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INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT, 2016 VOL. 12, NOS. 1–2, 96–107 http://dx.doi.org/10.1080/21513732.2016.1158209 Special Issue: Synergies between biodiversity and timber management Effect of overstory tree species diversity and composition on ground foraging ants (Hymenoptera: Formicidae) in timber plantations in Ghana a b a S. Sky Stephens , Paul P. Bosu and Michael R. Wager a b School of Forestry, Northern Arizona University, Flagstaff, AZ, USA; Principal Research Scientist-Entomology & Forest Health, Head- Ecosystem Services & Climate Change Division, CSIR-Forestry Research Institute of Ghana University, Kumasi, Ghana ABSTRACT ARTICLE HISTORY Received 27 February 2015 Plantation forests are becoming an increasingly important component of the world’s forested Accepted 17 February 2016 ecosystem. However, relatively little is known about how forest plantation management, overstory tree species composition and diversity impact biodiversity of nontree components EDITED BY of the forest. We assessed changes in ant functional group composition as related to changes Sheila Ward in overstory tree diversity (monocultures vs. polycultures), species composition (native African KEYWORDS species vs. exotic teak), and time (one and two years after planting). A pitfall trapping scheme Formicidae; timber was implemented during the summer months of 2006 and 2007. A total of 7473 specimens plantations; Khaya ivorensis; were collected representing six subfamilies, 22 genera, and 65 species. We found no sig- Tectona grandis; biodiversity nificant differences in traditional diversity measures or functional group composition between treatments one year after planting. Two years after planting, we found that species richness of ground foraging ants had significantly increased (F = 4.60, d.f. = 4, 15, p = 0.01). Several observed trends may have indicated that these ant communities were in transition and will likely become more distinct over time as the different plantation types recover from disturbance and diverge from each other in overstory structure. Tectona grandis monocultures. In spite of their high Introduction productivity and ability to meet some timber produc- Tropical forests are under numerous economic, poli- tion goals, exotic species cannot satisfactorily fill the tical, and social pressures including land conversion ecological and socioeconomic roles of native species. for agricultural and infrastructure development and Little research has been undertaken to understand the tropical timber production. Africa has seen the high- relationship between forest management and biodi- est rate of change in forest cover with an annual loss versity (Hartley 2002). In addition, the majority of rate of just under 1 million hectares per year between assessments made have not satisfactorily evaluated a 1990 and 2010 (FAO 2011). Decreasing tropical forest variety of alternative land uses such as native species area, a decline in native timber stock, and increasing plantations, polycultures, and nonforested land uses demand for timber products have led to plantation (Stephens & Wagner 2007). forests becoming an increasingly important compo- With the increasing loss of habitats and biodiver- nent of the world’s forested lands. Between 2000 and sity around the world, there is an urgent need for 2010, the rate of planted forest establishment rose to biodiversity assessment (Agosti et al. 2000). There is 115,000 ha per year (FAO 2011). Planted forest area considerable interest in the identification of robust represented approximately 187 million hectares bioindicators for use in land monitoring and assess- worldwide in 2000 (FAO 2001) and rose to 260 ment programs (Noss 1990). Living organisms can million hectares worldwide by 2010 (FAO 2011). integrate a variety of effects over time that short-term Plantation forests are depended on for wood biomass physical and chemical measures cannot, making them production, soil and water conservation, and wind suitable indicators of environmental conditions protection (Carnus et al. 2003) and are key sources (Danks 1992). Traditionally, bioindicators have been of fuel wood and nontimber forest products (FAO used to assess ecosystem response to human-induced 2015). environmental perturbation (Agosti et al. 2000). In 2010, planted forests represented almost 2.5 Terrestrial invertebrates are an important component million hectares in West Africa (FAO 2011). In of forest ecosystems and comprise a major part of 2005, Ghana had an estimated 160,000 ha of forest their biological diversity (Beattie et al. 1992). In con- plantations (FAO 2005), an increase in 85,000 ha trast to larger more mobile animals, terrestrial inver- since 2001 (FAO 2001). Of this approximately 90% tebrates are less likely to move between treatment was estimated to be exotic species, predominantly units, so their presence is a better indication of and CONTACT S. Sky Stephens email@example.com United States Forest Service, Forest Health Protection, 704 Simms St, Golden CO 80401 USA © 2016 Informa UK Limited, trading as Taylor & Francis Group INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 97 more strongly related to site condition (Bromham 34ʹ33ʺ N and 7°10ʹ49ʺ N. The moist semi-deciduous et al. 1999). Terrestrial invertebrates dominate the forest zone is the most extensive closed canopy forest biomass, are highly diverse, occur ubiquitously, and type