. pumilio distribution is often influenced by high densities of female frogs and territory is selected for areas with an absence of competing males (Murasaki, 2010). Female territorial ranges are typically larger with movement within the home range occurring much more frequently than males (Murasaki, 2010). Females are involved with much of the parental responsibility, securing territory for better access to clean water or food sources (Meuche, Linsenmair & Prohl, 2011). Since female O. pumilio are involved with raising young individuals, it is common to find female home ranges next to tadpole-rearing sites such as water filled axils of plants or small, calm bodies of water (Meuche, Linsenmair & Prohl, 2012). The axils of bananas or heliconias provide suitable sites for such rearing locations as well (Meuche, Linsenmair & Prohl, 2011). Once females have established their home range, they commonly spend much of their time within that area to tend to their young and secure food. Female O. pumilio home ranges are typically located in locations which allow access for multiple male mating (Murasaki, 2010). While the home range of male and female O. pumilio are undoubtedly distinct from one another, overlap does occur. Due to the frogs preferring moisture to prevent dehydration, it is quite common to find both male and female frogs in cool, damp areas beneath the organic leaf litter or within the roots of plants (Meuche, Linsenmair & Prohl, 2011). 4
5 The objective of this analysis was to determine whether there was an association between the gender of O. pumilio and common locations in which it is found. By analyzing if such an association is present, valuable insight on species site preference could be attained. Research has shown that the roles and responsibilities of O. pumilio are dependent upon gender, with both male and female individuals being involved with different aspects (Murasaki, 2010). Could gender responsibilities and habits then be used to explain why individuals occupy certain features of the habitat and distributions are situated the way they are? Perhaps females are more commonly found in trees or on leaf axils due to their parental and offspring rearing responsibilities. Perhaps male distributions are often found on the ground or within the leaf litter due to their competitive and reproductive habits. Research conducted in the past by Seiichi Murasaki, looked to determine rates of movement as well as how both genders of O. pumilio distributed themselves within environments. This particular study determined that females and juveniles occupied significantly larger home ranges then males (Murasaki, 2010). However, the objective of our analysis focuses on the actual natural features the frog was found upon (tree, ground or leaf) and the association that factor has with gender. Size of home range of male and female frogs was not studied in this analysis. Understanding how O. pumilio situates itself and operates within its home range would be beneficial when looking to protect and conserve the species. If such an association with gender is present, data collected could be used to better mitigate and avoid humanrelated damage to specific preferred O. pumilio habitat based on gender. Location information would also increase efficiency when conducting studies on O. pumilio as less time would theoretically be spent locating the species. 5
6 2.0 Methods 2.1 Study Region The study region was located on Isla Colon, Bocas Del Toro, Panama. Isla Colon is the northernmost Island in the Bocas Del Toro Archipelago, located east of mainland Panama in the Caribbean Sea. Bocas del Toro lies 9 degrees above the equator in the wet tropical zone. The average annual rainfall and temperature are 4000 mm and 30 C (86 F) respectively (ITEC, 2015). The island is highly diverse and offers habitat for several thousand different organisms. Many ecosystems occur within the region such as tropical lowland rainforest, which was the focus of this study. We collected our study samples from one population located in Dixon Forest, near the Bocas Del Toro Biological Station. 2.2 Dixon Forest Dixon Forest is approximately 6km 2 and is comprised of tropical lowland rainforest; a primary growth forest located on the northwest corner of Isla Colon. These forests are home to many large ficus trees, which are more than 100 feet tall. These trees are a keystone species in the area, and their buttresses create micro ecosystems for many species including O. pumilio. The ground is layered in leaf litter with many species of low-lying vegetation such as philodendrons and dieffenbachia. Nearby areas are cattle pastures, secondary forest, marshes and human settlement. 6
