Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID -(http://jwb.araku.ac.ir/) Research Article DOI: 10.22120/jwb.2019.115621.1099
Faunistic diversity and distribution of Wolf spiders (Lycosidae: Araneae) in Western and Northern Mindanao, Philippines
and sampled specimens usually clumped in 1* Philip Noel O. Banaag , Olga M. agroecosystems near water bodies. Generally, Nuñeza1, Aimee Lynn B. Dupo2, Myla Shannon-Weiner values were found to be very R. Santiago-Bautista3 low (H’= 0 to 1.29) but tend to exhibit higher values (H’= 1.23 to 1.29) in sites with field 1* Department of Biological Sciences and Premier margins that are located near streams or forest Research Institute of Science and Mathematics, patches. Results indicate the importance of Mindanao State University- Iligan Institute of riparian areas and ecotones for the diversity of Technology, Iligan City, Philippines, 2Institute of Biological Sciences, University of wolf spiders. the Philippines Los Baňos, Laguna, Philippines, Keywords: Ecotones, Lycosid, Philippines, 3Department of Biochemistry, Faculty of Pharmacy University of Santo Tomas; UST species richness, Wolf spiders. Research Center for Natural and Applied Sciences; and the Graduate School, University of Introduction Santo Tomas, Manila, Philippines With over 2,300 described species (Platnick *email: [email protected] 2005), the family Lycosidae (wolf spiders or Received: 11 October 2019 / Revised: 28 October2019 / Accepted: lycosids) is one of the most abundant spider 11 November 2019 / Published online: 07 January 2020. Ministry families in the world. Described with four of Sciences, Research, and Technology, Arak University, Iran. distinct morphological characters such as eye Abstract arrangement, absence of retro lateral tibial Despite being one of the most abundant spider apophysis on the male pedipalp, egg sac carried families in the world, wolf/lycosid spiders are on the spinnerets of females, and young carried poorly studied in the Philippines. In this study, on specialized setae on the dorsal surface of the we determined the faunistic diversity and mother’s abdomen, the said spider group tends distribution of lycosid spiders from nine to exhibit monophyly (Dondale 1986, Griswold sampling sites in Western and Northern 1993). Mindanao. An opportunistic sampling method The abundance of the group is attributed to the was used to collect samples. Non-parametric abovementioned morphological features estimators and GIS mapping were utilized in particularly the female traits. The female’s this study to determine richness (genus level) ability to carry its young on the back enables and distribution. Results showed that the the group to thrive in areas with high accumulation curves of the observed richness anthropogenic disturbance such as and non-parametric estimators did not reach an agroecosystems, grasslands, coastal, and asymptotic value suggesting that the true riparian zones (McKay 1974, Manderbach and richness of the sampling areas is much higher Framenau 2001, Framenauet al. 2002, Morse than the estimated values. Six genera were 2002). Dispersal methods of the early instars documented in which one genus is a new of some member species such as ballooning is Mindanao record and another one is new to the also very influential in making the lycosid Philippines. Distribution results showed that spiders virtually abundant in every terrestrial fewer specimens are found in forested areas habitat (Murphy et al. 2006) by eliminating www.SID.ir 41| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID some of the problems caused by geographical generic diversity and distribution of wolf factors (Richter 1970, Greenstone 1982, spiders in selected areas of Western and Greenstone et al. 1987, Crawford et al. 1995, Northern Mindanao. Edwards and Thornton 2001). Some lycosid species are also considered as possible Material and Methods bioindicators of organophosphates due to their Study Area This study was carried out in nine sampling abundance in agro-ecosystems (Hodge and sites on the island of Mindanao (8.4961° N, Vink 2000, Van Erp et al. 2000). 123.3034° E) particularly on the Northern and Like all spiders, lycosids are predatory in Western regions (Fig. 1). Geographically, the nature and most of the species belonging to island of Mindanao is located below the lycosid groups do not build webs but utilize a Visayas and is considered as the second largest “sit-and-wait” predatory behavior by building Philippine island. Of the nine sampling sites burrows (Kronk and Riechert 1979, Vink established in the two regions, five can be 2002). Some member species are also vagrant found in the provinces of Bukidnon, two in predators in which they actively hunt for prey Zamboanga del Sur, and two on Camiguin (Murphy et al. 2006). In some agroecosystems Island. Descriptions of each sampling site are like rice fields, hunting species such as shown in table 1. Pardosa pseudoannulata can regulate population densities of leafhoppers and Sampling and identification of specimens
