Diversity and Abundance of Carabidae and Staphylinidae 66 2 67 3 (Insecta: Coleoptera) in Four Montane Habitat Types on Mt

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55 HOSTED BY Contents lists available at ScienceDirect 56 57 HAYATI Journal of Biosciences 58 59 journal homepage: http://www.journals.elsevier.com/ 60 hayati-journal-of-biosciences 61 62 63 Original research article 64 65 1 Diversity and Abundance of Carabidae and Staphylinidae 66 2 67 3 (Insecta: Coleoptera) in Four Montane Habitat Types on Mt. 68 4 69 Q10 Bawakaraeng, South Sulawesi 5 70 6 1 fi 1* 2 71 7 Q9 Agmal Qodri, Rika Raf udin, Woro Anggraitoningsih Noerdjito 72 8 Q1 1 Department of Biology, Faculty of Mathematics and Natural Sciences, Bogor Agricultural University, Bogor, Indonesia. 73 9 2 Zoology Division (Museum Zoologicum Bogoriense), Research Center for Biology-LIPI, Cibinong, Indonesia. 74 10 75 11 76 12 Q2 article info abstract 77 13 78 14 Article history: Carabidae and Staphylinidae are the two beetle families frequently found to be most abundant and 79 Received 5 May 2015 15 diverse in forest ecosystem. Their roles especially as generalist predators are important in forest 80 16 Accepted 6 April 2016 ecosystem. No studies reported diversity and abundance of Carabidae and Staphylinidae in forest Available online xxx 81 ecosystem on Mt. Bawakaraeng, specifically in montane habitat yet. The aim of this study was to analyze 17 82 diversity and abundance of Carabidae and Staphylinidae in four montane habitat types, i.e. agricultural 18 83 KEYWORDS: area, pine forest, eucalypts and natural forest (1835 m asl), and natural forest (2165 m asl). They were 19 Carabidae, collected using pitfall traps. A total of 42 carabid beetles belonging to nine species and 260 staphylinid 84 20 diversity, beetles belonging to 37 species were collected. Diversity and abundance of Staphylinidae were higher 85 21 montane habitat, than Carabidae, this is predicted because of higher mobility in Staphylinidae compared to Carabidae. In 86 22 Mt. Bawakaraeng, Staphylinidae, Carabidae, the highest species richness was recorded in agricultural area, whereas the highest species 87 23 Sulawesi richness of Staphylinidae occurred in natural forest (2165 m asl). In Staphylinidae, the two largest 88 24 subfamilies are Aleocharinae and Staphylininae. Aephinidius adelioides occupied the highest abundance 89 25 of Carabidae and found in agricultural area. The differences in each montane habitat type are presumed 90 26 to cause variation in species richness of soil beetles, especially for Carabidae. 91 © 27 Copyright 2016 Institut Pertanian Bogor. Production and hosting by Elsevier B.V. This is an open access 92 article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). 28 93 29 94 30 95 31 96 32 1. Introduction In forest ecosystems, soil beetles show important roles in soil 97 33 food chain being predators, decomposers, and phytophagous. Car- 98 34 Q3 The geological history of Sulawesi Island in forming four abidae is one of the beetle families, which are predominantly 99 35 different arm (north, south, east, southeast) was caused by tectonic predators (Nitzu et al. 2008), especially as generalist predators (feed 100 36 processes of active plate boundaries (Hamilton 1979). Mt. Bawa- on a variety of arthropod preys, such as Collembola, soil mites, and 101 37 karaeng (2830 m asl) is located in the south arm in Gowa district, larvae of Diptera) (Ribera et al. 1999). Besides Carabidae, Staph- 102 38 South Sulawesi, and the mountain was formed through the volcanic ylinidae is also a generalist predator (Pohl et al. 2008). As a 103 39 activity in the Pleistocene (Hasnawir & Kubota 2008). Sulawesi has generalist predator, they serve as seed eaters and pollen feeders, 104 40 several types of forest, i.e. lowland (0e400 m asl), hill respectively (Steel 1970; Hanski & Hammond 1986). 105 41 (400e850 m asl), upland (850e1500 m asl), montane The species richness of soil beetle decreased with the increasing 106 42 (1500e2500 m asl), and tropalpine forest (>2500 m asl) (Cannon altitude in montane forest (Hanski & Hammond 1986; Maveety 107 43 et al. 2007). Montane forest is often considered as less diverse et al. 2011). Thus, different types of montane forest or altitudes 108 44 area than lowland forest, but high endemism occurs in this area may inhabit different diversity of Carabidae and Staphylinidae. 109 45 (Anderson & Ashe 2000). Anotylus sp. (Oxytelinae) and Philonthus sp. (Staphylininae) are the 110 46 common staphylinids that occur in montane forests in Gunung 111 47 Mulu National Park, Sarawak, Malaysia (Hanski & Hammond 1986). 112 48 Another example, the three genus of three tribes, i.e. Pelmatellus 113 49 (Harpalini), Dyscolus (Platynini), and Bembidion (Bembidiini) are 114 50 * Corresponding author. 115 fi fi commonly found in tropical montane forest, Andes mountains, 51 E-mail address: rraf [email protected] (R. Raf udin). 116 Peer review under responsibility of Institut Pertanian Bogor. southeastern Peru (Maveety et al. 2011). 52 117 53 http://dx.doi.org/10.1016/j.hjb.2016.04.002 118 54 1978-3019/Copyright © 2016 Institut Pertanian Bogor. Production and hosting by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http:// 119 creativecommons.org/licenses/by-nc-nd/4.0/).