in Ghana, characterized by multistory canopy are fundamentally important in ecosystem function layers and approximately equal proportions of ever- (Samways 1994; Andersen 1995). Recent work on green and deciduous species. This is the major tim- bioindicators identification has focused on soil inver- ber-producing area in Ghana and is suitable for the tebrates and microfauna (Cluzeaua et al. 2012). production of agricultural crops, orchard crops, and Specific insect guilds are highly sensitive to habitat exotic timber plantations (Wagner et al. 2008). disturbance (Day et al. 1993; Samways 1994; Mendez During July of 2005, experimental plantations were et al. 1995). For example, carabids (Kromp 1990; established on degraded forest sites within the South Fomangsu Forest Reserve (6°35ʹN, 0° 57ʹW) and Beaudry et al. 1997; Villa-Castillo & Wagner 2002), Lepidoptera (Holloway & Stork 1991; Kremen 1992), Afram Headwaters Forest Reserve (7°10ʹN, 1°40ʹW). Odonata (Samways 1996), and Formicidae Both reserves are located within the moist semi- deciduous forest zone and are approximately 90 km (Buffington 1967; Majer 1982; Andersen 1990, 1991, 1995, 1997a; Roth et al. 1994; York 1994, 1999, 2000; apart. Degraded areas were identified within the Brown 1997; King et al. 1998; Peck et al. 1998; reserves as lacking overstory trees and being domi- Bromham et al. 1999; Andrew et al. 2000; nated by weedy grass species and some woody shrubs. Vanderwoude et al. 2000; Stephens & Wagner The degradation within the reserves was initially due 2006), have all been used successfully as bioindicators to encroachment of illegal agriculture and chainsaw (Peck et al. 1998). felling of timber. The degraded areas were within the We selected ants as potential bioindicators of land- bounds of the forest reserves and approximately use type, because of their high diversity and biomass, 25–40 ha in size. We anticipated that the neighboring their ecological importance at all trophic levels, their forests would provide a source for colonizing ant ease of sampling, and their well-understood commu- fauna. These plantations were established as part of nity dynamics (Andersen & Sparling 1997). a larger study examining the impacts of overstory tree Furthermore, ants consistently show strong succes- diversity and species composition on ecosystem sional patterns in ecosystems and their functional functions. diversity and composition are related to land man- agement practices and disturbances (Andersen & Experimental design Sparling 1997), including forest management (Andersen & Sparling 1997; Vanderwoude et al. The treatments selected included (1) a native species 2000; Willett 2001; Stephens & Wagner 2006). monoculture, of Khaya ivorensis (an African maho- Majer (1983) showed that ants could be used to gany), (2) exotic species monoculture, Tectona monitor ecosystem recovery and disturbance and grandis (teak), and two polyculture conditions of (3) have been used extensively in Australia to observe a six native timber species mixture, and (4) a 10 rehabilitation of mine sites. Additionally, ants have native timber species mixture (Table 1). The experi- high spatial and temporal fidelity to sites (Nakamura mental plantations included tree species that repre- et al. 2015). sent different successional stages, growth forms and The objectives of this study were to experimentally economic values ranging from timber to non-timber evaluate changes in ant species diversity and func- forest products (Table 2). Two replicates of each tional group composition under different levels of treatment were installed at each Forest Reserve. overstory tree diversity (a monoculture and six and Plantations were set in complete randomized blocks 10 species polycultures of native species) and in plan- with each block containing 1 replicate of each of the tations of native African species and an exotic species four treatments, for two replicates of each treatment (Tectona grandis, or teak). We also looked for indi- cator species and functional groups for each type of Table 1. Species diversity and composition of treatments. plantation. We tested the hypothesis that changes in Description Species ant species diversity and ant functional group com- Treatment 1 Khaya ivorensis position would be related to differences in overstory Native species Monoculture tree diversity and/or native vs. exotic species. Treatment 2 Tectona grandis Exotic species Monoculture Treatment 3 Albizia zygia, Ceiba pentandra, Khaya ivorensis, Materials and methods Six native timber Mansonia altísima, Milicia excelsa, Pericopsis elata Species mixture Study sites Treatment 4 Albizia zygia, Ceiba pentandra, Khaya ivorensis, Ten native timber Mansonia altísima, Milicia excelsa, Pericopsis elata, This study was conducted in Ghana, West Africa in Species mixture Tieghemella heckelii, Terminila superba, Tetrapleura tetraptera the moist semi-deciduous forest zone between 6° 98 S. S. STEPHENS ET AL. Table 2. Tree species characteristics. Common Native vs. Nonpioneering light Non-pioneering Nitrogen Economic Species Name Exotic Pioneer demanding shade bearing Fixer Importance Threatened Khaya ivorensis African NX X X Mahogony Milicia excesla Iroko N X Pericopsis elata Afromosia N X X Albiza zygia Okuro N X Ceiba pentandra Onyina N X Mansonia altissima Oprono N X X Terminalia superba Ofram N X X Tetraplura tetraptera Prekese N Tieghenella heckelli Baku N X Tectona grandis Teak E X X at each forest reserve. Each individual