planthoppers (Barrion 2001). Due to their Intermittent field sampling was conducted from predatory behavior, lycosid venom peptides are August to December 2017 in the two regions. also considered as pharmacologically The sampling method was mainly opportunistic important. One species, Lycosa singoriensis using the vial tapping method and modified can initiate hemolysis of human erythrocytes from Coddington et al. (1991), Sørensen et al. (Liu et al. 2009). The possible importance of (2002), and other existing spider diversity lycosids (wolf spiders) in agriculture and studies in the Philippines (Patiñoet al. 2016, medicine is seemingly a good reason to Dacar and Nuneza 2016, Elias and Nuñeza conduct ecological studies about this spider 2016). Specimens were collected for a total of group. 81 person-hours. Specimens were then In the Philippines, ecological studies that are preserved in 5ml screw-capped plastic vials solely focused on wolf spiders are fairly filled with 95% ethanol. Each vial containing a limited. The most recent studies regarding the specimen was labeled with information such as diversity and distribution of wolf spiders are by locality, sex of the specimen, and habitat. Barrion and Litsinger (1981a, 1995) and Identification of the specimens up to the genus Barrion (2001). However, the studies were only level was carried out at the University of the focused on rice field ecosystems leaving other Philippines Los Baños and voucher specimens areas such as forest patch and stream were preserved in the Mindanao State ecosystems partially undocumented. Mindanao University – Iligan Institute of Technology. islands situated within deep-ocean channels like Camiguin were also not documented. The GIS Mapping and Abundance advent of publicly accessible mapping soft To visualize the relative abundance and wares or methods such as GIS mapping and evenness of each genus in all sampling sites, a non-parametric estimators could provide some rank-abundance curve was generated (Fig. 4). useful data to create a form of baseline GIS Mapping was utilized to determine the information regarding the ecology of wolf distribution of the specimens. spiders in the Philippines. As such, the general aim of this research was to determine the www.SID.ir 42| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID Statistical analysis determine and visualize the abundance and To determine the diversity of a certain species, evenness of the collected specimens. Finally, one must first calculate the richness Shannon-Weiner Diversity Index (H’) was (Garcianoet al. 2014, Quiñoneset al. 2016, utilized to determine the diversity of lycosid Maandiget al. 2017) and determine the spider genera per sampling site. H’ index was abundance and evenness of the said species calculated using the Paleontological Statistics (Stirling and Wilsey 2001). To measure Software Package for Education and Data richness, accumulation curves and non- Analysis (PAST) version 3.14 (Hammer et al. parametric estimators were utilized in this 2001). The Shannon-Weiner Diversity Index study. Asymptotic curves or values in the (H’) is commonly used in diversity studies accumulation curve are possible indications (Krebs 1989). The computed H’ index was then that the species inventory is already complete interpreted based on a scale modified from (Cardoso et al. 2008, Responteand and Nuñeza Fernando Biodiversity Scaling System 2016). Non-parametric estimators that were (Fernando 1998). In a modified Fernando commonly used in related literature include Biodiversity Scaling System, H’ values Chao1, Chao2, Jackknife1, Jackknife2, and between 3.5 and 4.0 are considered to be very Bootstrap (Colwell 1999, Castanheira et high, 3.0-3.49 are considered to be high, 2.5- al.2016). EstimateS, richness estimator 2.99 values are moderate, 2.0-2.49 values are program, version 9.1 (Colwell 2013) was used low and lastly, H’ values ranging from 1.99 and to calculate the estimator values. A rank- below are considered to be very low. abundance curve was also generated to
Figure 1. GIS map of the sampling sites in Northern and Western Mindanao. Sampling sites are represented by color-coded markers. The image at the top left corner shows the location of Mindanao in the Philippine Archipelago. www.SID.ir 43| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID Table 1. A detailed description of the sampling sites based on vegetation type, dominant or common plantspecies distance to nearest body of water, and anthropogenic disturbances observed in each site.
