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Journal of Wildlife and Parks, 30: 1-13 (2015)

ESTIMATING (: FORMICIDAE) DIVERSITY OF TASEK BERA RAMSAR SITE, PAHANG, PENINSULAR MALAYSIA

Noor Izwan, A. & Amirrudin, B.A.*

School of Marine and Environmental Sciences, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Malaysia

*Corresponding author’s email: [email protected]

ABSTRACT

Tasek Bera Ramsar Site, Pahang is the largest freshwater swamp-lake in Peninsular Malaysia and the ecosystem is diverse. The ground-dwelling fauna at Tasek Bera Ramsar Site is still understudied as there is no checklist available to date. The objective of this study is to survey the diversity of ground-dwelling ants (Formicidae) and to provide a checklist for Tasek Bera Ramsar Site. In this study, leaf-litter samples from 1 m2 plot in five line transects were collected, and ants were taken manually. At each transect, plots were established between 20 m and 40 m in distance with a total of 16 plots. Six of ants were recorded comprising of 26 genera and 53 . Almost half of the samples collected were from the (45.3%), followed by (20.8%) and (17.0%). This finding is not surprising since Myrmicinae contains the most species in Formicidae. The most common genera recorded were (13.2%), Camponotus (7.5%), (7.5%), and (7.5%). According to the estimates of the expected species richness, leaf litter sifting and manual collection methods allowed for the collection of 76 to 175 species of ants which is different from the total observed species in this survey. This difference is possibly due to the low sampling effort and short sampling duration. Raup- Crick’s similarity index was calculated between plots resulting into four clusters. This might be due to the condition of each plot in which certain plots were in wet or dry conditions as a result from the tidal influence from Tasek Bera Ramsar Site. The finding from this study could serve as the baseline information on the Formicidae fauna at Tasek Bera Ramsar Site and can be used for further study for the conservation and management of the site.

Keywords: Formicidae, estimation, similarity, Tasek Bera Ramsar Site, ground-dwelling ants

Received (06-November-14); Accepted (24-August-15); Available online (03-November-15)

Citation: Noor Izwan, A. & Amirrudin, B.A. (2015). Estimating ants (Hymenotera: Formicidae) diversity of Tasek Bera Ramsar Site, Pahang, Peninsular Malaysia. Journal of Wildlife and Parks, 30: 1-13. 2 Noor Izwan, A. & Amirrudin, B.A.

INTRODUCTION

Ants can be found almost everywhere (Brian, 1978) and their species richness is very diverse in the tropics. Worldwide, there are about 300 genera and approximately 15,000 species of ants were described (Bolton, 1994). However, this number is very conservative. Ants are also abundant and can be easily sampled. Some groups have well-known taxonomic keys and these provide great advantages in studying them (Hölldobler & Wilson, 1990). They are well adapted to a particular site, food resources, microhabitat, and temperature regimes (Anderson, 2000). In the Brazilian forest alone, there were more ant species compared to all primate species in the world (Wilson, 1975). This shows that this tiny creature is a mega diverse group (Hölldobler & Wilson, 1990).

Many studies on ants have been conducted in Asia, including India, China and countries in the Southeast Asia region including Malaysia. However, the high temperature of the tropical forests makes it the least favourable place to study ants albeit containing more than 2,000 species (Hölldobler & Wilson, 1990). Studies on ants’ diversity in Peninsular Malaysia are rather limited. Only a few studies on their ecology have been done so far such as in Pasoh Forest Reserve, Negeri Sembilan which successfully recorded 427 species from nine subfamilies (Malsch et al., 2003). In another study, 78 species of ants were recorded at Sungai Ber, Gua Musang, Kelantan (Bakhtiar & Maryati, 2008). A less comprehensive study was conducted at Melawi Beach, Bachok, Kelantan wherein 28 species from 21 genera of ants were recorded (Nurul Ashikin & Rosli, 2010). Recently, a research was done at Temenggor Forest Reserve in which 211 ant species from nine subfamilies were recorded (Nur-Zati et al., 2011). In general, ant diversity in Peninsular Malaysia is very much underestimated. This study aims to explore and examine the diversity of the ant fauna living in the inundated freshwater area of Tasek Bera Ramsar Site (TBRS), Pahang. We hope this study could provide some inputs on Formicidae diversity especially in Peninsular Malaysia and fill the knowledge gap on ant diversity from such habitat type.