Please cite this article in press as: Qodri, A., et al., Diversity and Abundance of Carabidae and Staphylinidae (Insecta: Coleoptera) in Four Montane Habitat Types on Mt. Bawakaraeng, South Sulawesi, HAYATI J Biosci (2016), http://dx.doi.org/10.1016/j.hjb.2016.04.002 HJB31_proof ■ 9 July 2016 ■ 2/7

2 A. Qodri, et al

1 Research on diversity and abundance of Carabidae and Staph- Laboratory of Entomology, Zoology Division, and Research Center 66 2 ylinidae in the area of Mt. Bawakaraeng particularly related to for Biology LIPI. 67 3 montane habitats has not been reported yet. Therefore, it is 68 4 69 necessary to conduct a research on the diversity and abundance of 2.3. Identification 5 70 Carabidae and Staphylinidae in various montane habitat types on Family level identification was identified using Triplehorn and 6 71 Mt. Bawakaraeng. The objective of this study was to analyze di- Johnson (2005). Carabidae was identified according to Darlington 7 72 versity and abundance of Carabidae and Staphylinidae in four (1970), Sawada and Wiesner (2000), Ito (2009), and specimen 8 73 montane habitat types on Mt. Bawakaraeng, South Sulawesi. This references in Museum Zoologicum Bogoriense (MZB) LIPI Cibinong. 9 fi 74 result is expected to provide scienti c information on the area The further identification for Staphylinidae was performed using 10 75 richness related to diversity and abundance of Carabidae and Cameron (1930) and specimen references in MZB. For morpho- 11 76 Staphylinidae. species level was identified based on external morphology. Almost 12 77 all species of Carabidae and Staphylinidae were verified by Co- 13 78 2. Materials and Methods leopterists listed at Table 2. All specimens were deposited in MZB. 14 79 15 2.1. Study sites 80 2.4. Environmental variables 16 Collecting sample of Carabidae and Staphylinidae was con- 81 We measured the environmental variables, i.e. temperature, 17 ducted once in dry season from September until October 2013 in 82 humidity, soil temperature, soil moisture, and soil pH. Temperature 18 four montane habitat types on Mt. Bawakaraeng. Those habitats are 83 and humidity were measured using digital thermohygrometer. Soil 19 (1) agricultural area (1465 m asl), consisted of tomatoes (Sol- 84 temperature was measured using soil thermometer. Soil moisture 20 anaceae), gourds (Cucurbitaceae), leeks (Alliaceae), and mustards 85 and pH were measured using soil tester Takemura. The range of soil 21 (Brassicaceae). There was no canopy cover in this habitat. (2) Pine 86 moisture value at soil tester is 1%e8% and soil pH value ranges from 22 forest (1545 m asl), dominated by pine trees and ferns. There were 87 3to8. 23 also weeds of Ageratum sp. in this habitat. (3) Mixed forest, i.e. 88 24 partly eucalypts forest, partly natural forest (1835 m asl). The nat- 89 25 ural forest in this altitude was dominated by trees of Magnolia 2.5. Data analysis 90 26 vrieseana (Magnoliaceae), and the trees of other families were Data of Carabidae and Staphylinidae were analyzed for species 91 27 coated by mosses and epiphytic plants. This habitat was overgrown richness (S), Shannon's diversity index (H'), Pielou's evenness index 92 28 by shrubs Leucosyke capitellata (Urticaceae). (4) Natural forest (E), and Simpson's index/dominance of species (D)(Magurran 93 29 (2165 m asl), dominated by trees of Melicope accedens (Rutaceae) 1988). The relationship among species, habitats, and environ- 94 30 and the trees of other families were coated by bryophytes and mental variables was analyzed using canonical correspondence 95 31 epiphytic plants, as well as had the most dense canopy cover analysis (CCA) implemented in the Paleontological Statistics (PAST) 96 32 (Table 1). program version 1.93 (Hammer & Harper 2006). 