plantation was sites in this study, in comparison to more time-inten- 40 m × 40 m with an initial planting distance of 1 m. sive quadrant counts, which are better suited to open Within each mixed native species plantation, trees of habitat types. The use of pitfall traps may underre- each species were randomly distributed. All seedlings present cryptic, soil-litter-dwelling ant species, were of local provenance and reared in the FORIG although they do occur in small numbers. nursery until approximately 3 months in age. A 4 m The contents of each trap were collected individu- buffer was left between treatments. ally and transported back to the lab for sorting, pre- servation, and preparation of museum vouchers. Ants were identified to species and placed into functional groups as adapted from Andersen (2007 and personal Ant sampling communication) and described in Table 3; identifica- Sampling of ants via pitfall traps was conducted dur- tions were based on Taylor (2007). Members of the ing July/August in 2006 and 2007, after the onset of genus Dorylus were excluded from this study because the short rainy season, at one and two years after they are highly mobile and do not establish long-term plantation establishment. Five pitfall traps were stationary colonies, and therefore are not associated placed on each of two parallel 25-m transects at 5- with long-term site characteristics. This behavior pre- m spacing, for a total of ten pitfall traps per treatment cludes them from being suitable bioindicators. replica. Transects were placed at the center of each Specimens were vouched by Dr. Brian Taylor and replicate plantation with a random start and direction have been deposited into the Northern Arizona within the stand. The traps were placed in two par- University School of Forestry voucher collection and allel transects instead of a single transect to reduce in the Forestry Research Institution of Ghana edge effects. insectarium. A pitfall trap consisted of a plastic deli style con- tainer (11 cm diameter by 8 cm deep) buried flush to ground level. During the 48-hour sampling periods, Ant species diversity and functional groups traps were set flush with the soil surface and with approximately 250 ml of soapy water in the trap. Any The traditional diversity measures of species rich- debris inside the trap was carefully removed when the ness (R), the Shannon-Wiener Diversity Index (H), traps were set. Preliminary trap testing conducting in and the Simpson Dominance Index (D) were calcu- the field ranged from 24 hours to 10 days. Analysis of lated for each treatment. Species richness is the these trap catches showed a 48-hour trapping period number of species observed. The Shannon-Wiener allowed for sufficient ant capture and reduced the Diversity Index was calculated as a measure of rela- risk of trap disturbance and flooding, especially dur- tive diversity per treatment type per year. The ing the rainy season. In addition, species richness Simpson Dominance Index was used to determine curves were optimized at between six and eight the strength of species numerical abundance. These traps (unpublished data). traditional diversity measures fail to assess species Pitfall trapping has been widely used in studies of assemblages or groups with particular traits, so spe- Australian ant communities (Andersen 1995) and has cies were also grouped into functional groups for been shown to provide a reliable estimate of species further analysis. Functional groups were determined composition (Andersen 1991; Agosti et al. 2000). according to behaviors, habitat preferences, and Andersen (1991) showed that pitfall trapping for trophic level interactions, using existing functional ants was an adequate collection method for habitats group assignments developed by Andersen (2007 with dense litter and vegetation as occurred at the and personal communication). INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 99 Table 3. Classification of ant functional groups collected from the semi-deciduous forest zone of Ghana, West Africa. Functional Species Species Groups (code) Characteristics Species Code Species Code Subordinate Ecologically separated due to large body size Camponotus sp. CAMY Camponotus furvus CAFU Camponotini and nocturnal foraging. Several arboreal (Myrmacrhaphe) (SC) nesters. Camponotus acvapimenis CAAC Camponotus maculatus CAMA Camponotus CAFL flavomarginatus Tropical Climate Distribution centered on tropical climatic zone. Aenictus guineensis AEGU Pristomyrmex orbiceps PROR Specialist (TCS) Unspecialized ants. Aenictus species 2 AESP Cryptic Species Forage and nest primarily within soil and litter, Cerapachys nitidulus CENI Pyramica cryptura PYCR (CS) having relatively little interaction with epigaetic Hypoponera HYCA Pyramica membranifera PYME ants cammerunensis Oligomyrmex diabolus OLDI Strumigenys petiolata STPE Oligomyrmex vorax OLVO Strumigenys rufobrunea STRU Opportunist (O) Unspecialized, poorly competitive species, Cardiocondyla emeryi CAEM Tetramorium angulinode TEAN characteristic of disturbed sites Cardiocondyla neferka CANE Tetramorium calinum TECA Decamorium decem DEDE Tetramorium flavithorax TEFL Odontomachus troglodytes ODTR Tetramorium intonsum TEIN Lepisiota cacozela LECA Tetramorium minimum TEMI Lepisiota laevis LELA Tetramorium sericeiventre TESE Paratrechina albipes PAAL Tetramorium sillimimum TESI Paratrechina arlesi PAAR Tetramorium quadridentatum TEQU Tapinoma lugubre TALU Generalist (G) Ubiquitous