7 Fig 1. Dixon Forest study location, ITEC, Study Site Two survey locations (Plot A and B) were randomly selected within Dixon Forest. Plots were located approximately 400 meters apart. Each plot was designed as a 20x20 meter quadrat. Quadrats were created using a tape measure and a compass to ensure that the plot remained square. Once the quadrat was created, surveying flags were placed to form the shape for the remainder of the study. 2.4 Sampling Methods Plots were surveyed for 60 minutes every second day, for a total of three times each. When a frog was located, the searcher would place a piece of surveying tape to mark the location. The searcher would than capture the frog, place the frog in a zip lock 7
8 bag and place the bag at the first observed location. Careful consideration were given that the frogs were not injured during the process. The frogs were only handled for a short amount of time, as amphibians have permeable skin and should not be touched with dry hands for long periods. Once the 60 minute period was complete, we began to record the data. Each frog was measured, sexed and its first observed location documented on a field message pad. Each frog was photographed with a digital camera to ensure that is was not a recaptured frog. O. pumilio on Isla Colon all have different spotted patterns making individuals distinguishable. Recaptured frog data observations were not included in this study. Fig 2. O. pumilio on Isla Colon displaying the variation of pattern. McRae, 2013 O. pumilio are easily sexed on Isla Colon as male frogs have a dark colouration on their throats while females have lime green colouration. Frogs were measured with the use of a pair of calipers. When we returned to the biological station, data recorded on the field message pad was transferred to a Microsoft Excel spreadsheet. 8
9 Table 1: O. pumilio contingency table, McRae, Tree Leaf Ground Total Male Female Total Statistical Analysis A chi-square analysis was used to determine significant differences in observed location in relation to frog gender. All assumptions were met to perform this test; the sampling method is simple random sampling, the study variables are categorical. The value of the number of sample observations for each variable is at least 5. A 2x3 contingency table was used to determine gender and perch. Frogs from both samples plots were separated into two categories: male and female. The second variable, first observed location was separated into three categories: trees, leaf and ground. The data was compiled in a contingency table seen in Table 1. All data was analyzed in IBM SPSS Statistics Results It was determined that there is sufficient evidence to support the null hypothesis and that the locations of sighted O. pumilio does in fact occur independent of their gender. 9
10 3.1 Descriptive Statistics Prior to initiating the inferential analysis of the O. pumilio information, the descriptive statistics should be collected, measured, and analyzed. The three types of measures within descriptive statistics to be analyzed consist of central tendency, measure of dispersion, and measure of shape. The purpose of utilizing the method of measuring descriptive statistics within the scope of the research at hand is to efficiently and concisely summarize the collected information. The descriptive data collected for the O. pumilio frogs and their respected locations are represented below in Tables 2 and 3. Table 2: O. pumilio descriptive statistics, Elphick, 2016 Location Gender N Valid Missing 0 0 Mode a Sum a. Multiple modes exist. The smallest value is shown Table 3: O. pumilio location frequencies, Elphick, 2016 Frequency Percent Valid Percent Cumulative Percent Tree Leaf Ground Total
11 With the information utilized in this study consisting of nominal data, expanding upon the many aspects of descriptive analysis proves challenging. Nevertheless, upon examining the central tendency measures affixed with the data in Table 2, it can be seen that the mode is 3.00, which throughout testing represents the ground. Taking the modal unit into account it may be considered that perhaps no study is needed as the data shows that the frogs are most observed upon the ground. However, this value considers both genders of frogs whereas it is the locational difference between genders being evaluated. Furthermore, Table 3 shows that the ground location does not far surpass the amount of strawberry poison dart frogs observed amongst the trees. Lastly, considering the variables are comprised of gender and location of the O. pumilio frog it is clear that a median as well as a mean cannot be calculated as the variables cannot be ranked nor does an average value in the data set exist. As all other aspects