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AGRICULTURE AND BIOLOGY JOURNAL OF NORTH AMERICA ISSN Print: 2151-7517, ISSN Online: 2151-7525, doi:10.5251/abjna.2014.5.3.123.134 © 2014, ScienceHuβ, http://www.scihub.org/ABJNA Termite species composition in the central rift valley of Ethiopia Daniel Getahun Debelo 1 and Emana Getu Degaga 2 1 Department of Biology, Adama Science and Technology University, P. O. Box 1888, Adama, Ethiopia 2 Faculty of Life Sciences, Addis Ababa University, P. O. Box 1176, Addis Ababa, Ethiopia E-mail: [email protected] ; [email protected] ABSTRACT In Ethiopia, particularly in the Central Rift Valley (CRV) where high density of epigeal termite mounds is found, little systematic studies were done to know the termite fauna. Hence, the current study was conducted in three districts of East Shawa Zone of Oromia Regional State to know termite species composition in the CRV of Ethiopia from August to November 2013. Termites were sampled from different land-use types: protected lands, rangelands, and farmlands using the standardized belt transect and maize stalk baiting. Termites were also collected randomly from different places and habitats. The collected samples belonged to seven genera: Macrotermes, Odontotermes, Microtermes, Amitermes, Microcerotermes, Angulitermes, and Trinervitermes. Angulitermes and Trinervitermes were very rare and recorded only from protected lands. Protected lands housed all genera of the collected termites. Microcerotermes were the most abundant in protected lands, Macrotermes in rangelands, and Microtermes in farmlands. The results showed that land-use type determine termite abundance and diversity. The baiting method captured all genera of the collected termites, while, the standardized belt transect method and random collection methods captured 43% and 71% of the termite genera, respectively and this indicated that maize stalk baiting was highly suitable for sampling wood-based food feeders. The result of this survey study can form a baseline for designing a national termite species inventory. Keywords: Abundance, land-use, maize stalk baiting, species diversity, standardized method. INTRODUCTION Termites are social insects of the order Isoptera with about 3000 species (Munthali, 1999; Glaciela et al., 2006; Tathiane et al., 2009) in 281 genera (Grohmann et al., 2010), fifteen subfamilies and seven families (Logan et al., 1990; Pearce, 1997). Termites are predominantly distributed in tropical environment, with the highest species richness in equatorial rainforest, and generally declining with increasing latitude (Yanyong et al., 2003). Termites are dominant arthropod detritivores important in decomposition process. Their influence on decomposition processes at any site is likely to be governed to a large extent by the species composition and the local termite assemblage (Jones and Eggleton, 2000). Termite diversity, composition and their associated functions vary within and between ecosystems and these may shift under changing land-uses (Zeidler et al., 2002). Many of the records of termite damage on crops lack the necessary information regarding the stage of crops attacked and the extent of damage caused by the different species. Some of the studies even did not report the name of the species responsible for the crop damage (Abdurahman, 1990). Detailed knowledge of local species diversity as well as the diversity of functional groups of termites is a prerequisite for evaluating and quantifying the type and extent of their functional roles in ecosystem processes. Such information is vital to understand the economic importance of the species found in an area, so that studies on their biology and control may be directed at the most important species (Abdurahman, 1990; Mugerwa et al., 2011). High density of epigeal termite mounds is a peculiar feature of the Maki-Batu area which is part of the CRVE (Central Rift Valley of Ethiopia). However, no systematic termite inventory was available. Knowledge is also lacking on the impact of different land-use practices (anthropogenic activities) on termite diversity. Therefore, the major objective of this study was to assess termite species composition and the effect of different land-use practices on the

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Page 1: Termite Species Composition in the Central Rift Valley of ... · High density of epigeal termite mounds is a peculiar feature of the Maki-Batu area which is part of the CRVE (Central

AGRICULTURE AND BIOLOGY JOURNAL OF NORTH AMERICA ISSN Print: 2151-7517, ISSN Online: 2151-7525, doi:10.5251/abjna.2014.5.3.123.134

