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1 Graduate Institute of Entomology National Taiwan University Master s Thesis Dr. Wu, Wen-Jer Comparison of ant community composition on the ground of two types of habitat in Fushan experimental forest Pan, Chein-An January,

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3 Abstract For comparing the ant community composition the ant species were sampled by pitfall traps quantitatively from two different habitats, the forest and an artificial open area among Fushan experimental forest, from August 2000 to July Overall, 5 subfamilies, 31 genera and 53 species of ants were collected. Forty-five species were collected on the ground of the open area, while only 31 species were collected on forest floor. The species richness and abundance of the ant were higher in the open area, and Shannon coefficient of diversity and evenness were similar between two habitats. The species assemblage of the ant was analyzed by functional group method. The ratio of two functional groups, the competition group and disturbance group, was higher in the open area suggested that the effect of artificial removal of original plantation was similar to the effect of natural formation of treefall gaps, which may increase disturbance and available resources. Lower similarity of ant community composition were found between two sampling plots with more distance, but the ant community composition were not evidently different between these two habitats. 3

4 ... i...iii... iv... v ( ) ( )

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9 (Insecta) (Hymenoptera) (Vespoidea) (Formicidae) 10,000 (Bolton, 1995a, b) (bioindicator) (Majer, 1983) 9

10 (pitfall trap) 10

11 (Insecta) (Hymenoptera) (Vespoidea) (Formicidae) ( ) (caste) (generation) ( ) ( ) (Michener, 1969) (division of labor) 11

12 (temporal caste) (Hölldobler and Wilson, 1990; Robinson, 1992) (Wilson, 1971) (plasticity) (Robinson, 1992) (Calabi and Traniello, 1989) (Wilson, 1984) (Gordon, 1989) (superorganism) (Wilson, 1971) 12

13 30 % 75 % (Beck, 1971) (Papua New Guinea) (Wilson, 1959) 2500 m (Andersen, 1983) (Nash et al., 2000) (abundance) (diversity) (opportunist) Acromyrmex Atta (leaf cutting ant) 13

14 (Dacetonini) (Collembola) (harvesting ant) (Hölldobler and Wilson, 1990) (trophic level) (Dryer and Letourneau, 1999) (Abe and Darlingtion, 1985; Darlington, 1985; Higashi and Ito, 1989; Hölldobler and Wilson, 1990; Leponce et al., 1999) (Eggleton et al., 1995, 1996) 14

15 Leprogenys Processionalis (Hölldobler and Wilson, 1990) (symbiosis) (Homoptera) (Lepidoptera) (Hölldobler and Wilson, 1990) (Acacia sphaerocephala) (Pseudomyrmex) (Carolina) (Argentine ant, Linepithema humile) (regeneration) (Christian, 2001) 15

16 (mineralization) (Pêtal, 1998) (hydraulic properties) (Wang et al., 1996) (condition) (resource) Andersen (1995) Hölldobler and Wilson (1990) (temperature-humidity envelope) ( ) Levings (1983) Barro Colorado Island 16

17 (extrafloral nectaries) (Blüthgen et al., 2000) (Perfecto and Snelling, 1995) (competition) (Andersen, 1995) 17

18 (Argentine ant, Linepithema humile) (Fire ant, Solenopsis invicta) (Way et al., 1997; Human and Gordon, 1999; Sanders and Gordon, 2000) (Wilson, 1971) (Oklahoma) (ant lion) (Gotelli, 1996) (Hölldobler and Wilson, 1990) Aenictus (army ant) (Hirosawa et al., 2000) 18

19 (disturbance) (White and Pickett, 1985) Northern Chihuahuan Desert Conomyrma insana Pogonomyrmex desertorum (Nash et al., 2000) Chaco (Bestelmeyer and Wiens, 1996) (North Carolina) species assemblage (Peck et al., 1998) Roth and Perfecto (1994) (Shannon s diversity index) (evenness) 19

20 (bioindicator) ( Majer, 1983; De Bruyn, 1993; Andersen, 1997; King et al., 1998; Nash et al., 1998) Majer (1983) Wang et al. (2000) (functional groups) Andersen (1995) (competition) (stress) (disturbance) 20

21 Andersen (1995) 7 Dominate Dolichoderinae Generalized Myrmicinae Opportunists Dominate Dolichoderinae Dominate Dolichoderinae ( ) Dominate Dolichoderinae Opportunists (species inventory) Delabie et al. (2000) Winkler sacks Berlease funnel 17 Wang et al. (2001) 21