beyond mode in descriptive statistics are based upon the mean, the measure of dispersion and shape also cannot be further analyzed. Due to the nature of the variables allowing only a limited analysis, Figure 3 provides an additional and potentially more beneficial representation of the data in which to infer further information as it provides a visual representation of the distribution and observed frequency of male and female O. pumilio frogs. It is evident that for both genders, frogs found on leaves were observed at the lowest frequency, while ground and tree location data was observed at two to three times larger frequencies. Though when focusing on gender, it is apparent that males were most commonly observed on the ground while females were most often found in trees. 11
12 Fig 3. O. pumilio double bar graph of distributions. Borton, Chi-Square Assumptions In proving whether or not the location of where poison dart frogs are found is dependent upon gender, determination resides upon performing a Chi-Square test on the observed data obtained within the Dixon forest, Panama. In conducting the Chi-square test of Independence three assumptions required to carry out such a test must to be met. The assumptions in this case include that the data come from a single random sample, that the variables be organized by nominal or ordinal categories, and lastly that no more than one fifth of the categories contain less than five observations. To initiate the hypothesis testing, data was collected using a single random sample of sixty-two O. pumilio frogs. Sixteen females and eleven males of which found in trees, two females and five males found on leaves, and thirteen females with fifteen males found on the ground. Due to the data being a single random sample, as discussed 12
13 previously within the methods section, it is clear that the first assumption is met. Secondly, based on the variables being both the strawberry poison dart frog s gender along with their quantifiable locations, the condition of nominal data can be easily assumed. Lastly, as only one variable category, being the leaf location, contains less than five observations it can be ascertained that this last crucial requirement is also fulfilled. For additional clarification, Table 4 below further illustrates the data discussed and assumptions verified by such. Table 4: O. pumilio observational statistics, Elphick, 2016 Gender Measure Location Total Tree Leaf Ground Male Count % within Gender 35.5% 16.1% 48.4% 100.0% % within Location 40.7% 71.4% 53.6% 50.0% % of Total 17.7% 8.1% 24.2% 50.0% Female Count % within Gender 51.6% 6.5% 41.9% 100.0% % within Location 59.3% 28.6% 46.4% 50.0% % of Total 25.8% 3.2% 21.0% 50.0% Total Count % within Gender 43.5% 11.3% 45.2% 100.0% % within Location 100.0% 100.0% 100.0% 100.0% % of Total 43.5% 11.3% 45.2% 100.0% 13
14 3.3 Chi-Square Test of Independence With the assumptions shown to be satisfied, commencing the Pearson s Chi- Square test of independence can be perceived as viable. Based on the characteristics of the study being conducted, determination will be established using a significance value of 0.05 or 5% using a two tailed analysis. Though the final outcome of the test will be decided using the p-value approach, to further verify the conclusion it is helpful to establish the critical statistic relevant to the data. With the degrees of freedom equal to two and the significance level set at 0.05, the Chi-Square table provides a critical statistic value of Furthermore, as it is the independence of the O. pumilio frog s location contingent upon gender being investigated, the null and alternative hypothesis in this case are as follows: Ho: The locations of sighted O. pumilio frogs occurs independent of their gender Ha: The locations of sighted O. pumilio frogs are not independent of their gender With all prior necessary steps taken in order to perform the hypothesis test, the O. pumilio data calculated through SPSS provide the below statistics in Table 5. Table 5: Chi-Squared Test Results, Elphick, Value Degrees of Freedom Asymp. Sig. (2-sided) Pearson Chi Square Likelihood Ratio Linear-by-Linear Association N of Valid Cases