© 2014, ScienceHuβ, http://www.scihub.org/ABJNA

Termite species composition in the central rift valley of Ethiopia

Daniel Getahun Debelo1 and Emana Getu Degaga2

1Department of Biology, Adama Science and Technology University, P. O. Box 1888, Adama, Ethiopia

2 Faculty of Life Sciences, Addis Ababa University, P. O. Box 1176, Addis Ababa, Ethiopia E-mail: [email protected]; [email protected]

ABSTRACT

In Ethiopia, particularly in the Central Rift Valley (CRV) where high density of epigeal termite

mounds is found, little systematic studies were done to know the termite fauna. Hence, the current study was conducted in three districts of East Shawa Zone of Oromia Regional State to know termite species composition in the CRV of Ethiopia from August to November 2013. Termites were sampled from different land-use types: protected lands, rangelands, and farmlands using the standardized belt transect and maize stalk baiting. Termites were also collected randomly from different places and habitats. The collected samples belonged to seven genera: Macrotermes, Odontotermes, Microtermes, Amitermes, Microcerotermes, Angulitermes, and Trinervitermes. Angulitermes and Trinervitermes were very rare and recorded only from protected lands. Protected lands housed all genera of the collected termites. Microcerotermes were the most abundant in protected lands, Macrotermes in rangelands, and Microtermes in farmlands. The results showed that land-use type determine termite abundance and diversity. The baiting method captured all genera of the collected termites, while, the standardized belt transect method and random collection methods captured 43% and 71% of the termite genera, respectively and this indicated that maize stalk baiting was highly suitable for sampling wood-based food feeders. The result of this survey study can form a baseline for designing a national termite species inventory.

Keywords: Abundance, land-use, maize stalk baiting, species diversity, standardized method.

INTRODUCTION

Termites are social insects of the order Isoptera with about 3000 species (Munthali, 1999; Glaciela et al., 2006; Tathiane et al., 2009) in 281 genera (Grohmann et al., 2010), fifteen subfamilies and seven families (Logan et al., 1990; Pearce, 1997). Termites are predominantly distributed in tropical environment, with the highest species richness in equatorial rainforest, and generally declining with increasing latitude (Yanyong et al., 2003).

Termites are dominant arthropod detritivores important in decomposition process. Their influence on decomposition processes at any site is likely to be governed to a large extent by the species composition and the local termite assemblage (Jones and Eggleton, 2000). Termite diversity, composition and their associated functions vary within and between ecosystems and these may shift under changing land-uses (Zeidler et al., 2002).

Many of the records of termite damage on crops lack the necessary information regarding the stage of

crops attacked and the extent of damage caused by the different species. Some of the studies even did not report the name of the species responsible for the crop damage (Abdurahman, 1990). Detailed knowledge of local species diversity as well as the diversity of functional groups of termites is a prerequisite for evaluating and quantifying the type and extent of their functional roles in ecosystem processes. Such information is vital to understand the economic importance of the species found in an area, so that studies on their biology and control may be directed at the most important species (Abdurahman, 1990; Mugerwa et al., 2011).

High density of epigeal termite mounds is a peculiar feature of the Maki-Batu area which is part of the CRVE (Central Rift Valley of Ethiopia). However, no systematic termite inventory was available. Knowledge is also lacking on the impact of different land-use practices (anthropogenic activities) on termite diversity. Therefore, the major objective of this study was to assess termite species composition and the effect of different land-use practices on the

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termite diversity and abundance in the CRVE by sampling termites using standardized belt transect, maize stalk baits and random/casual collection.

MATERIALS AND METHODS

Study area description: The study was conducted in three land-use types: farmland, rangeland, and protected land in three districts of East Shawa Zone

of Oromia Regional State (Figure 1) from August to November 2013 after the long rainy season when termite activity was at peak. From each district 1-3 Farmers’ Associations and in total 18 study sites were selected. The districts are located in the Central Rift Valley of Ethiopia and they have similar climate. More descriptions of the districts are presented in Table 1.