species occurring in most habitats. Crematogaster censor CRCE Pheidole concinna PHCO Rapid recruitment to, and successful defense of Crematogaster zavattarii CRZA Pheidole costauriensis PHCOS food resources. Predominate moderately Monomorium bicolor MOBI Pheidole impressifrons PHIM productive habitats. Monomorium exiguum MOEX Pheidole megacephala PHME Monomorium pharaonis MOPH Pheidole melancholica PHMEL Monomorium rosae MORO Pheidole nigritella PHNIG Monomorium tanysum MOTA Pheidole occipitalis PHOC Monomorium trake MOTR Pheidole pusilla PHPU Monomorium vaguum MOVA Pheidole retronitens PHRE Pheidole aeberlii PHAE Pheidole schoutedeni PHSC Pheidole aurivillii PHAU Pheidole speculifera PHSPE Pheidole bayeri PHBA Pheidole weissi PHWE Pheidole caffra PHCA Specialized Medium to large bodied strictly predaceous Anochetus bequaerti ANBE Pachycondyla silvestrii PASI Predators (SP) species. Well-developed sight, highly active Cerapachys sudanensis CESU Pachycondyla caffraria PACA foragers mostly solitary. Little competitive Leptogenys conradti LECO Pachycondyla tarsata PATA interactions with other ants behaving more like Pachycondyla pachyderma PAPA other predatory arthropods. Adapted from Andersen (1997b, 2007). Dominant Dolichoderine, Cold and Hot Climate Specialist functional groups removed, none were expected or observed. Dorlyus spp. were excluded from the study because of their characteristic nesting and foraging behavior. Statistical analyses treatments, we applied a two-sample t-test after meeting assumptions of normality and equal var- Prior to any statistical interpretation of the ant data, iance (Hays 1963). we decided to pool the trap collections across treat- ment replicate to obtain one value per treatment replicate per year, because we were primarily inter- Functional group analysis ested in observing indicator species and guild Multiple response permutation analyses (MRPP) was responses to treatment. Effect of treatment was eval- performed for the relative abundance measures of uated in two separate analyses: (1) comparing planta- each ant functional group to test the null hypothesis tions with different levels of diversity of native species of no difference in functional group composition (1, 6, and 10 native species; Table 1), and (2) compar- among treatments (McCune & Mefford 2006). ing native African species (all plantations) with exotic Indicator species value analysis (referred to as indi- teak monocultures. We used data from each year to cator value) by the Dufrene and Legendre (1997) determine if changes in communities occurred over method was performed to determine the specific con- time. tributions of each species to the functional group and We applied standard analysis of variance to determine the strength of indication of treatment (ANOVA) tests to determine differences between provided by either a functional group or an indivi- traditional diversity indices in overstory tree diver- dual species (Stephens & Wagner 2006). sity treatments using JMP In 5.1.2 (SAS 2004)after meeting assumptions of normality and equal var- iance. Nonparametric Welch ANOVA was applied Results when standard ANOVA assumptions were not met (SAS 2004). To determine differences between spe- A total of 7473 specimens were collected representing cies diversity indices for the native vs. exotic six subfamilies, 22 genera and 65 species (Table 4). 100 S. S. STEPHENS ET AL. The number of specimens collected per treatment p = 0.01). Species richness was highest in the planta- tions of six native timber species (R = 19.75 ±SE 1.48) was similar and ranged from 746 to 1262. The great- est number of species was trapped in the genus and lowest in the native African mahogany mono- Pheidole (16 species). The most commonly trapped culture (R = 13.25 ±SE 1.48) and (Figure 1). Shannon-Weiner Diversity Index (F = 2.28; d.f. = 4, ants were members of Pheidole, represented by three to six species per treatment. The most frequently 15; p = 0.1) was highest in the plantations of six observed ant was Odontomachus troglodytes native timber species (H = 2.22 ±SE 0.24) and lowest in native African mahogany monocultures (H = 1.38 Santschi, which represented 17–55% of ants collected per treatment. Composition of ant communities by ±SE 0.24) (Figure 2). Simpson’s Dominance Index functional group varied little between treatments. (F = 1.64; d.f. = 4, 15; p = 0.2) was highest in the plantations of six native timber species (D = 0.83 ±SE Results from two years after planting showed increases in ant species richness and nonsignificant 0.09) and lowest in native African mahogany mono- differences in ant community composition. cultures (D = 0.54 ±SE 0.09) (Figure 3). Ant species diversity and overstory tree species Ant species diversity and native species vs. teak diversity overstories At one year after planting, there were no significant At one year after planting, there were no significant differences in measures of ant species diversity in differences between the plantations of native African relation to overstory diversity of native tree species species and teak in measures of ant species diversity, (Table 4). Insignificant measures included species including species richness (native species R = 11.25; richness (F = 0.96; d.f. = 4, 15; p = 0.5), Shannon- exotic teak R = 8.75) (t = −1.43, d.f. = 18, p = 0.1); Weiner Diversity Index (F = 0.82; d.f. = 4, 15; Shannon-Weiner Diversity Index (t = −1.24, d.f. = 18, p = 0.5), and Simpson’s Dominance Index p = 0.1); or Simpson’s Dominance Index (t = −1.26, (F = 0.76; d.f. = 4, 15;p = 0.6) (Table 4). At two d.f. = 18, p = 0.2) (Table 4). However, at two years years after planting species richness was significantly after planting, species richness (t = −2.54, d.f. = 18, different between treatments (F = 4.60; d.f. = 4, 15; p = 0.02) was significantly higher in the plantations of Table 4. Composition of ant genera by functional group. Data are number of species per genera, with percent total ants per treatment per year (column) in brackets. Native African mahogany Six native timber species Ten native timber species Functional group Exotic teak monoculture monoculture mixture mixture Genera 2006 2007 2006 2007 2006 2007 2006 2007 Subordinate Camponotini Camponotus 3 (15.67) 1 (2.39) 2 (5.74) 1 (4.77) 4 (5.14) 1 (1.27) 3 (7.91) 1 (4.83) Tropical Climate Specialist Aenictus 1 (0.13) Pristomyrmex 2 (0.22) Cryptic Species Cerapachys 1 (0.26) 1 (0.10) 1 (0.18) 1 (0.08) Hyperponera 1 (0.16) Oligomyrmex 1 (0.11) 1 (0.10) 1 (0.51) Pyramica 1 (0.10) 1 (0.36) 1 (0.16) Strumigenys 1 0.13) 1 (0.20) 1 (0.12) 1 (0.73) 1 (0.13) 1 (0.24) Oppurtunist Tapinoma 1 (15.52) 1 (4.27) 1 (9.44) 1 (8.45) 1 (12.60) Lepisiota 2 (5.41) 1 (1.33) 2 (7.42) 1 (1.32) 2 (8.30) 2 (6.09) 2 (5.23) 2 (2.46) Paratrechina 1 (4.96) 2 (9.81) 2 (3.04) 1 (1.42) 1 (1.64) 2 (3.00) 1 (3.89) 2 (1.35) Cardiocondyla 1 (3.15) 1 (0.56) 1 (0.79) 2 (0.35) 1 (4.00) 1 (3.25) Decamorium 1 (0.11) Tetramorium 1 (1.35) 2 (1.86) 3 (3.26) 3 (0.81) 7 (5.36) 3 (2.06) Odontomachus 1 (55.02) 1 (24.80) 1 (17.32) 1 (46.90) 1 (41.82) 1 (25.18) 1 (45.04) 1 (23.53) Generalist Crematogaster 1 (1.62) 1 (0.64) 1 (0.13) 1 (1.23) Monomurium 2 (3.16) 3 (1.59) 1 (1.46) 2 (0.71) 1 (1.99) 4 (2.65) 4 (2.68) 5 (2.38) Pheidole 5 (2.37) 3 (27.19) 6 (34.53) 2 (19.19) 5 (22.43) 4 (24.45) 4 (19.97) 3 (34.15) Specialized Predators Cerapachys 1 (0.10) Anochetus 1 (0.34) 1 (0.20) 1 (0.09) 1 (0.08) Leptogenys 1 (0.11) 1 (0.10) Pachycondyla 2 (11.95) 2 (14.19) 2 (21.60) 2 (9.64) 2 (12.97) 3 (16.82) 2 (15.01) 3 (11.33) Total Individuals 887 754 889 985 856 1100 746 1262 Total # of species 18 20 25 24 18 33 19 29 Mean R (SE) 8.75 (1.25) 11.75 (1.48) 11.25 (1.25) 13.25 (1.48) 9.50 (1.25) 19.75 (1.48) 10.25 (1.25) 17.00 (1.48) Mean H (SE) 1.36 (0.27) 1.84 (0.24) 1.76 (0.27) 1.38 (0.24) 1.47 (0.27) 2.22 (0.24) 1.67 (0.27) 1.94 (0.24) Mean D (SE) 0.58 (0.11) 0.78 (0.09) 0.75 (0.11) 0.54 (0.09) 0.63 (0.11) 0.83 (0.09) 0.70 (0.11) 0.78 (0.09) INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 101 AB BC Native Species Six Native Ten Native Teak All Native Monoculture Species Plantations Species Plantations Monoculture Species Plantations (a) Overstory Tree Species Diversity (b) Exotic Vs. Native Species Figure 1. Ant species richness (R) comparisons with standard error bars in 2-year-old plantations for (a) overstory tree species diversity, and (b) teak monocultures s. all native species plantations. Significance shown at α = 0.05. Bars with different letters are statistically different based on a standard mean separation test. 2.5 2.5 AB AA 1.5 1.5 0.5 0.5 Teak All Native Species Native Species Six Native Species Ten Native Species Monoculture Plantations Monoculture Plantations Plantations (b) Exotic Vs. Native Species (a) Overstory Tree Species Diversity Figure 2. Shannon Weiner Index (H) comparisons with standard error bars for ant species in 2-year-old plantations for (a) overstory tree species diversity, and (b) teak monocultures vs. all native species plantations. Significance shown at α = 0.05. Bars with different letters are statistically different based on a standard mean separation test. AB 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 Native Species Six Native Species Ten Native Species Teak All Native Monoculture Plantations Plantations Monoculture Species Plantations (a) Overstory Tree Species Diversity (b) Exotic vs. Native Species Figure 3. Simpsons Dominance Index (D) with standard error bars for ant species in 2-year-old plantations for (a) overstory tree species diversity, and (b) teak monocultures vs. all native species plantations. Significance shown at α = 0.05. Bars with different letters are statistically different based on a standard mean separation test. native species (R = 13.25) than in the exotic teak among the one, six, and 10 species treatment, nor plantations (R = 11.75) (Figure 1). between the native African species and teak plantations (α = 0.05, Table 5; Figures 4 and 5). The A statistic for the MRPP comparisons of both overstory diversity and Functional group analysis native vs. teak plantations was very close to zero for both ant functional groups and individual species in During both the 2006 and 2007 collection periods, both years (Table 5), indicating that the heterogeneity neither ant functional group composition nor indivi- within groups was equal to that expected by chance. dual ant species composition differed significantly at (D) (H) (R) (R) (H) (D) 102 S. S. STEPHENS ET AL. Table 5. Multiple response permutation procedure (MRPP) to determine ant functional group and individual species relative abundance response to treatment per year. Individual species Functional groups Teak vs. all native plantations Overstory diversity Teak vs. all native plantations Overstory diversity 2006 2007 2006 2007 2006 2007 2006 2007 Test statistic T 1.32 −0.01 1.51 0.45 −0.36 −0.96 0.81 −1.12 Observed Delta 138.72 120.43 143.53 123.67 114.49 119.31 119.57 114.55 Expected Delta 135.08 120.48 135.08 120.48 115.41 122.76 115.41 122.76 A* −0.03 0.0003 −0.06 −0.02 0.007 0.02 −0.04 0.06 p(α = 0.05) 1.00 0.38 0.97 0.62 0.29 0.15 0.78 0.13 *A statistic chance corrected within group agreement, A(max) = 1 when all items are identical within groups. A = 0 when heterogeneity within groups is equal to expected by chance. A < 0 when heterogeneity is greater within groups than expected by chance. Small value for A indicates that groups are appropriately assigned, members within groups are more similar to each other than to members of other groups. p-value indicates probability of smaller or equal delta. 