15 As it is the Pearson test of independence being conducted, the results being considered will entail the Pearson Chi-Square row of values as they correspond with the hypothesis test at hand. Furthermore, as standard with hypothesis testing, if the p-value is greater than the value of alpha, one would accept the null hypothesis. With a calculated p-value of.308 seen in Table 5 as the Pearson s Chi-Square Asymp. Sig. it can determine to not reject the null in the case of independence concerning O. pumilio frog s gender and their identified locations. This conclusion can be inferred from the prior established alpha value of.05 being less than the p-value of.308. To further ascertain this conclusion one can look to the critical value and test statistic of and respectively, additionally confirming the stated conclusion. In decisive summation of the Pearson s Chi-Square test of independence it can be deduced that at the 5% level of significance there is sufficient evidence to support the null hypothesis and that the locations of sighted O. pumilio frogs does in fact occur independent of their gender. 4.0 Conclusion 4.1 Discussion In concluding that locations of sighted O. pumilio is independent of gender, possible solutions for this occurrence should be examined. Based on the data collected, we concluded that there is no association between gender and where the frogs are found, leading us to analyze why both genders of the frog are distributed the way they are. Our samples show that male and female frogs were both found more often in trees and on the ground then on leaves. This could potentially be due to leaves representing far less surface area within O. pumilio s home range than trees or ground. It is interesting to note 15
16 however, that while frog location was independent of gender, both male and female frogs appeared to occupy the same locations in relatively similar frequencies. This could potentially be explained by how male distributions are influenced by female distributions, often times with males being drawn to areas of high female density (Meuche, Linsenmair & Prohl, 2012). To better understand why gender and locational distributions are situated the way they are, it is important to analyze parental responsibilities as well as reproductive and behavioral characteristics of the frog. Location of frogs may be the result of many different stimuli. It may be related to gender, which would also help explain why frogs are more frequently found at these sites. This could help explain the duration male frogs spend courting females or the duration female frogs spend transporting and caring for their tadpoles. Frogs use bromeliads axils to deposit their tadpoles and when they return to feed their young. Therefore, it would be a feasible hypothesis to believe that female frogs spend more time situated near bromeliads than male frogs. However, frog location may be due to a predator response or feeding, both, which are independent of gender. If behaviour was documented, it may help explain why a frog is perched in a specific area. A better understanding of O. pumilio home range and territory may also help us better understand perch location. In total, male and female frogs were most often found on ground locations. This concept could potentially be explained by some of the parental responsibilities the frog possesses. Both sexes are involved with aspects of rearing young O. pumilio individuals. Male and female frogs consistently return to tadpole rearing sites, often times located on the ground next to small bodies of water, to feed young tadpoles nutritive material (Murasaki, 2010). Consistent visitation to ground locations could potentially explain why 16
17 many of the frogs were found on ground locations. Another reason why both sexes were commonly found on the ground could be due to reproductive factors. Oviposition sites are mainly located within the dry leaf litter on the ground (Meuche, Linsenmair & Prohl, 2012). If the time frame happened to line up with the time courtship activities were taking place between male and female frogs, the higher frequency ground observations make sense, as both genders would be spending more time on the ground. Lastly, another reason why ground observations were higher may have to do with the fact that O.pumilio frogs require consistent moisture to survive. Many ground locations, such as beneath the leaf litter or within the roots of trees provide such ideal locations (Meuche, Linsenmair & Prohl, 2011). Perhaps frogs of both genders were found more often on the ground, as they were seeking refuge from hot, dehydrating conditions. Leaves were likely selected for the least by both genders due to their lack of benefits for the frog. At times, it has been noted that females will raise tadpoles in the axils of heliconias and other plants (Meuche, Linsenmair & Prohl, 2011). However, if there was a more suitable body of water within our sampling location, it could be suggested that the frogs would aggregate there instead. While gender was concluded to be independent of location, female frogs were still found at the highest frequency when located on trees. It has been suggested that female frogs move much more frequently within their home range than do male frogs (Murasaki, 2010). Furthermore, movement within female O. pumilio home range is typically to accomplish food collection or to access refugium (Meuche, Linsenmair & Prohl, 2012). A common food source in which O. pumilio feeds upon are ants, which are typically found in permanently humid microhabitats. (Meuche, Linsenmair & Prohl, 2011) Such microhabitats can be found 17