METHODOLOGY

The survey was conducted at TBRS, Pahang from 13th–16th August 2014 in conjunction with the 2014 Inventory Programme held by the Department of Wildlife and National Parks (DWNP). TBRS is part of freshwater swamp-lake system located in the southwest of Pahang (Figure 1). This wetland area is also the largest freshwater swamp in Peninsular Malaysia and protected under the Ramsar Convention since 1994 (Mokbolhassan, 2014). Ants diversity of Tasek Bera 3

Figure 1 Map showing the TBRS, Pahang. The red rectangle shows the DWNP inventory field site where the ant survey was conducted. Inset is the map of Peninsular Malaysia showing the location of TBRS (black triangle).

In this study, surface leaf-litter within 1 m2 plot in five out of six transect lines were collected and examine for ant fauna (Figure 2). Six transect lines were established by the DWNP inventory team on both sides of the access road to the TBRS jetty, just before the DWNP TBRS office. At each transect line, two to four plots were established with 20 m to 40 m gap. A total of 16 plots was established and leaf- litter sifting was thoroughly done to collect ant samples. Manual technique was also employed within the plot to increase data collection. The data from leaf-litter sifting and manual collection were combined and analysed to determine the ant diversity. Ants’ samples were placed in a vial containing 70% alcohol and were brought back to the laboratory for sorting, pinning, and identification. Keys for ants’ identification were based on Fayleet al. (2014) and Hashimoto (2003) but some taxonomic revisions within the subfamily (Brady et al., 2014) and Myrmicinae (Ward et al., 2014) were noted. Identifications were made to the lowest possible taxa. We also referred to two ant websites, Antweb (Antweb, 2014) and antbase.net (Pfeiffer, 2014) for further details.

To estimate species richness, present-absent data were used. The data were analysed using non-parametric, incidence-based estimators of ICE, Chao2, Jack1 and Jack2 in EstimateS v.9 (Colwell, 2013) to estimate species richness 4 Noor Izwan, A. & Amirrudin, B.A.

Figure 2 A schematic diagram (not to scale) of the location of the five transect lines wherein two to four plots were established to sample ant fauna at TBRS, Pahang. Transects A and D were located nearby swamp areas while transect B was not surveyed. and determine which estimator are the best to estimate the diversity of ants at TBRS. The relationships between ant assemblages and plots were then measured based on their similarity (Raup-Crick’s similarity measure) using PAST software v.2.17 (Hammer et al., 2001).

RESULTS

A total of six subfamilies (, Dorylinae, Formicinae, Myrmicinae, Ponerinae, and ) comprising 26 genera and 53 species of ants were recorded from five transect lines set up in inundated areas at TBRS. The most abundant subfamily was Myrmicinae (45.3%), contributing almost half of the verified species followed by Formicinae (20.8%), Ponerinae (17.0%), Dorylinae (7.5%), Pseudomyrmecinae (5.7%), and Dolichoderinae (3.8%) (Figure 3). The most diverse ant recorded was Crematogaster with six species followed by four species for Camponotus, Carebara and Tetramorium each, three for Pachycondyla and Tetraponera and two species each for , , , , Leptogenys, , , , and . Meanwhile only one species was recorded for each of Ants diversity of Tasek Bera 5 Number of species Number of spedes

Figure 3 Composition of ant subfamilies in TBRS, Pahang. the genus , , , , , Odontomachus, Odontoponera, Oecophylla, , , and (Appendix 1).

The estimated species richness of ant fauna at TBRS was high compared to the observed curve and none of the estimated curves showed any signs of reaching an asymptote. Figure 4 shows the species accumulation curve (Sobs-Mao Tau) and the species estimation curves generated using EstimateS. The Chao2 estimator curve was deemed to be the best among others incidence-based estimator such as ICE, Jacknife 1 and Jacknife 2. The species accumulation curve showed no indication of levelling off. Estimation from the Chao2 estimator suggested a total of 106 species of Formicidae at TBRS but only 53 species were recorded in this survey. The estimated proportion of unsampled potential species was between 30.3% and 69.7%. The unique species recorded more than half of the total species collected (39 of 53 species) while 12 more species were duplicates (see inset of Figure 4). The unique curve did not indicate any tendency to decrease and the duplicates curve still increasing indicate that sampling was far from complete.