Table 1 Coordinates, elevations, mean annual rainfall and temperatures of the study sites

District Coordinates Elevation (m)

Mean annual rainfall (mm)

Mean annual temperature (°C)

Maximum Minimum

Bora 8°29'N, 38°91E 1656 814.5 28.3 13.0

Dugda 8°15N, 38°82'E 1662 762.8 27.8 14.4

ATJK 7°56'N, 38°43'E 1650 742.4 27.2 14.0

Source: National Meteorological Agency of Ethiopia ATJK = Adami Tulu Jido Kombolcha

Fig. 1 Map showing study sites in three districts of Eastern Shawa Zone of Oromia Regional State

The area has a semi-arid and arid climate (Mengistu, 2008) and has a bimodal rainfall patterns. Highest precipitation occurs between July and September. There is some additional rainfall between February and the end of April, but this usually varies. Rain-fed crops are most often not planted until mid June because of the unpredictable rainfall pattern in the preceding period (Haimanot, 2002). Generally, the soils of the area range from sandy to loam, loam to sandy clay loam with some clay loam and a few clay soils. In addition to salt crust (white and deep shiny black salt crust), the soils are characterized by

numerous gray-to-gray brown termite mounds (Kefayalew, 2008) which belong to the genus Macrotermes (Getachew et al., 2003). The vegetation is characterized by acacia open-woodland and savanna (Huib and Herco, 2006).

Different types of field and pulse crops are grown and these were teff, wheat, barley, millet, maize, sorghum, faba bean, peas, and haricot bean. Maize (Zea mays), haricot bean (Phaseolus vulgaris L.), teff (Eragrostis teff L.), wheat, barley and sorghum were the major crops widely grown by small scale farmers in the area (Source: East Shawa Zone of Agricultural Development Office).The majority of the farmers are

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engaged in rain-fed agriculture. The farmers who are situated along the water sources have traditionally practiced irrigation. In recent years irrigation has substantially increased, mainly because of moisture stress during the growing seasons (Haimanot, 2002).

Termite sampling and identification: Termites were sampled using the standardized belt transect and maize stalk baiting methods. To maximize the number of species recorded to represent the local termite fauna, termites were also collected randomly/casually from different places and habitats.

The standardized belt transect method was used as described by Jones and Eggleton (2000). It was especially designed for a rapid assessment of termite species richness and functional diversity in tropical forests. However, as the vegetation of the area in the current study was of open woodland savanna type ecosystem dominated by agriculture, it was believed that the method might not be suitable for sampling the existing local termite fauna to the maximum extent. Accordingly, a maize stalk baiting technique was devised by the researcher of this study during the study period as described below. In each site, one transect (100 m x 2 m) was run and divided into 20 sections (5 m x 2 m). One 20 cm long maize stalk used as termite bait was half-buried at about 2 m away from each section of the standardized belt; a total of twenty maize stalk baits were used for each transect. Twenty-eight days later, termites were sampled from each transect and simultaneously the maize stalk baits were excavated and checked for termites and when present termites were sampled and preserved in 80% ethanol labeled vials for later identification.

The standardized belt transect and maize stalk bait sampling methods were replicated at six sites for each land-use type. Termite species recorded by the two methods from the six replicates of each land-use type were combined and regarded as the termite species occurred in that specific land-use type.

Whereas, all the termite species recorded in all the land-use types together with the ones sampled by random/casual collection, were considered as the termite species composition (fauna) of the area. The number of encounters with termites of a given genus within a land-use type was taken as a relative abundance of the genus within that land-use-type.

Taxonomic identification was done at the genus level using Keys to the Genera of Ethiopian Termites based on soldier characters (Abdurahman, 1991). Voucher specimens from this study are deposited in the laboratory of the Department of Biology, Adama Science and Technology University.

Data analysis: Descriptive statistics and cross tabulation were used to summarize collected data. Termite species encounter from the six sites of each land-use type were analyzed using the SAS software (SAS Institute Inc., 2003). One-way analysis of variance (ANOVA) was performed and the Least Significant Difference (LSD) test at 5% was used to separate significant means. Data were square root transformed (X + 0.5)

1/2 before analysis (Gomez and

Gomez, 1984).