Distance Site Dominant/Common to nearest Anthropogenic Site Name Vegetation Type Number Plant species water Disturbances bodies Sugarcane and Gmelina sp., Musa sp., Sto. Domingo, Situated cassava Secondary Schismatoglottis sp., 1 San Fernando, along a plantations (20 Forest/Agricultural Homalomena sp., Bukidnon stream. meters from the Artocarpus sp. site). (Center for Ecological Homalena sp., Situated Public swimming Development Primary/Secondary Schismatoglottis sp., 2 along a pool (10 meters and Recreation), Forest Rattan sp., Heliconia stream. from the site). Impasug-ung, sp., Shorea sp. Bukidnon 50 meters Kisolon, Cocos nucifera and National Highway from 3 Sumilao, Agricultural Musa sp. (25 meters from Alalum Bukidnon Cogon Grass the site). Falls 600 meters from a Spring onion and Mt. Kalatungan, Allium porrum,Zea sp., stream 4 Agricultural corn plantations, Bukidnon cogon grass located in hiking trails. the Primary Forest Bangcud, Corn and Sugarcane Adjacent to Pumping Station, 5 Malaybalay, Agricultural (Saccharum sp.) Matin-aw plantations Bukidnon Carabao grass spring. Calumanggi, 30 meters National Highway Dumingag, 6 Agricultural Rice, carabao grass from a (10 meters from Zamboanga del stream the rice field). Sur Bobongan, 10 meters Ramon Cocos nucifera, carabao from a Fishponds and 7 Magsaysay, Agricultural grass, cassava, corn dried-up cornfields Zamboanga del stream. Sur Nearby spillway Pandan, Cocos nucifera, Ficus Adjacent to and cement road 8 Mambajao, Secondary/Agricultural sp., Gmelina sp., a stream. to Katibawasan Camiguin carabao grass Falls. Baylao, Mimosa sp., Paspalum 20 meters Abandoned 9 Mambajao, Agricultural sp., Tree ferns, Gmelina from a cornfield, cement Camiguin sp. stream road.
Result Of the 181 adult specimens collected, six Venonia, and Draposa were not mentioned or genera were identified (Table 2). These are described in previous Philippine ecological or Artoria, Draposa, Hippasa, Pardosa, Venonia, taxonomic publications hence they can be and Wadicosa. Genus Hippasa and Genus considered as new records from the Pardosa were previously recorded and Philippines. The genus Artoria was mentioned described by Barrion and Litsinger (1981a, by Barrion and Litsinger (1995) however it was 1981b, 1995). On the other hand, Wadicosa, only recorded in Luzon Island. This is the first
www.SID.ir 2| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID record of the genus Artoria in Mindanao Island. and 12.15 % adult females and adult males A pie chart was also generated to represent the respectively). sex ratio of the collected specimens (87.84 %
Table 2. List of the collected and identified lycosid genera and their respective individual counts per sampling site (S1-S9)
Genera S1 S2 S3 S4 S5 S6 S7 S8 S9 Total Genus 0 8 0 0 0 0 0 0 0 8 ArtoriaThorell, 1877 Genus Draposa 0 0 0 0 0 1 0 6 0 7 Kronestedt, 2010 Genus Hippasa 3 0 0 4 0 3 7 0 3 20 Simon, 1885 Genus Pardosa C. L. Koch, 1847 22 0 8 18 4 11 28 2 4 97 Genus Venonia 4 0 0 0 0 0 0 0 0 4 Thorell, 1894 Genus Wadicosa Zyuzin, 1985 5 1 0 11 6 6 0 7 9 45 Total per site 34 9 8 33 10 21 35 15 16 181
Richness estimators automatically suggest to the user to consider Nonparametric species estimators particularly the larger Chao2 value if the estimated CV of Chao1, Chao2, Jackknife 1, Jackknife 2, and the abundance or incidence distribution is Bootstrap were utilized in this study to measure greater than 0.5 (Colwell 2013). On the other the richness of each genus from all sampling hand, the results showed that the highest sites (Fig. 2). To visualize the estimated values, estimate of species richness is from Chao2 at accumulation curves for each estimator were 25.89 or 26 genera followed by Jackknife2 at generated along with the accumulation curve of 8.65, Jackknife1 at 7.78, bootstrap at 7.69 (14), the observed richness, which was 6. The larger and the lowest is Chao1 at 6. The generated value for Chao2 was utilized in the results since accumulation curves failed to reach an the CV (Coefficient of Variation) of the asymptote and seem to be rising as well. abundance or incidence distribution of the However, Jackknife2 seems to approach provided data is equal to 0.534. EstimateS asymptotic values.