Finally, the clustering analysis was used to examine plot similarity. Figure 5 shows that four main clusters were established at TBRS, based on the presence of ants at each plot. However, we did not examine the physical environment 6 Noor Izwan, A. & Amirrudin, B.A. Species accumulation based on observed species and Chao 2 estimator curves for ant fauna collected at the study site. Observed study site. the at collected fauna curves for ant and Chao 2 estimator based on observed species accumulation Species Figure 4 Figure unique (73.6%) and 106 species of ants can be found in the study site. Inset is graph total 53 while Chao 2 estimated species recorded duplicates (22.6%) species that were recorded in this survey.

Ants diversity of Tasek Bera 7 Similarity

Figure 5 A dendrogram showing the similarity among ants assemblages found at 16 plots in TBRS. The cluster analysis of paired group algorithm was measured using Raup-Crick’s method with 0.79 cophenetic coefficient indicates how well the dendrogram reflects the similarity. The dash-line at 35% similarities separated the ant assemblages into three major groups indicate by respective colours. Black line colour indicates the most similar assemblages in this clustering.

of each plot. At 35% of similarity, the plots were grouped into three different clusters. Plots 1 and 4 formed a distinct cluster in the first splitting point. Plots 10 and 16 shared the most similar species with 87% similarity while plots 10, 12, 14, and 16 were clustered together and its share almost similar species. On the other hand, plot 3 and 6 recorded the lowest similarity between the pair with less than 40% similarity.

DISCUSSION

In this study, Myrmicinae was found to be the most dominant subfamily of ants at TBRS where almost half of the total species recorded (45.3%) were 8 Noor Izwan, A. & Amirrudin, B.A. from this subfamily. Subsequently, Formicinae contributing about 20.8% of the species recorded. The result was not surprising because Myrmicinae is the most diverse and largest subfamily in Formicidae with more than 130 genera and 6,000 species in the world (Bolton, 2014). For this reason, these two subfamilies dominated many checklists of ants fauna of the region, e.g., in Borneo, these two subfamilies covered almost two third of the Formicidae species recorded (Pfeiffer et al., 2011). However, the methods used in this survey were biased towards the two subfamilies in which many species within both subfamilies are ground dwellers. For other subfamilies such as Dolichoderinae and Pseudomyrmecinae, many of the species inhabits lower vegetation stratum such as tree barks, twigs and decaying woods (Malsch et al., 2003) which were missing in the plots studied and thus were less recorded. In this study, manual collection yielded the highest number of species from the subfamilies of Formicinae (Camponotus spp.1, Camponotus spp.2, Camponotus spp.3 and Camponotus gigas), Pseudomyrmecinae (Tetraponera spp. 1, Tetraponera spp. 2 and Tetraponera spp. 3) and Dolichoderinae (Dolichoderus cuspidatus and Dolichoderus sp.) suggesting that multiple methods are necessary to obtain a good representative of species richness of ants. Therefore, we would urge that future works need to employ more variety of collections gears in to obtain more species. Therefore, the findings from this study should be considered as preliminary and any comparison in the future should take this factor into account.

The estimated species richness calculated in this study was higher than the observed species number with an estimated of between 30.3–69.7% of the unsampled species. This shows that the sampling effort was inadequate to represent the complete Formicidae fauna in TBRS. This led to the collection of high number of unique species (73.6%) but low duplicates species (22.6%). Such findings will impinge the accuracy of species richness estimation of the ant fauna of TBRS. Among the incidence-based species richness estimators, Chao2 depends on the unique (species that only being found once in samples) and duplicate species (species that were recorded twice in samples) to estimate species richness and is reliable to be used to determine the local ant diversity. Chao2 estimator performed better when more less species are caught during sampling than common species. If more duplicate species were observed, this will lead the estimator to reach an asymptote. On the other hand, if the unique species are higher, more sampling or repeated samplings are needed to reduce the unique species. Thus, longer sampling time and greater sampling efforts may overcome this shortcoming.