RESULTS

Species composition: In total, 105 termite samples were collected and of these 61 were collected by standardized belt transect and maize stalk baiting methods from the three land-use types, while 44 samples were collected randomly. Overall, termites belonging to seven genera representing one family (Termitidae) and three sub-families were identified from the total collections of the study area. The combined collected termite species from all the sites are shown in Table 2. All the genera recorded from the area were also recorded from protected lands. Trinervitermes and Angulitermes were very rare and collected only by maize stalk baiting method, the former from one site and the latter genus from two sites only, from protected (Plates 1-3).

Table 2. Termite species recorded and their occurrences within the three land-use types from the combined standardized, baiting and random sampling methods

Subfamily Land-use type/Termite genera

Protected land Rangeland Farmland

Macrotermitinae Macrotermes Microtermes Odontotermes

Macrotermes Microtermes

Macrotermes Microtermes Odontotermes

Termitinae Amitermes Microcerotermes Angulitermes

Amitermes Microcerotermes

Amitermes Microcerotermes

Nasutitermitinae Trinervitermes

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Plate 1 Protected area by Maki-Batu Fruit and Vegetables Growers Cooperative Union Ltd. from where the rare

Angulitermes was sampled. Photo: Daniel Getahun (October 2013).

Plate 2 Warja Hill: A protected area from human activities and domestic animals from where only Trinervitermes was sampled. Photo: Daniel Getahun (October 2013).

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Plate 3 Adami Tulu Agricultural Research Center: A protected land from human activities and domestic animals, from where six of the seven termite genera were sampled. Photo: Daniel Getahun (October 2013).

Plate 4 Macrotermes mounds without opening

Relative abundance of termites: The relative abundance based on the number of encounters of each genus within land-use types is given in Table 3. A total of 61 termite occurrences were recorded in the belt transect and baiting methods. Macrotermes occurred 20 (32.8%) times out of the 61 total

occurrences and 10 (16.4%) in rangeland out of its total 20 occurrences. Macrotermes, Microcerotermes, Amitermes, and Microtermes occurred in all of the land-use types. Macrotermes were the most abundant in rangelands, Microtermes in farmlands, and Microcerotermes in protected lands.

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Angulitermes and Trinervitermes were only sampled from protected lands, while Odontotermes were sampled only from farmland. However, Odontotermes was sampled in two protected lands from constructed soil sheeting on bark of trees (Adami Tulu Agricultural Research Center and Maki-Batu Fruit and Vegetables Growers Cooperative Union Ltd.) by random sampling. Macrotermes mounds of both

closed and open type of different forms were recorded (Plates 4A-C and 5).

Macrotermes occurrence was significantly (P < 0.05) higher in protected land and rangeland than in farmland (Table 4). However, there was no significance difference (P > 0.05) of occurrence of the other species in each land-use type.

Table 3 Percent (n) encounters/occurrences of termite genera in three land-use types sampled by standardized belt transect and baiting methods in the Central Rift Valley of Ethiopia in the year 2013.

Termite genera Percent (n) termite genera encounters in Total encounters

Protected land Rangeland Farmland

Macrotermes 9.8 (6) 16.4 (10) 6.6 (4) 32.8 (20)

Microtermes 4.9 (3) 11.5 (7) 11.5 (7) 27.9 (17)

Odontotermes 0.0 (0) 0.0 (0) 1.6 (1) 1.6 (1)

Microcerotermes 11.5 (7) 3.3 (2) 8.2 (5) 23.0 (14)

Angulitermes 3.3 (2) 0.0 (0) 0.0 (0) 3.3 (2)

Trinervitermes 1.6 (1) 0.0 (0) 0.0 (0) 1.6 (1)

Amitermes 4.9 (3) 3.3 (2) 1.6 (1) 9.8 (6)

Total 36.0 (22) 34.5 (21) 29.7 (18) (100) 61

Table 4 Mean (± SE) of individual termite species encounters in three land-use types

Land-use type

Macr. Micr. Odon. Microc. Angul. Trin. Amit.