97 33 98 34 99 2.2. Specimen collection 3. Results 35 100 Carabid and staphylinid beetles were collected using pitfall 36 101 traps. Fifteen pitfall traps were positioned systematically in agri- 3.1. The diversity and abundance of Carabidae 37 102 cultural area and set up randomly in pine forest, mixed forest, and Compared to Staphylinidae, number of collected Carabidae was 38 103 natural forest. The space between pitfall traps are 10e20 m in a plot less, i.e. a total of 42 individuals, those from three subfamilies and 39 104 area of 2400 m2 in all the habitats, and with reference to Green- nine species of carabids. The subfamily Harpalinae was the most 40 105 Q8 slade (1973) we kept the traps for 2 days. The differential common carabids collected in the areas of Mt. Bawakaraeng, 41 106 arrangement model of pitfall traps between the three montane belonging to seven species. From each subfamily Cicindelinae and 42 107 habitat types (pine forest, mixed forest, and natural forest) and Scaritinae one species was collected, i.e. Hipparidium shinjii and 43 108 agricultural area was due to the obstacles in those three montane Clivina sp., respectively. The highest Carabidae diversity was in 44 109 types, such as the disarrangement of trees disposition and oblique natural forest (2165 m asl). However, agricultural area revealed 45 110 soil surface contour. The pitfall traps were made of a plastic cup home for the highest abundance and species richness of carabids 46 111 (height 10 cm, base diameter 5.7 cm, and diameter of upper base and Aephnidius adelioides (Harpalinae) was the most dominant in 47 112 9.2 cm) and were placed in the ground with the mouth flat against this area. Nevertheless, the dominance of A. adelioides (Figure 1A) in 48 113 the soil surface. Each trap was filled with 70% alcohol which serves agricultural area led to the lower diversity in that area compared to 49 114 as a preservative agent of the specimens. Transparent roofs mixed forest and natural forest (2165 m asl). The other carabid 50 115 (20 20 cm) were used as the protective of pitfall trap from rain species, i.e. H. shinjii (Cicindelinae) (Figure 1B), Trigonotomi (Les- 51 116 and litter. The distance between the roof and soil surface is 10 cm. ticus sp.1) (Harpalinae) (Figure 1C), and Platynini sp.1 (Harpalinae), 52 117 Collected Carabidae and Staphylinidae were identified in the respectively were found in pine forest, in mixed forest: natural 53 118 54 119 55 Table 1. Description of montane habitat types at study sites on Mt. Bawakaraeng 120 56 121 57 Code Coordinate Montane habitat types Altitude (m asl) 122 58 AA S 05 150 03.100 Agricultural area 1465 123 59 E 119 530 56.700 124 0 00 60 PF S 05 15 29.3 Pine forest 1545 125 E 119 540 31.800 61 MF S 05 160 42.500 Mixed forest: eucalypts forest and natural forest 1835 126 62 E 119 540 58.500 127 63 NF S 05 170 11.400 Natural forest 2165 128 0 00 64 E 119 55 47.6 129 65 In the four montane habitat types were found cattle dungs. 130

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1 Table 2. Veriﬁcation of Carabidae and Staphylinidae species collected on Mt. Bawakaraeng Q6 66 2 67 ﬁ 3 No. Family Species Veri er Institution/address 68 4 1 Carabidae Hipparidium shinjii David L. Pearson School of Life Sciences, Arizona State University 69 5 Jürgen Wiesner Dresdener Ring 11, 38444, Wolfsburg, Germany 70 2 Aephnidius adelioides Noboru Ito Konica Minolta Business Technologies, Toyokawa, Japan 6 3 Kipling Will Essig Museum of Entomology, University of California, Berkeley 71 7 4 Harpaloxenus sp. Noboru Ito Konica Minolta Business Technologies, 72 8 5 Pseudotrichotichnus