The clustering analysis paired the most similar neighbour together followed by the next pairs (when the value of the paired similarity index reaching 1) (Pielou, Ants diversity of Tasek Bera 9

1984). In this study, the most similar plots were plot 10, 12, 14, and 16 which shared almost 80% similarity. These plates consist of the same species and have a similar environment condition and food resource which promotes similar species to congregate (Magurran, 1988). In general, most of the paired plots share high similarity (> 50%) except for plots 3 and 6. This could be contributed by the nature of the study area which occasionally inundated by high water level of the swamp-lake especially during wet season. Such event only permits a number of ants groups to flourish and these may come from a limited number of subfamilies or even from specific genera. We strongly believe that the ant fauna recorded in this study may represent the species that able to utilise the submerged environment at TBRS. We concur that the overall diversity is underestimated but species composition might symbolise the common landscape of TBRS.

CONCLUSION

Ant fauna at TBRS is diverse and intensive and further extensive surveys should be able to obtain a better picture of its diversity. The result of this study is far from comprehensive, but we hope the findings could provide the baseline information on the Formicidae fauna at TBRS and the dataset can be used for the planning of conservation and management of the area.

ACKNOWLEDGEMENT

We wish to thank the DWNP for their permission to conduct this research in TBRS. Special thanks to M. Izzat-Husna, M. Fahmi-Ahmad, Syed Ahmad Rizal and staffs of DWNP for their valuable help during field trips.

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Appendix 1 List of ants’ fauna collected from TBRS, Pahang.

Subfamily Genus Species Dolichoderinae Dolichoderus Dolichoderus cuspidatus Dolichoderinae Dolichoderus Dolichoderus spp. Dorylinae Aenictus Aenictus spp.1 Dorylinae Aenictus Aenictus spp.2 Dorylinae Cerapachys Cerapachys spp.1 Dorylinae Cerapachys Cerapachys spp.2 Formicinae Anoplolepis Anoplolepis gracilipes Formicinae Camponotus Camponotus gigas Formicinae Camponotus Camponotus spp.1 Formicinae Camponotus Camponotus spp.2 Formicinae Camponotus Camponotus spp.3 Formicinae Nylanderia Nylanderia spp. Formicinae Oecophylla Formicinae Plagiolepis Plagiolepis spp. Formicinae Polyrhachis Polyrhachis spp.1 Formicinae Polyrhachis Polyrhachis spp.2 Formicinae Pseudolasius Pseudolasius spp. Myrmicinae Cardiocondyla Cardiocondyla spp. Myrmicinae Carebara Carebara spp.1 Myrmicinae Carebara Carebara spp.2 Myrmicinae Carebara Caebara spp.3 Myrmicinae Carebara Carebara spp.4 Myrmicinae Cataulacus Cataulacus spp.1 Myrmicinae Cataulacus Cataulacus spp.2 Myrmicinae Crematogaster Crematogaster inflata Myrmicinae Crematogaster Crematogaster rogenhoferi Myrmicinae Crematogaster Crematogaster spp. 1 Myrmicinae Crematogaster Crematogaster spp.2 Myrmicinae Crematogaster Crematogaster spp.3 Myrmicinae Crematogaster Crematogaster spp.4 Myrmicinae Lophomyrmex Lophomyrmex bedoti Myrmicinae Meranoplus Meranoplus castaneus Myrmicinae Meranoplus Meranoplus mucronatus Myrmicinae Monomorium Monomorium spp. Myrmicinae Pheidole Pheidole spp. 1 Continued Ants diversity of Tasek Bera 13

Appendix 1 Continued Myrmicinae Pheidole Pheidole spp.2 Myrmicinae Strumigenys Strumigenys spp. Myrmicinae Tetramorium Tetramorium spp.1 Myrmicinae Tetramorium Tetramorium spp.2 Myrmicinae Tetramorium Tetramorium spp.3 Myrmicinae Tetramorium Tetramorium spp.4 Ponerinae Leptogenys Leptogenys spp.1 Ponerinae Leptogenys Leptogenys spp.2 Ponerinae Odontomachus Odontomachus rixosus Ponerinae Odontoponera Odontoponera transversa Ponerinae Pachycondyla Pachycondyla spp.1 Ponerinae Pachycondyla Pachycondyla spp.2 Ponerinae Pachycondyla Pachycondyla spp.3 Ponerinae Ponera Ponera spp.1 Ponerinae Ponera Ponera spp.2 Pseudomyrmecinae Tetraponera Tetraponera spp.1 Pseudomyrmecinae Tetraponera Tetraponera spp.2 Pseudomyrmecinae Tetraponera Tetraponera spp.3