Protected land

1.00 ± 0.26ab 0.50 ± 0.34a 0.00 ± 0.00a 1.17 ± 0.98a 0.33 ± 0.21a 0.17 ± 0.17a 0.50 ± 0.22a

Rangeland 1.67 ± 0.42a 1.17 ± 0.40a 0.00 ± 0.00a 0.33 ± 0.33a 0.00 ± 0.00a 0.00 ± 0.00a 0.33 ± 0.21a

Farmland 0.67 ± 0.21b 1.17 ± 0.65a 0.17 ± 0.17a 0.67 ± 0.33a 0.00 ± 0.00a 0.00 ± 0.00a 0.17 ± 0.17a

LSD 0.94 1.46 0.29 1.89 0.37 0.29 0.61

Means within a column followed by the same letter do not differ significantly by Least Significant Difference Test (LSD) at 5% of probability. Macr. = Macrotermes; Micr. = Microtermes; Odon. = Odontotermes; Microc. = Microcertermes; Angul. = Angulitermes; Trin. = Trinervitermes; Amit. = Amiterems; SE = Standard error

Even though the abundance of termite species recorded was not equal in unmanaged (protected land), statistically there was no significant difference (P > 0.05) among termite species encounters except Macrotermes were higher (P < 0.05) than Odontotermes. Occurrences of Macrotermes and

Microtermes in rangeland were significantly higher (P < 0.05) than the other species. Microtermes were the most frequently encountered species in farmlands (Table 3), although there was no significant difference (P > 0.05) between them and Macrotermes and Microcerotermes (Table 5).

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Table 5 Mean (± SE) encounters of all the termite genera in three land-use types sampled by standardized belt transect and baiting methods in the Central Rift Valley of Ethiopia in the year 2013.

Termite genera Protected land Rangeland Farmland

Macrotermes 1.00 ± 0.36a 1.67 ± 0.60a 0.67 ± 0.30ab

Microtermes 0.50 ± 0.49ab 1.17 ± 0.58a 1.17 ± 0.92a

Odontotermes 0.00 ± 0.00b 0.00 ± 0.00b 0.17 ± 0.24b

Microcerotermes 1.17 ± 1.39ab 0.33 ± 0.47b 0.67 ± 0.47ab

Angulitermes 0.33 ± 0.30ab 0.00 ± 0.00b 0.00 ± 0.00b

Trinervitermes 0.17 ± 0.24ab 0.00 ± 0.00b 0.00 ± 0.00b

Amitermes 0.50 ± 0.32ab 0.33 ± 0.30b 0.17 ± 0.24b

LSD 1.22 0.76 0.88

Means within a column followed by the same letter do not differ significantly by Least Significant Difference Test (LSD) at 5% of probability. Table 6 Termites collected by random, standardized belt transect, and maize stalk baiting methods in three land-use types in the Central Rift Valley of Ethiopia in the year 2013

Land-use type Sampling methods Total species collected from each land-use type

Random Standardized belt transect

Maize stalk baiting

Protected land Macrotermes Amitermes Odontotermes

Macrotermes Microcerotermes Amitermes

Macrotermes Microcerotermes Amitermes Angulitermes Microtermes Trinervitermes

7 (100%)

Rangeland Macrotermes Odontotermes Microcerotermes

Macrotermes

Macrotermes Microtermes Amitermes Microcerotermes

4 (57.1%)

Farmland Macrotermes Odontotermes Microtermes

Macrotermes

Macrotermes Microtermes Amitermes Microcerotermes Odontotermes

5 (71.4%)

Total species collected using each method

5 (71.4%) 3 (42.9%) 7 (100%) -

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A

B

C

Plates 4A-C Macroterems mounds of different forms with openings. Ventilation openings are off center of the nest (A). Photo: Daniel Getahun (October 2013).

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Plate 5 Cone-shaped Macortermes mound without opening. Photo: Daniel Getahun (October 2013).

Efficacy of sampling methods: The relative efficacy of the different methods in sampling termites and the number of genera captured from different land-use types are shown in Table 6. The maize stalk baiting method captured all (100%) the recorded genera in the study area, while the standardized belt method and random (casual) sampling method captured three (42.9%) and five (71.4%) of the total record, respectively. With respect to land-use type, all the seven genera collected from the area were also captured from protected lands, while only four (57.1%) and five (71.4%) genera were captured from rangeland and farmland, respectively.