sp. Toyokawa, Japan 73 9 6 Clivina sp. 74 10 7 Trigonotomi (Lesticus sp.?) Kipling Will Essig Museum of Entomology, University of California, Berkeley 75 8 Platynini sp. Dmitri N. Fedorenko A.N. Severtsov Institute of Ecology and Evolution, Russian Academy 11 of Sciences, Moscow 119071, Russia 76 12 9 Staphylinidae Aleochara sp. Munetoshi Maruyama Kyushu University Museum, Fukuoka, Japan 77 13 10 Anomognathus sp. 78 14 11 Atheta sp. 79 12 Athetini sp. 15 13 Coenonica sp. 80 16 14 Drusilla sp. 81 17 15 Homalotini sp. 82 18 16 Neosilusa sp. 83 19 17 Tropimenelytron sp. 84 18 Paederus sp. Adam J. Brunke Naturhistoriches Museum Wien 20 19 Medonina sp. 85 21 20 Erichsonius sp. 86 22 21 Philonthus sp. 87 23 22 Xantholinini sp. 88 23 Mycetoporini sp. 24 89 24 Sepedophilus sp. 25 25 Anotylus sp. Harald Schillhammer Natural History Museum Vienna, International 90 26 26 Astenus sp. Research Institute for Entomology, Burgring 7 91 27 27 Stenus sp. A e 1010 WIEN, Austria 92 28 28 Phloeonomus sp. Margaret K. Thayer Field Museum of Natural History, Chicago, USA 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 Figure 1. Species of Carabidae. (A) Aephnidius adelioides, (B) Hipparidium shinjii, (C) Trigonotomi (Lesticus sp.?). Scale bar: 1 mm. 115 51 116 52 117 forest (1835 m asl) under the forest canopy adjacent to open area, Athetini sp.1 (Aleocharinae). No species dominated in mixed forest. 53 118 and in both natural forest (1835 and 2165 m asl) (Table 3). Surprisingly, Staphylinidae diversity increased from agricultural 54 119 area to natural forest (2165 m asl). The highest abundance was in 55 120 natural forest (2165 m asl), whereas the lowest abundance was in 56 3.2. The diversity and abundance of Staphylinidae 121 mixed forest. The other staphylinid species, i.e. Paederus sp.1 57 A total of 260 individuals from eight subfamilies and 37 species 122 (Paederinae) was found in agricultural area, and Paederus sp.2 58 were collected and the two largest subfamilies were Aleocharinae 123 (Paederinae) (Figure 2B) and Mycetoporini sp.2 (Tachyporinae) 59 and Staphylininae. The subfamily Aleocharinae comprised 13 spe- 124 were collected in both natural forest (1835 and 2165 m asl) 60 cies and subfamily Staphylininae comprised eight species. Most of 125 (Table 4). 61 them dominated in three montane types, i.e. (1) agricultural area 126 62 was dominated by Aleochara sp. (Aleocharinae) and Xantholinini 127 63 sp. (Staphylininae) (Figure 2A), (2) pine forest was dominated by 3.3. Environmental variables 128 64 Tropimenelytron sp.1 (Aleocharinae) and Philonthus sp.3 (Staph- We recorded environmental data in Mt. Bawakaraeng, and 129 65 ylininae), and (3) natural forest (2165 m asl) was dominated by showed that the agricultural area has the highest for temperature 130

4 A. Qodri, et al

1 Table 3. Diversity and abundance of Carabidae of the three subfamilies in four montane types on Mt. Bawakaraeng 66 2 67 3 No Subfamily Species Montane habitat types Total 68 4 Agricultural area Pine forest Eucalypts and natural forest Natural forest 69 5 (1465 m asl) (1545 m asl) (1835 m asl) (2165 m asl) 70 6 71 1 Cicindelinae Hipparidium shinjii 010 01 7 2 Harpalinae Aephnidius adeliodes 30 0 0 0 30 72 8 3 Egadroma sp. 1 0 0 0 1 73 9 4 Harpaloxenus sp. 3 0 0 0 3 74 10 5 Pseudotrichotichnus sp. 0 0 0 1 1 75 6 Trigonotomi (Lesticus sp.?) 0 0 1 0 1 11 7 Platynini sp.1 0 0 1 1 2 76 12 8 Platynini sp.2 0 0 0 1 1 77 13 9 Scaritinae Clivina sp. 2 0 0 0 2 78 14 N (Abundance) 36 1 2 3 42 79 S (Species richness) 4 1 2 3 9 15 80 H’ (Diversity) 0.619 0 0.693 1.098 16 E (Evenness) 0.446 0 1 1 81 17 D (Dominance) 0.705 1 0.5 0.33 82 18 83 19 84 20 The two staphylinid species that were only found in natural 85 21 forest (2165 m asl) were Athetini sp.1 (Aleocharinae) and Phloeo- 86 22 nomus sp. (Omaliinae). Both were clustered in natural forest 87 23 (2165 m asl) together with Anotylus sp.1, Anotylus sp.2, and Anotylus 88 24 sp.3 (Oxytelinae), and had positive correlation with humidity and 89 25 soil moisture (Figure 4). 