18 around tree areas and could present a solution on why female distributions are situated near trees. Another reason could be female frogs occupy tree sites as a way to escape unfavorable conditions. This in turn could explain why male distributions were lower around trees, as male frogs tend to remain more sedentary within their home range (Murasaki, 2010). Perhaps the added energy needed to ascend trees are not worth the benefit occupying tree sites provides. 4.2 Problems with current research The observed location variable in this study vaguely describes O. pumilio s habitat. In tropical rainforests, the ground and trees both create their own microhabitats. Better variables could have been used such as observations below 1m, between 1-3m and above 3m. Large trees such as ficus trees where O. pumilio frequent are often over 100 tall and the trunk buttress can be over 12 wide. Frogs use trees for calling and to transport tadpoles to water sources such as bromeliads. The ground offers a microhabitat with leaf litter for them to lay their eggs, fallen trees and a wide variety of vegetation to hide and feed on small insects. 4.3 Research Improvements Further research should be conducted in the region and sampling frequency should be increased to allow for in order to gather a larger data set. A larger sample size is more representative of the population and it limits the risk of outliers. More plots could be established in the Dixon forest and analyzed. Nearby islands could be surveyed in order to compare results between different populations within the archipelago. Radio telemetry would allow for researchers to monitor the frequency of time that a frog spends at each location. Telemetry has been used in the past to research terrestrial 18
19 herps, including frogs. Small radio transmitters have been used to monitor O. pumilio successfully (Murasaki, 2010). Recently internal radio transmitters have been successfully used to study the home range of Hyla versicolour (Johnson et al, 2007). This would allow for more consistent and accurate data collection as many observations are missed during visual surveying (McGarrity et al, 2016). With the use of these transmitters, researchers could monitor long term surveys, in order to determine why frogs choose these perches. Observation of O. pumilio behaviour could also be documented, such as feeding, courting (calling) or tadpole transporting. Recording behaviour may give biologists a better understanding of why frog gender may be related to perch location. Telemetry would also help gain valuable data on territory size and the dispersal of metamorphosed tadpoles. However, due to the small size of these frogs, battery lives will be greatly reduced and will have to be serviced more frequently. 19
20 5.0 Literature Cited Galeano, SP., Harms, KE Coloration in the polymorphic frog Oophaga pumilio associates with level of aggressiveness in intraspecific and interspecific interactions. Behavioural Ecology and Sociobiology 70(1): Institute of Tropical Ecology and Conservation. Bocas del Toro Biological Station. Map 2. Lund, M. & Lund, A. (2013). Testing for Normality using SPSS Statistics. Retrieved from McGarrity, Monica, Johnson Steven, Radio telemetry study of invasive Cuban tree frogs. Gulf Coast research and Education Center, University of Florida. McGrew, Jr. C., Lembo, Jr. A., & Monroe, C. B. (2014). An Introduction to Statistical Problem Solving in Geography. Long Grove, IL: Waveland Press. Meuche, I., Linsenmair, KE., Prohl, H Female territoriality in the Strawberry Poison Frog (Oophaga pumilio). Copeia (3): Meuche, I., Linsenmair, KE., Prohl, H Intrasexual competition, territoriality and acoustic communication in male strawberry poison frogs (Oophaga pumilio). Behavioural Ecology and Sociobiology 66(4): Murasaki, S Sex specific patterns of movement and space use in the strawberry poison frog, Oophaga pumilio. FIU Electronic Theses and Dissertations. Paper 226. Murasaki, Sex Specific Movements and Space use in Strawberry Poison Dart Frog Oophaga pumilio. Florida International University. Nowakowski, AJ., Jimenez, BO., Allen, M., Diaz-Escobar, M., Donelly, MA Landscape resistance to movement of the poison frog, Ophaga pumilio, in the lowlands of northeastern Costa Rica. Animal Conservation 16(2):
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