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23 30 km 16 km ha 400 m 1400 m (1419 m) mm 96 % RH (1993~1999 ) 23

24 1000 m 3 ( ) a. b. a. b. Fig. 1. Fushan experimental forest. a. area divisions; b. artificial installations. ( ) 4 50 ml ( cm) 25 ml 95 1 m 24

25 ( ) 7 1m 1m ( ) Fig. 2. Sampling design. 1. (Special use plant area, S) 40 m 120 m 25

26 ( ) 1-1 ~ 4-4 W S (S W ) Fig. 3. Sampling area map (S: Open area; W: Forest). 26

27 2. (First watershed area, W) 54 3 m 100 m 1 40~60 m W1 W12 12 (Shannon s diversity index, H ) (eveness, E) 1 pi i H' = n i= 1 p i ln p i 1 27

28 2 S E H ln S =. 2 Andersen (1995) Stress group Strumigenys solifontis 2. Competition group Oligomyrmex oni 3. Disturbance group 28

29 Monomorium chienese (linear regression) (Correlation Coefficient) 29

30 (1-1~4-4) (W1~W12) Bray-Curtis Cluster Analysis (Pielou, 1984) log(n+1) 30

31 (Myrmicinae) % (Ponerinae) 10 (19 %) (Formicinae) 5 (10 %) (Dolichoderinae) 2 (4 %) (Aenictinae) 1 (2 %) (Pachyconera luteipe) (Pheidole noda) (S) (W) Table 1. Ant species, abundance and functional group captured by pitfall trap on the ground of open area (S) and forest floor (W) in Fushan experimental forest during August, 2000 to July, 2001 Subfamily Scientific name Chinese name S W Total Functional group* MYRMICINAE Aphaenogsters gracilento C Aphaenogsters caperatus C Aphaenogsters tipuna R Aphaenogsters isthmus St Aphaenogsters lepida R Aphaenogsters cruciatus C Cadiocondyla monticola C Cadiocondyla wroughtonii C Kyidris mutica C Leptothorax cerius R Monomorium chinense R Monomorium intrudens St Monomorium caecus St Myrmecina taiwana C 31

32 ( ) Table 1. (continued) Subfamily Scientific name Chinese name S W Total Functional group* Oligomyrmex sauteri St Oligomyrmex oni C Pheidole noda St Pheidole rinae tipuna St Pheidole fustis C Pheidole pieli R Pheidole ernsti St Pheidole inocula C Pheidologeton yanoi R Pristomyrmex pungens R Rhaopalomsatix sp St Smithistruma leptothix C Strumigenys solifontis St Strumigenys hispida St Strumigenys fomosensis St Tetramorium pacificum St Tetramorium nipponense C Vollenhovia acanthinus C Vollenhovia formosae C Vollenhovia galvus St PONERINAE Crytopone taivanensis C Gnamptogenys taivanensis St Leptogenys kitteli St Leptogenys confucii St Odontomachus monticola St Pachyconera luteipe R Pachyconera javanus St Pachyconera chinensis R Pachyconera darwinii St Ponera alisana St FORMICINAE Camponotus sp C Formica japonica R Lasius talpa St Paratrechina flavipes R Pseudolasius binghami taivanae St DOLICHODERINAE Tapanoma melanocephalum R Technomyrmex albipes St AENICTINAE Aenictus sp C Total *C: Competition group; D: Disturbance group; St: Stress group. 32

33 ( ) ( ) ( ) 5 (Pachyconera luteipe) (Pachyconera javanus) 3 ( ) 33

34 Table 2. The number of ant species, mean individuals per trap, Shannon Habitat coefficient of diversity and evenness captured from pitfall trap on the ground of open area and forest floor in Fushan experimental forest during August, 2000 to July, 2001 Number of species Mean individuals per trap Shannon s diversity index, H Evenness, E Ground of open area ± Forest floor ± Total Accumulated number of species Number of pitfall trap (S) (W) Fig. 4. The species accumulation curve over the number of pitfall trap captured on the ground of open area (S) and forest floor (W) in Fushan experimental forest during August, 2000 to July,

35 Table 3. The dominant ant species captured from pitfall trap on the ground of open area and forest floor in Fushan experimental forest and their abundance percentage during August, 2000 to July, 2001 Rank Ground of Open Area Abundance (%) Forest Floor Abundance (%) 1 Pheidole noda Pachyconera luteipe Pachyconera luteipe Pheidologeton yanoi Monomorium chinense 8.48 Pheidole pieli Pachyconera javanus 6.47 Pachyconera javanus Formica japonica 4.84 Aphaenogsters gracilento 6.56 Others Others ( ) ( a.) ( b.) 35