90 26 91 27 92 28 4. Discussion 93 29 94 30 The results of this study indicated that the diversity and abun- 95 31 dance of Staphylinidae were higher than Carabidae. The presence or 96 32 absence of the ability to fly is predicted to affect carabid ecology 97 33 and distribution. Carabidae in Australia do not have ability to flyin 98 34 both lowlands and highlands. In contrast, Carabidae in Papua New 99 35 Guinea in the lowlands mostly have the ability to fly, but in the 100 36 highlands they are incapable to fly(Darlington 1961). 101 37 The great diversity and abundance of Staphylinidae were sus- 102 Figure 2. Species of Staphylinidae. (A) Xantholinini sp., (B) Paederus sp.2. Scale bar: 38 1 mm. pected caused by their movement capability which are higher than 103 39 Carabidae. The high movement of Staphylinidae is supported by its 104 40 well-developed wings (Pohl et al. 2008), shown by Dolatia coriaria 105 41 (33.77 C) and soil temperature (21.63 C), whereas this area had the (Aleocharinae) that has rapid movement across surface of the 106 42 lowest for humidity (31.67%) and soil moisture (1.33%) compared to ground and is able to escape quickly when there is a disturbance 107 43 others. As expected, the natural forest (2165 m asl) was recorded to (Meihls & Hibbard 2009). Distribution of Orsunius (Paederinae) in 108 44 have the lowest temperature (22.77 C) and soil temperature oriental region was supported by ability of the rear wings to fly 109 45 (18.23 C), and the highest humidity (66%) and soil moisture (7%) (Assing 2011). Although in general, species richness and abundance 110 46 (Table 5). of Staphylinidae were higher than Carabidae (Ferro et al. 2012; Lee 111 47 et al. 2014), however, several studies indicated that abundance of 112 48 3.4. Correlation among Carabidae species to environmental Carabidae is greater than Staphylinidae but species richness is 113 49 variables contrary (Nitzu et al. 2008; Yu et al. 2013). 114 50 A. adelioides was the most abundant carabid in agricultural area. Aleocharinae and Staphylininae collected were more abundant Q4 115 51 The carabid species that was only collected in pine forest was and diverse than others (Table 4) presumably because both sub- 116 52 H. shinjii. Based on CCA, A. adelioides that was clustered in agri- families have the ability to forage, detect and locate dung and 117 53 cultural area together with Egadroma sp., Harpaloxenus sp. (Har- decaying mushroom from a long distance (Pohl et al. 2008). Thus, 118 54 palinae), and Clivina sp. (Scaritinae) had positive correlation with their ability in finding food makes them able to have fast adaption 119 55 temperature and soil temperature, however, H. shinjii was not to the environmental change and further to the longer temporal 120 56 correlated to all environmental variables (Figure 3). change (Pohl et al. 2008). 121 57 The great abundance, diversity, and species richness of Staph- 122 58 3.5. Correlation among Staphylinidae species to ylinidae in natural forest (2165 m asl) (Table 4) were expected 123 59 environmental variables because of their role as saproxylic in addition as predator. Sap- 124 60 The highest abundance of Staphylinidae occurred in agricultural roxylic is defined as organisms that play a role in the process of 125 61 area was Aleochara sp. (Aleocharinae) and Xantholinini sp. fungal decay in trees or in association with living and dead trees 126 62 (Staphylininae). CCA revealed that Aleochara sp. and Xantholinini (Alexander 2008). Our environment data in natural forest 127 63 sp. were clustered in agricultural area together with Paederus sp.1 (2165 m asl) showed the natural forest has the highest humidity 128 64 (Paederinae) and had positive correlation with temperature and (Table 5) thus in agreement with fungi habitat. The member of 129 65 soil temperature (Figure 4). three subfamilies Aleocharinae, Omaliinae, and Staphylininae 130