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Figure 2. Accumulation curves for the observed richness [S(est)] and richness estimators (Chao1, Chao2,Jackknife 1, Jackknife 2, and Bootstrap).
Abundance Pardosa is the most abundant with a relative The abundance curve exhibited a steep gradient abundance of 53.59%. This is followed by implying a low evenness between genera with Wadicosa and Hippasa at 24.86% and 11.04%, higher-ranked genera such as Pardosa respectively. Artoria is at 4.41% and Draposa exhibiting a much larger abundance gap at 3.86%. The least abundant genus was compared to lower-ranked genera. The genus Venonia at 2.20%.
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Figure 3. Rank-abundance curve of the observed Lycosid genera found in Western and Northern Mindanao. The steep gradient indicates a low evenness.
Distribution grasslands or rice fields situated near Distribution maps based on the genus were freshwater bodies. Genus Wadicosain this generated through GIS mapping. The study was found in Sites 1, 2, 5, 8, and 9 which distribution map of Genus Pardosa (Fig. 5) are situated along or beside a stream. Venonia indicates that the representative specimens of specimens were only collected in Site 1. the said genus can be found in all sampling sites except in CEDAR (S2) and Baylao, Species Diversity Mambajao (S9). The distribution map of the Table 3 shows the observed richness and said genus also suggests that the bulk of its Shannon-Weiner Diversity values for each sample population can be found in Sto. sampling site. Based on the Modified Fernando Domingo (S1), Mt. Kalatungan (S4), and Biodiversity Scale, the Shannon-Weiner Bobongan (S7). On the other hand, all diversity indices of all sampling sites are very specimens of the Genus Hippasa were low (1.99 and below). The highest Shannon- observed to be distributed in agroecosystems as Weiner value is from Site 6 with H’ = 1.29 shown in Fig. 4. The bulk of the sample followed by Site 1 with H’ = 1.23. The highest population of Hippasacan also is found in S7 richness value (number of recorded genera which is a coconut plantation. The distribution per/site) can also be attributed to both sites at S of Artoria seems to be limited only to CEDAR. = 4. The lowest Shannon- Weiner value is from The sampled Draposa specimens, on the other Site 3 with H’ = 0. This is expected since only hand, are distributed on habitats such as one genus was recorded in this site.
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Figure 4. Distribution map of genus Artoria, Draposa, Venonia(a) and Hippasa (b).