36 a. b. Number of species Percentage (%) (S) (W) a. b. (St C D ) Fig. 5. a. The species number and b. the ratio of functional groups composition captured by pitfall trap on the ground of open area (S) and forest floor (W) in Fushan experimental forest during August, 2000 to July, (St: Stress group; C: Competition group; D: Disturbance group) 36

37 ~ ( ) ~10 2~3 ( ) Individuals per trap Aug-00 Sep-00 Oct-00 Nov-00 Dec-00 Jan-01 Feb (S) Mar-01 Apr-01 May-01 (W) Fig. 6. Monthly change in the mean individuals per pitfall trap on the ground of open area (S) and forest floor (W) in Fushan experimental forest during August, 2000 to July, Month Jun-01 Jul-01 S W 37

38 35 30 Number of species S W T 0 Aug-00 Sep-00 Oct-00 Nov-00 Dec-00 Jan-01 Feb-01 Mar-01 Apr-01 May-01 Jun-01 Jul-01 Month (S) (W) Fig. 7. Monthly change in the number of ant species captured from pitfall trap on the ground of open area (S); forest floor (W) and sum of both habitats (T) in Fushan experimental forest during August, 2000 to July, (13.3 ) (24.2 ) ( a.) 88.3 % % ( b.) mm mm ( c.) 38

39 a. Temperature ( ) b. Relative humidity (%) c. Precipitation (mm) Month a. b. c. Fig. 8. The weather condition of a. temperature; b. relative humidity; c. precipitation during the period of sampling by pitfall trap in Fushan experimental forest from August, 2000 to July, 2001 each month. 39

40 3 0.7 ( ) Table 4. The correlation coefficient between the monthly change of ant abundance captured from pitfall trap in Fushan experimental forest and the monthly change of temperature, relative humidity and precipitation during August, 2000 to July, 2001 Correlation Coefficient Temperature Relative Humidity Precipitation p

41 Pielou (1984) Bray-Curtis Cluster Analysis (1-1 ~ 4-4) (similarity) 65 % (W1 ~ W12) 50 ~ 65 % ( ) (1-1 ~ 4-4 W1 ~ W12 ) Fig. 9. Dendrogram of similarity between sampling plots captured from pitfall trap in Fushan experimental forest during August, 2000 to July, 2001 (1-1 ~ 4-4: sampling plots on the ground of open area; W1 ~ W12: sampling plots on the forest floor) 41

42 ( ) ( ) 6 7 (Campontus) (Protanilla) Majer (1983) 1 km 42

43 (Majer, 1983; Roth and Perfecto, 1994; Perfecto and Snelling, 1995) Feener and Schupp (1998) (treefall gaps) (Fetcher et al., 1985; Denslow, 1987) (Schupp and Feener, 1991) (Levings, 1983; Torres, 1984; Hölldobler and Wilson, 1990) 43

44 ( ) (>75%) 50 % Majer et al. (1997) 25 ~ 45 m 44

45 45

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53 Appendix 1. Profiles of three ant functional groups in Australia (Modified from Andersen, 1995) Functional groups * Dominate Dolichoderinae Generalized Myrmicinae Competivity Stress Disturbance tolerance tolerance Note ++ - Equal to arbor Equal to shrub Opportunists Equal to grass * Still including Subordinate Camponotini; Climate specialists; Cryptic species; Specialist predators, to discriminate secondary habitat characteristics. 53

54 Appendix 2. Functional group discriminance of ants in Taiwan Attribute St Stress Group C Competition Group D Disturbance Group Colony Size Small Variable Large Queen Number Monogyny Monogyny or Polygyny Polygyny Soldier Caste None Few species Most species Territorial Exclusiveness Low High High Feeding Type Oilgophagus Oilgophagus or Monophagus Polyphagus Foraging Strategies Solitary or Large group or Solitary or (Serching) Small group Territoriality Territoriality Foraging Strategies (Retrieve) Solitary or Small group Territoriality or Large Group Retrieve Daily Diurnal or Nocturnal Diurnal Diurnal or Nocturnal Swarming Period Fixed Fixed Unfixed Tolerance to Extreme Temperature and Humidity High Low Low Colony Productivity Low High Low Settlement Ability High Low High Special Case Parasitism, Army Ant, Ant-eater Vagabond 54