Diversity and Abundance of Carabidae and Staphylinidae 5

1 Table 4. Diversity and abundance of Staphylinidae of the eight subfamilies in four montane types on Mt. Bawakaraeng 66 2 67 3 No Subfamily Species Montane habitat types Total 68 4 Agricultural area Pine forest Eucalypts and natural forest Natural forest 69 5 (1465 m asl) (1545 m asl) (1835 m asl) (2165 m asl) 70 6 71 1 Aleocharinae Aleochara sp. 17 0 0 0 17 7 2 Anomognathus sp. 0 1 0 0 1 72 8 3 Atheta sp. 0 0 2 0 2 73 9 4 Athetini sp.1 0 0 0 17 17 74 10 5 Athetini sp.2 5 5 1 1 12 75 6 Athetini sp.3 1 8 0 2 11 11 7 Athetini sp.4 0 3 0 0 3 76 12 8 Coenonica sp. 0 0 0 1 1 77 13 9 Drusilla sp. 0 0 0 3 3 78 14 10 Homalotini sp. 0 0 1 0 1 79 11 Neosilusa sp. 1 0 0 0 1 15 80 12 Tropimenelytron sp.1 0 24 2 11 37 16 13 Tropimenelytron sp.2 0 2 3 13 18 81 17 14 Omaliinae Phloeonomus sp. 0 0 0 4 4 82 18 15 Osoriinae Eleusis sp. 0 0 0 2 2 83 19 16 Oxytelinae Anotylus sp.1 0 0 0 13 13 84 17 Anotylus sp.2 0 0 0 3 3 20 18 Anotylus sp.3 0 0 0 1 1 85 21 19 Anotylus sp.4 0 0 1 0 1 86 22 20 Anotylus sp.5 0 1 0 0 1 87 23 21 Anotylus sp.6 1 0 0 0 1 88 22 Paederinae Astenus sp. 0 0 2 0 2 24 89 23 Paederus sp.1 2 0 0 0 2 25 24 Paederus sp.2 0 0 4 6 10 90 26 25 Medonina sp. 1 0 1 0 2 91 27 26 Staphylininae Erichsonius sp. 0 0 1 0 1 92 28 27 Philonthus sp.1 8 0 0 0 8 93 28 Philonthus sp.2 4 0 0 0 4 29 29 Philonthus sp.3 0 16 1 2 19 94 30 30 Philonthus sp.4 0 7 1 1 9 95 31 31 Philonthus sp.5 0 0 0 1 1 96 32 32 Philonthus sp.6 0 3 0 1 4 97 33 33 Xantholinini sp. 36 0 0 0 36 98 34 Steninae Stenus sp. 0 0 0 2 2 34 35 Tachyporinae Mycetoporini sp.1 0 0 0 1 1 99 35 36 Mycetoporini sp.2 0 0 3 5 8 100 36 37 Sepedophilus sp. 1 0 0 0 1 101 37 N (Abundance) 77 70 23 90 260 102 S (Species richness) 11 10 13 20 37 38 H’ (Diversity) 1.632 1.863 2.426 2.525 103 39 E (Evenness) 0.68 0.809 0.945 0.842 104 40 D (Dominance) 0.286 0.202 0.1 0.106 105 41 106 42 107 (Table 4) were supposed to play role as saproxylic beetles. These Mycetoporini sp.2 (Tachyporinae), respectively. The dark body 43 108 data are supported by the finding of Toivanen & Kotiaho (2010) that coloration of Mycetoporini sp.2 in this instance is expected in 44 109 several species of Staphylinidae which act as saproxylic are accordance with habitat that has low light intensity to support it for 45 110 Phloeonomus pusillus (Omalinae), Homalota plana (Aleocharinae), foraging. The most dense canopy cover in natural forest specifically 46 111 and Quedius xanthopus (Staphylininae). is supposed to cause the low light intensity, which was also sup- 47 112 Although Staphylinidae has many species, however, only the ported by existence of bryophytes and high humidity (Table 5). The 48 113 agriculture area and natural forest showed the species character- genus Ischnosoma and Mycetoporus (Tachyporinae: Mycetoporini) 49 114 ized for these two habitats. Montane type at low altitude are often collected from moist leaf litter and moss in mature forests 50 115 (1465 m asl), i.e. agricultural area (Table 1), was characterized by (Webster et al. 2012). 