Specimens are represented by color-coded dots. The larger the marker, the greater the abundance range. size of the dots represents the abundance (in range) The same set of legends are also used for Fig. 5. of the collected specimens in a specific site. The
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Figure 5. Distribution maps of Genus Pardosa (a) and Wadicosa (b)
Discussion about singletons which can affect diversity Richness indices and species richness. No singletons Results imply that the species/genus inventory were recorded (in relation to genus) in this is not yet complete and it is also probable that study but some results of this study are the richness of the sampling areas is much seemingly similar to the results of Coddington larger than the estimated richness generated in et al. (2009) particularly the presence of this study. Possible reasons for the occurrence female-based sex- ratio bias. A study by of the above-mentioned estimator values Vollrath and Parker (1992) which was also include cryptic habitat, biased sex ratio, mentioned in Coddington et al. (2009) stated clumped distribution, and small size which is that adult male spiders, particularly wandering based on a study by Coddingtonet al. (2009) males, exhibit higher mortality rates than www.SID.ir 49| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID females, hence the higher occurrence of values of this study. Morphological females which is again, a plausible reason for conservatism is a considerable deterrent in the occurrence of the resulting estimator values. identifying spider assemblages particularly in However, Coddington et al. (2009) also pointed lycosids (Bond et al. 2001, Vink 2002). out that the abovementioned reasons cannot Philippine species such as Pardosa daniloi, P. affect the frequency of singletons and in effect asacayi, and Pardosahawakana are previously the occurrence of the generated estimator described as separate species and were values in this study. According to Coddington reclassified as synonyms of P. et al. (2009), the main culprit for negative magkasalubonga (Barrion and Litsinger 1995). richness bias is undersampling. The mentioned Framenau (2006) then transferred the same study along with other previous studies like species to the genus Venatrix. A study about Walther and Morand (1998) and Mao and trapdoor spiders by Bond et al. (2001) suggests Colwell (2005) showed that non-parametric that using morphology-based species concepts species estimators can only properly measure to identify spiders can possibly affect our the true species richness of a given data set if understanding of spider evolution and diversity. the observed species of the said data set reach Habitat utilization could also be a factor but 66.6% or 80% threshold. This means that the needs further information (Elias and Nuñeza current study needs a fairly large amount of 2015). Despite the conclusions of Coddington samples or a greater sampling intensity to et al. (2009), factors like morphological generate realistic species/generic estimates. But conservatism and female based sex-ratio bias one could still speculate for other possible could possibly still affect species richness or reasons. Morphological conservatism can also diversity but further studies should be be used to discuss the generated estimator conducted to draw more accurate conclusions.
Table 3. Observed richness and Shannon-Weiner Diversity (H’) values in the nine sampling sites of Western and Northern Mindanao
Sampling Site Observed Richness Shannon-Weiner (H’) 1 4 1.23 2 2 0.63 3 1 0 4 3 1.06 5 2 0.69 6 4 1.29 7 2 0.64 8 3 1.08 9 3 1.09
Distribution and Abundance be probably attributed to its resistance to some Except for CEDAR, all sampling sites are pesticides. A study by Tahir et al. (2016) considered as agroecosystems or buffer zones. suggested that when compared to susceptible Genus Pardosa, particularly P. asumatrana, are (control) groups, resistant groups of P. known to dominate (in terms of abundance) sumatrana contained higher levels of non- agroecosystems when compared to other specific esterases and monooxygenases in species as confirmed by similar studies like the which both enzymes play a role in metabolic assessment of spider fauna in Punjab, Pakistan resistance. However, the Pardosa specimens by Mukhtar et al. (2012). The seemingly high from Sto. Dominggo seems to be different from abundance of Pardosa in agroecosystems can the rest. Specimens from Sto. Dominggo was www.SID.ir 50| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID collected from a stream within a Secondary species were found in forested sites like forest patch and the morphology of the said CEDAR. Most of the sampled species seem specimens seems to be different from were found to prefer open areas and buffer specimens found in grasslands and fields. zones situated near bodies of freshwater such as However, further taxonomic and molecular streams and rivers. analyses are required to confirm this. Results Diversity indicate that Hippasa specimens are heavily The very low diversity values can also be distributed in agroecosystems. A web-building attributed to the low richness since it can affect genus, Hippasa is considered being commonly diversity indices (Stirling and Wilsey 2001). found in agroecosystems as stated