55 (1997 ~ 2000) Appendix 3. Comparison of the result in this study and in the past investigation in Fushan (1997~2000) Category Present study Number of taxon Past study Capture (%) Subfamily Genus Species

56 Appendix 4. The result of sampling plots on the ground of open area Subfamily Scientific name Chinese name Total MYRMICINAE Aphaenogsters gracilento Aphaenogsters caperatus Aphaenogsters tipuna Aphaenogsters isthmus Aphaenogsters lepida Aphaenogsters cruciatus Cadiocondyla monticola Cadiocondyla wroughtonii Crematogaster laboriosa Crematogaster pia taivanae Crematogaster subnuda formasae Kyidris mutica Leptothorax cerius Lordmyrma formosa Monomorium chinense Monomorium intrudens Monomorium caecus Oligomyrmex sauteri Myrmecina taiwana

57 ( ) Appendix 4. (continued) Subfamily Scientific name Chinese name Total Oligomyrmex oni Pheidole noda Pheidole rinae tipuna Pheidole fustis Pheidole pieli Pheidole ernsti Pheidole inocula Pheidologeton yanoi Pristomyrmex pungens Rhaopalomsatix sp Smithistruma leptothix Strumigenys solifontis Strumigenys hispida Strumigenys fomosensis Tetramorium pacificum Tetramorium nipponense Vollenhovia acanthinus Vollenhovia formosae Vollenhovia galvus

58 ( ) Appendix 4. (continued) Subfamily Scientific name Chinese name Total PONERINAE Crytopone taivanensis Gnamptogenys taivanensis Leptogenys kitteli Leptogenys confucii Odontomachus monticola Pachyconera luteipe Pachyconera javanus Pachyconera chinensis Pachyconera darwinii Ponera alisana FORMICINAE Camponotus sp Formica japonica Lasius talpa Paratrechina flavipes Polyrjachis sp Pseudolasius binghami taivanae

59 ( ) Appendix 4. (continued) Subfamily Scientific name Chinese name Total DOLICHODERINAE Tapanoma melanocephalum Technomyrmex albipes AENICTINAE Aenictus sp Total 59

60 Appendix 5. The result of sampling plots on forest floor Subfamily Scientific name Chinese name W1 W2 W3 W4 W5 W6 W7 W8 W9 W10 W11 W12 Total MYRMICINAE Aphaenogsters gracilento Aphaenogsters caperatus Aphaenogsters tipuna Aphaenogsters isthmus Aphaenogsters lepida Aphaenogsters cruciatus Cadiocondyla monticola Cadiocondyla wroughtonii Crematogaster laboriosa Crematogaster pia taivanae Crematogaster subnuda formasae Kyidris mutica Leptothorax cerius Lordmyrma formosa Monomorium chinense Monomorium intrudens Monomorium caecus Myrmecina taiwana Oligomyrmex sauteri

61 ( ) Appendix 5. (continued) Subfamily Scientific name Chinese name W1 W2 W3 W4 W5 W6 W7 W8 W9 W10 W11 W12 Total Oligomyrmex oni Pheidole noda Pheidole rinae tipuna Pheidole fustis Pheidole pieli Pheidole ernsti Pheidole inocula Pheidologeton yanoi Pristomyrmex pungens Rhaopalomsatix sp Smithistruma leptothix Strumigenys solifontis Strumigenys hispida Strumigenys fomosensis Tetramorium pacificum Tetramorium nipponense Vollenhovia acanthinus Vollenhovia formosae Vollenhovia galvus

62 ( ) Appendix 5. (continued) Subfamily Scientific name Chinese name W1 W2 W3 W4 W5 W6 W7 W8 W9 W10 W11 W12 Total PONERINAE Crytopone taivanensis Gnamptogenys taivanensis Leptogenys kitteli Leptogenys confucii Odontomachus monticola Pachyconera luteipe Pachyconera javanus Pachyconera chinensis Pachyconera darwinii Ponera alisana FORMICINAE Camponotus sp Formica japonica Lasius talpa Paratrechina flavipes Polyrjachis sp Pseudolasius binghami taivanae

63 ( ) Appendix 5. (continued) Subfamily Scientific name Chinese name W1 W2 W3 W4 W5 W6 W7 W8 W9 W10 W11 W12 Total DOLICHODERINAE Tapanoma melanocephalum Technomyrmex albipes AENICTINAE Aenictus sp Total

64 ( NSC B A10 NSC B ) 64