51 116 species Aleochara sp. (Aleocharinae), Xantholinini sp. (Staph- Despite lower diversity of Carabidae compare to Staphylinidae 52 117 ylininae) (Figure 2A), and Paederus sp.1 (Paederinae). They are in this present study, it was showed that every single montane type 53 118 presumed to have strong responses with vegetation in this area on Mt. Bawakaraeng has several characterized carabid species. 54 119 that serve food sources for them as shown by Aleochara bipustulata Abundance and species richness of Carabidae were the highest in 55 120 (Aleocharinae) that occur in cruciferous crops (Brassicaceae) of agricultural area (low montane type), presumed in this area Cara- 56 121 their roles as predators on larva of cabbage root flies Delia radicum bidae has a role as generalist predator, and it could be an effective 57 122 (Balog et al. 2008). The genus Xantholinus (Staphylininae: Xantho- regulator of crop pests (Bukejs et al. 2009). A. Adelioides (Harpali- 58 123 linini) often inhabit and dominate in open and disturbed areas nae) (Figure 1A) has the greatest abundance in agricultural area 59 124 (Brunke & Majka 2010). Paederus sp.1 is similar to P. fuscipes based (Table 3) which is a bioindicator for open areas (Fujita et al. 2008). 60 125 on external morphology characters (Cameron 1930). Paederus fus- The high abundance of A. adelioides was supposed to be due to its 61 126 cipes occurs only in agricultural area (in large number, collected in inability to fly(Baehr 2007), thus its dispersal power is low. 62 127 gourd), and not found in forest area (Nasir et al. 2012). H. shinjii (Cicindelinae) was the only carabid collected in pine 63 128 The natural forest at altitudes 1835 and 2165 m asl was char- forest (Table 3). Lots of ferns and litters of pine trees in pine forest 64 129 acterized by species Paederus sp.2 (Paederinae) (Figure 2B) and might be a suitable habitat for H. shinjii (Figure 1B). H. shinjii 65 130

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1 Table 5. Environmental variables in the four montane habitat types on Mt. Bawakaraeng 66 2 67 3 No. Variables Montane habitat types 68 4 AA SD PF SD MF SD NF SD 69 5 1 Temperature (C) 33.77 0.76 23.83 0 24.13 1.22 22.77 0.20 70 6 2 Humidity (%) 31.67 3.21 56.67 4.72 55.16 6.33 66 3.60 71 7 3 Soil pH 6.83 0.28 6.83 0.28 6 1.09 4 0 72 4 Soil moisture (%) 1.33 0.57 5.83 0.28 5.75 0.82 7 0 8 73 5 Soil temperature (C) 21.63 1.35 18.87 0.37 19.13 0.66 18.23 0.37 9 74 ¼ ¼ ¼ ¼ ¼ 10 AA agricultural area; MF mixed forest: eucalypts forest and natural forest; NF natural forest; PF pine forest; SD standard deviation. 75 11 76 closely related to H. shinjii has black and white body coloration that 12 77 camouflages its presence while hiding in vegetation and litters 13 Pse 78 3.5 (Mawdsley 2007). 14 Pla.2 II Hip 79 3.0 Trigonotomi (Lesticus sp.1) (Harpalinae: Pterostichini) was 15 80 2.5 IV collected only in natural forest (1835 m asl) (Table 3) in the closed- 16 81 2.0 canopy area near open area (Figure 1C). This result is in accordance 17 82 1.5 with Lesticus finisterrae (Harpalinae: Pterostichini) collected in 18 Aep, Ega 83 xis 2 xis Pla.1 closed-canopy area, but close to open area in upper montane moss A 1.0 19 Rha Rhs Har, Cli 84 forest, Finisterre Range, Papua New Guinea (Will & Kavanaugh 20 0.5 85 III 2012). The second carabid species characterized of closed area in 21 0.0 86 natural forest is Platynini sp.1 (Harpalinae), which occurred in both 22 -0.5 I pHs 87 Tri Ta Ts natural forests (1835 and 2165 m asl) (Table 3). The highest humidity 23 -1.0 Q5 88 -1.6 -0.8 0.0 0.8 1.6 2.4 3.2 4.0 4.8 and soil moisture in natural forest (Table 5)mightinfluence its 24 89 Axis 1 presence as found Dalatagonum (Harpalinae: Platynini) at the leaf- 25 90 litter in montane subtropical broad-leaved forests (Fedorenko 2011). 