by Barrion Another possible reason is that most of the and Litsinger (1995). A representative species, sampling sites are agro-ecosystems. Due to Hippasa holmerae, was also considered as a high anthropogenic disturbance, agro- predator of leafhoppers in rice agroecosystems ecosystems tend to yield low diversity values (Barrion and Litsinger 1981b). As shown in the when compared to undisturbed forest results, genus Artoria and genus Venonia were ecosystems (Bos et al. 2007). On the other only collected in a single site. Since the hand, Sites 1 and 6 exhibited the highest H’ species/genus inventory of this particular study values since they can be both considered as is considered to be incomplete, the field margins or ecotones based on their site abovementioned genera could probably be descriptions. Field margins are least productive found in other sites. Compiled results from a compared to the central field area but are review by Framenau (2002) suggested that particularly considered as a key factor for members of the genus Artoria are usually promoting wildlife in agro-ecosystems found in river banks and forest. CEDAR is an (Marshall and Moonen 2002). A study by Gallé ecological preserve and the Artoria specimens and Fehér (2006) also stated that spider were collected beside a stream situated along a diversity is high in ecotones. In Sites 1 and 6, forest. It is probable that streams located in the ecotones are characterized as streams and primary forest or forest patches are the habitat forest patches that are favorable to some preference of Artoria. More studies focused on lycosid species like Pardosa sumatrana and microhabitat preferences and utilization are Wadicosa fidelis due to the mix vegetation and again required for further confirmation. On the the presence of a nearby water source. This was other hand, the distribution of Draposa also confirmed by Mukhtar et al. (2012) in documented in this study is fairly similar to a which spider abundance was significantly study on spider faunal diversity conducted by higher in margins than in central field areas. Dhaliet al. (2017) at West Bengal, India in which the majority of Draposa specimens were Conclusion found in grasslands. Not much is known about Of the 6 genera that were observed and the habitat preferences of Genus Wadicosa but identified, Wadicosa, Venonia, and Draposa most taxonomic studies particularly Ahmed et are considered as new Philippine records. Non- al. (2014) and Kronestedt (2015) suggest that parametric species estimator values suggest Genus Wadicosa is usually found in water that the true estimate of the species richness in bodies like lakes, streams, or even in irrigated this study is much higher than the generated rice fields. This can explain why most of the estimates. The notion that female based sex- members of the Genus Wadicosa that were ratio bias and morphological conservatism as collected in this study were found in Sites 1, 2, possible factors that can affect species richness 5, 8, and 9. Generally, the results from the in given samples should be investigated further. generated species distribution maps suggest Overall results for distribution suggest that that only a handful of the sampled lycosid most of the sampled specimens are found in www.SID.ir 51| Journal of Wildlife and Biodiversity 4(1): 40-54 (2020) Archive of SID agroecosystems situated near freshwater leafhoppers and planthoppers. International bodies. In general, the Shannon-Weiner Rice Research Newsletter 6(4): 15. Diversity value of this study is considered to be Barrion A.T., Litsinger J.A. 1981b. The spider very low but is supported by the fact that most fauna of Philippine rice agroecosystems I. of the sampling sites are agroecosystems that Dryland.The Philippine Entomologist 5: tend to exhibit lower diversity values. 139-166. Diversity tends to be higher in sites such as Barrion A.T., Litsinger J.A. 1995. Riceland ecotones that are dotted with streams or forest Spiders of South and Southeast patches. Information on microhabitat Asia.Entomology Division International preferences of lycosid spiders is limited which Rice Research Institue. UK, Philippines, means that the collection of quantitative microhabitat data and GIS mapping, with pp. 700 emphasis on microhabitats, is highly Bond J.E., Hedin M.C., Ramirez M.G., Opell recommended for future studies. B.D., 2001. Deep molecular divergence in the absence of morphological and Acknowledgment ecological change in the Californian The authors would like to thank the coastal dune endemic trapdoor spider Commission on Higher Education; University Aptostichussimus. Molecular Ecology(10): of Santo Tomas Proteomics and Venomics 899–910. Group (ProVe); Department of Biological Bos M.M., Tscharntke T., Leuschner C., Zeller Sciences, Mindanao State University – Iligan M., Guhardja E., Bidin A.2007. Insect Institute of Technology; Department of Science and Technology; and the Department of diversity responses to forest conversion Environment and Natural Resources (Region and agroforestry management. Stability of IX and Region X). This research endeavor Tropical Rainforest Margins 277-294. would not be possible without the support of Cardoso P., Scharff N., Gaspar C., Henriques the above- mentioned institutions. 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