100% 100% 80% 80% CS CS 60% TCS 60% TCS SC SC SP SP 40% G 40% 20% 20% 0% Native African Ten Native Six Native 0% Mahogany Species Species Teak All Native Species Monoculture Plantations Plantations Monoculture Plantations (a) Overstory Tree Species Diversity (b) Exotic vs. Native Species Figure 4. Ant functional group composition 1 year after planting for (a) overstory tree species diversity, and (b) teak monocultures vs. all native species plantations. Different ratios of column fills indicate differences in functional groups composition – greater dissimilarity between columns equals greater dissimilarity between ant communities. TCS – tropical climate specialist, SP – specialized predators, SC – subordinate camponotini, O – opportunist, G – generalist, CS – cryptic species. While no indicator species or groups were identi- mixed native species plantations gained 10–15 species fied for any treatments, the Opportunist functional between years. We observed only small, nonsignificant group dominated all of the treatments in both years, changes in the proportions of functional group in each representing 36–67% of the ants collected in each treatment (Figure 7). Subordinate Camponotini species treatment. The Generalist functional group was typi- declined in all treatments, Cryptic species increased in cally the second most common functional group in all treatments, and Specialized Predators increased in both years representing an average of 26% of ants the plantations of one, six, and 10 native tree species collected in each treatment. Specialized predators and and declined in exotic teak monocultures. Opportunists Subordinate Camponotini averaged 13% and 11%, increased in the exotic teak monocultures and native respectively. Tropical climate specialist and Cryptic mixed species plantations and decreased in native species were very seldom observed and contributed to African mahogany monocultures. Generalists increased less than 2% of the ants collected from any treatment or remained unchanged in mixed native species planta- (Table 4, Figures 4 and 5). tions and decreased in the teak and African mahogany monocultures. Changes over time Discussion Over the 2 years of the study, we observed a significant change in species richness (F = 53.22, d.f. = 8, 30, Over the two years of the study, we observed a 20% p= < 0.0001) in all treatments (Figures 1 and 6). The increase in species richness across all treatments exotic teak and native African mahogany monocultures (Figure 6). If current trends persist, we predict species gained two and lost two species between years. The diversity will become stable over time. As supported Percent Composition (%) Percent Composition (%) INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 103 100% 100% 90% 90% 80% 80% 70% 70% CS CS 60% 60% TCS TCS SC 50% SC 50% SP SP 40% 40% 30% 30% 20% 20% 10% 10% 0% 0% Native African Six Native Ten Native Teak All Native Species Mahogany Species Species Monoculture Plantations Monoculture Plantations Plantations (a) Overstory Tree Species Diversity (b) Exotic vs. Native Species Figure 5. Ant functional group composition 2 years after planting for (a) overstory tree species diversity, and (b) teak monocultures vs. all native species plantations. Different ratios of column fills indicate differences in functional groups composition – greater dissimilarity between columns equals greater dissimilarity between ant communities. TCS – tropical climate specialist, SP – specialized predators, SC – subordinate camponotini, O – opportunist, G – generalist, CS – cryptic species. (Piper et al. 2009). Anderson (1991) indicated that ants Native Species are associated with structural components of their Monoculture 6 Native Species habitats. Peck et al. (1998) saw significant changes in 10 Native Species ant communities based on vegetation community diversity and structure. Disturbance resulting from fire and timber management has been observed to affect ant communities (Neumann 1992;Andrew et al. 2000;York 2000). Changes in ant diversity and overstory composition 1 year 2 year Overstory tree diversity had a positive impact on ant Time After Planting species diversity at two years after plantation estab- Figure 6. Change in ant species richness one and two years lishment (Figures 1–3, and Table 4), although not at after planting for (a) overstory tree species diversity, and (b) one year after planting. Increasing overstory tree teak monocultures vs. all native species plantations. One year diversity from one to six species resulted in a 30% after planting there were no differences in species richness among treatments. 