26 91 Figure 3. CCA showed the relationship among Carabidae species ( ), habitat ( ), and No Carabidae was found in eucalypts forest (1835 m asl) prob- 27 environmental variables ( ). Name of species: Hip, Hipparidium shinjii; Aep, 92 ably due to the allelopathic effects of eucalypts, and mostly does the 28 Aephnidius adelioides; Har, Harpaloxenus sp.; Les, Trigonotomi (Lesticus sp.?); Pse, 93 Staphylinidae. Only a few of staphylinid species, i.e. two individuals 29 Pseudotrichotichnus sp.; Ega, Egadroma sp.; Pla.1, Platynini sp.1; Pla.2, Platynini sp.2; Cli, 94 Clivina sp. Environmental variables: pHs, soil pH; Rha, humidity; Rhs, soil moisture; Ta, of Atheta sp. (Aleocharinae), one individual of Astenus sp. (Paeder- 30 95 temperature; Ts, soil temperature. Habitat: I (agricultural area), II (pine forest), III inae), and one individual of species Philonthus sp.4 (Staphylininae) 31 96 (mixed forest: eucalypts and natural forest), IV (natural forest). were collected in eucalypts forest. Eucalyptus essential oil possesses 32 97 a utility as unnatural pesticide or insecticide, which controls beetle 33 98 pests and other insects (Batish et al. 2008). 34 99 Besides living in soil habitat, tropical Carabidae have arboreal 35 Ani.1, Coe, Dru, Phl 100 Aty.6, Pae.1 e 1.5 Pae.2 Aty.1, Aty.2, Aty.3 morphological characters, and 30% 60% of them belong to the 36 Myc.2 Phi.1, Phi.2 101 1.0 Ele, Ste, Phi.5, Myc.1 Neo, Sep Harpalinae (Ober 2003). Arboreal characters allow Harpalinae to 37 Tro.2 102 0.5 IV Ale, Xan not only live on the ground but adapt to live on plants (barks and 38 Rha III Med 103 Ta I trunks) (Ober 2003). In addition to morphological characters of 39 0.0 104 Rhs Ts arboreal, Harpalinae also play a role as plant seed eaters (sper- 40 -0.5 105 Tro.1 pHs II Ani.2 mophages), such as herbaceous plants seeds (Honek et al. 2003). 41 2 Axis -1.0 Ani.3 106 Phi.6 Thus, this ability might be possible to lead to the great number of 42 -1.5 Phi.4 Ata, Hom, Aty.4 107 Phi.3 Harpalinae species richness in Mt. Bawakaraeng (Table 3). 43 -2.0 Ast, Eri 108 Amo Ani.4 We conclude that differences of diversity, abundance, and spe- 44 -2.5 109 Aty.5 cies richness in Carabidae and Staphylinidae are responses to each 45 110 -3.0 montane habitat type. Mobility of Staphylinidae is higher than 46 -0.9 -0.6 -0.30.0 0.3 0.6 0.9 1.2 1.5 1.8 111 Carabidae and becomes one of the reasons of greater staphylinid 47 Axis 1 112 diversity and abundance. The different species in the same genus 48 113 Figure 4. CCA showed the relationship among Staphylinidae species ( ), habitat ( ), could have different habitat and behavior, thus it is necessary to 49 114 and environmental variables ( ). Name of species: Ale, Aleochara sp.; Ano, identify Carabidae and Staphylinidae up to particular species for 50 Anomognathus sp.; Tro.1, Tropimenelytron sp.1; Tro.2, Tropimenelytron sp.2; Ath, Atheta 115 further research. 51 sp.; Ath.1, Athetini sp.1; Ath.2, Athetini sp.2; Ath.3, Athetini sp.3; Ath.4, Athetini sp.4; 116 52 Coe, Coenonica sp.; Dru, Drusilla sp.; Hom, Homalotini sp.; Neo, Neosilusa sp.; Phl, 117 Phloeonomus sp.; Ele, Eleusis sp.; Ani.1, Anotylus sp.1; Ani.2, Anotylus sp.2; Ani.3, Ano- 53 tylus sp.3; Ani.4, Anotylus sp.4; Ani.5, Anotylus sp.5; Ani.6, Anotylus sp.6; Ast, Astenus Uncited reference 118 54 sp.; Pae.1, Paederus sp.1; Pae.2, Paederus sp.2; Med, Medonina sp.; Phi.1, Philonthus sp.1; 119 55 Phi.2, Philonthus sp.2; Phi.3, Philonthus sp.3; Phi.4, Philonthus sp.4; Phi.5, Philonthus 120 Kahono and Setiadi 2007. Q7 56 sp.5; Phi.6, Philonthus sp.6; Xan, Xantholinini sp.; Ste, Stenus sp.; Myc.1, Mycetoporini 121 57 sp.1; Myc.2, Mycetoporini sp.2; Sep, Sepedophilus sp. 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