2 years after planting all treatments increase in ant species richness. The species present showed an increase in species richness which differed for in the six and ten native timber species plantations both overstory diversity and between teak monoculture and increased by 15 and 10 species respectively between the native species plantations (see also Figure 1). the first and second years (Table 4). This suggests that, within plantation forests, moderate increases in by other observations (Stephens et al. 2008), we overstory tree diversity can have a significant effect expect distinct ant communities will develop in on ant biodiversity, and perhaps on other taxa. these plantation types, which may include specific Native environments typically support higher num- indicator species or functional groups. ber of insect species than non-native environments This study highlights the potential of ants as bioin- (DeGomez & Wagner 2001). In this study, the native dicators of habitats conditions in tropical forested species plantations had approximately 20% more ant systems, where they have been generally underutilized species than the exotic teak plantation over both years Species Richness Percent Composition (%) Percent Composition (%) 104 S. S. STEPHENS ET AL. CS TCS SC SP –2 –4 Exotic Species Native Species Six Native Timber Ten Native Timber Monoculture Monoculture Species Plantations Species Plantations Treatment Figure 7. Changes in number of ant species representing each functional group over time for exotic teak and native African mahogany monocultures, 6 and 10 native species mixed plantations. Positive bars indicate an increase in species per functional group, negative bars indicate a decrease in the number of species per functional group. TCS – tropical climate specialist, SP – specialized predators, SC – subordinate camponotini, O – opportunist, G – generalist, CS – cryptic species. among plantation types in ant diversity indicators (Table 4). At two years, the native species plantations were detectable at only twoyears after plantation had significantly higher species richness teak planta- tions. A nonsignificant difference in species composi- establishment. Other work (Stephens et al. 2008) assessed ants as indicators of land use in Ghana at tion was also observed (Table 4) with six genera being 10–13 years after planting and found distinct patterns unique to the native African species overstories and in communities of ants and other organisms among only one genera unique to teak plantations. Since teak agriculture, citrus orchards, exotic species planta- is a major plantation species in West Africa and else- tions, mixed native species plantations, and native where (FAO 20001), the lower associated ant diversity forests. The plantations were similar in species com- in this study is of concern. position to the 10 native timber species plantation and teak monocultures used in this study. The agri- Ant functional groups and succession cultural areas and citrus orchards were also similar to the degraded conditions prior to plantation establish- Whereas traditional diversity measures are based on ment for this study at the Forest Reserve sites. numerical counts, functional group analysis is based Furthermore, indicator species and functional groups on differences in characteristics among species were identified for the more mature land uses groups. Unlike traditional diversity measures, func- (Stephens et al. 2008). These observations support tional group analysis permits examination of how the conjecture that the newly established plantations species habitat requirements and competitive interac- of this study are in the early stages of establishment tions may impact diversity patterns (Anderson 1991). for distinct ant communities. In this study, the relative abundances of each func- In Years 1 and 2 under different overstory types, the tional group were similar among treatments espe- ant communities were generally dominated by cially at one year after planting (Figures 4 and 5), Opportunists and Generalists (Figures 4 and 5). We with MRRP A values indicating high homogeneity have observed that Opportunists tend to move into dis- between treatments (Table 5). However, by Year 2, turbed habitats (Stephen et al. unpublished), whereas we see a trend of increasing heterogeneity in ant Generalists seem indifferent to successional stage. In functional groups among treatments in MRRP A this study, between Years 1 and 2, Opportunists values (although still insignificant) (Table 5). increased (except for the native African mahogany This study may have presented the early stages of monoculture) and Generalists showed no pattern establishment of distinct ant communities under dif- (Figure 7). In another study (Stephen et al. unpublished), fering overstory composition. As the treatment areas a similar relationship between Opportunist and recover from forest degradation, disturbance, and Generalist functional groups and different land uses plantation establishment (site prep, planting, and was observed, with Opportunists dominating agricultural early site maintenance), and their composition and fields. We anticipate future changes from Opportunist- structure become more distinct, more distinct ant dominated ant communities to more functional group communities may emerge. Significant differences Change in Number of Species INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 105 changes in understory vegetation and leaf litter will diversity and stronger links to overstory tree diversity support different ant communities. For example, as and composition. Nonsignificant shifts in functional group composi- individual trees increase in bole diameter and height during stand succession, several species of arboreal tion between Year 1 and Year 2 include decreases in nesters would be expected to appear, including car- the subordinate Camponotini (SC) and increases in the Cryptic species functional groups in all treat- ton-nesting Crematogastor spp. and Camponotus spp. (Holldobler & Wilson 1990). ments. The SC group was composed largely of arbor- Future research is required to fully assess the eal Camponotus species strongly associated with downed woody debris (Higgins & Lingren 2006; stages of stand development and management that are most influential on ant communities. As these Stephens & Wagner 2006). The Camponotus species plantations mature and are managed over time, that were found in Year 1 may have been residual populations using coarse woody debris and the few some tree species are likely to be thinned or har- small shrubs remaining from before overstory vested to meet management objectives. This would present the opportunity to assess how those changes removal and plantation establishment. By Year 2, since woody material on the treatment sites had les- in stand structure and diversity further impact ant sened and the young trees were not of sufficient size communities. A longer period of observation over to provide suitable habitat, Camponotus species were 5–10 years may reveal the full sequence of ant probably being displaced by more pioneering species community assemblage establishment and the more adapted to disturbance and early successional appearance of indicator species and functional conditions. Camponotus species may re-enter these groups associated with overstory tree diversity and areas when the overstory trees mature and provide composition. Additional work could be conducted habitat. to assess the association of ant communities with In contrast to the SC group, Cryptic species such changes in understory vegetation, leaf litter, and soil as Strumigenys species, specialize in predation on leaf characteristics. litter dwelling microinvertebrates (Holldobler & Wilson 1990). The increase in Cryptic species between Years 1 and 2 was likely a response to Conclusion changes in understory and soil conditions after removal of agricultural crops and plantation estab- Biodiversity is clearly an important component of lishment that improved microhabitats for their prey. ecosystems, supporting their function (Tilman et al. The number of species in the Specialist Predators 1996) and their anthropocentric value (Wilson 1984). (SP) functional group also increased in all of the Conventional wisdom has long held that biodiversity native overstory treatments (Figure 7) from Year 1 and plantation forests are mutually exclusive to Year 2, perhaps as a consequence of understory (Stephens & Wagner 2007). Stephens and Wagner changes. For example, Cerapachys sp. are predators of (2007) argued the importance of land use compari- termites and other ants, particularly Pheidole sp. sons prior to passing judgment on plantation forest (Holldobler & Wilson 1990). Pheidole is a impacts on biodiversity. Observed cases in the litera- Generalist, which was readily abundant in treatments ture indicate that plantation management, can impact where Specialist Predators were observed (Table 3, biodiversity positively and negatively, depending on Figure 7). the strategies used. Based on the results of this study As these young plantations transition through suc- and other work (Stephens et al. 2008), the six native cessional stages of stand development, it seems likely tree species mixture at only two years after planting that distinct ant community patterns will emerge. recouped an astonishing 80% of the total number of Our experimental plantations with managed unders- ant species, and from 40% to 100% of the individual tories and open canopies may not yet have developed ant functional groups observed at native forest sites. all of the characteristics that promote specific ant The native African mahogany monoculture recouped functional groups or species. Regular weeding was 60% of native forest ant diversity. These numbers used to control the understory vegetation during the show that biodiversity and plantation forests are not early years of plantation development, and the tree necessarily at odds. Casual observation of soil, micro- species used in these plantations have different climate, understory plants, and birds, reptiles and growth characteristics (Table 2) and grew at signifi- mammals in the more mature mixed native species cantly different rates, all of which may have impacted and exotic teak plantations suggest that strong com- ant community composition and diversity. The munity difference are likely occur across numerous cumulative differences in the plantation types as the floral and faunal groups in different kinds of planta- stands develop will impact both the overstory and tions. Our findings suggest that moderate changes in understory ant communities. As canopy structure management, such as focusing on mixed native spe- matures, shading of the understory will increase and cies plantations, can recapture a significant level of 106 S. S. STEPHENS ET AL. and global change. 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International Journal of Biodiversity Science, Ecosystem Services & Management – Taylor & Francis
Published: Jan 2, 2016
Keywords: Formicidae; timber plantations; Khaya ivorensis; Tectona grandis; biodiversity
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