DISCUSSION

Species composition: The Results show that termite species belonging to seven genera were recorded in the area and of these make 28% of the total 25 termite genera reported by Abdurahman (1990) earlier in the country. Of these recorded genera, Trinervitermes and Angulitermes were very rare species. Trinervitermes was recorded for the first time in the Central Rift Valley of Ethiopia in this study. Trinervitermes and Angulitermes are also uncommon in the country and Wood (1986) recorded Trinervitermes only in Southern Peoples Nations and Nationalities Regional State, 90 km from Sodo town towards Gofa and Gambela Regional State; Angulitermes only from Yaballo town. Except

Tinervitermes, all the genera in the current study were recorded by Abraham Tadesse (1987) cited in Abdurahman et al. (2010). The termite genera recorded in the area was 28% of the total 25 termite genera reported by Abdurahman (1990) earlier in the country.

The results have shown that all the termite species belonging to the seven genera recorded in the area were also collected from protected lands, while only species belonging to four and five genera were collected from rangelands and farmlands, respectively. The three land-use types were not equally managed by man, the protected lands being the least managed (the least disturbed) followed by rangelands and farmlands (the most disturbed). The difference implies that cultivation and grazing by livestock appear to eliminate some termite species. Deforestation followed by cultivation and overstocking increase climatic instability and decrease habitat heterogeneity. This situation eliminates the most susceptible termite species to disturbance and causes the outbreak of those species which tolerate disturbances, contributing to a biological situation characterized by a low biodiversity with few dominant species (Kooyman and Onck, 1987; Sileshi et al., 2009).

The absence of Angulitermes and Trinervitermes in managed lands in the current study may indicate that

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these species are sensitive to environmental disturbances. Trinervitermes species are specialist grass feeders and their presence in the protected land is due to availability of grass and loss of adequate grass vegetation would deprive the species of adequate feed resources for survival (Saran et al., 2008; Mugerwa et al., 2011; Dosso et al., 2012). Dosso et al. (2012) collected grass-feeding Trinervitermes spp. from protected areas restricted to savannas which provided an elevated quantity of grassy litter. Mugerwa et al. (2011) also noted the rare occurrence of Trinervitermes spp., by sampling only from plots with dense grass cover and attributed rarity of the genus to the escalating denudation of grass cover due to overgrazing and competition for resources with members of other genera. The degree of anthropogenic activities appears to contribute to the termite diversity and assemblages in the area. Therefore, habitat disturbance seems a key factor for the differences in species composition and abundance of termites among the different land-use types.

The mound-building termite genera include Macrotermes, Cubitermes, Amitermes, Odontotermes and Trinervitermes (Meyer et al., 1999). Odontotermes species build both subterranean and epigeal nests (UNEP, 2000). However, in the current study, Trinervitermes and Odontotermes were not sampled from mounds. Amitermes were collected only from peripheral wall of Macrotermes mounds from sites where grasses grew and by maize stalk baiting. If the Amitermes were mound builders, from Amitermes and the Macrotermes, the original builder of the nest was unknown. In line with this, Sileshi et al. (2010) reported that the original builder of a mound could be one species, each mound may be inhabited and modified by several species through time, and that it is often difficult to determine which one first established the mound. In Zambia species of over 20 different genera were found in the mounds of Macrotermes and in Nigeria, 37% of the mounds of Cubitermes fungifaber were occupied by a total of 31 termite species.

The occurrence of two types of Macrotermes mounds having cone and dome shapes, and mounds with and without external openings is an indicative of the existence of different species of the genus in the area. Wood (1986) has also reported the existence of the two types of Macrotermes mound in the area but did not mention the species building the mounds.

Contrary to the current study, Abdurahman (1990) reported that Macrotermes mounds found in western

Ethiopia and Maki-Batu area differ significantly both in size and shape and that no external openings are found on both mounds. In western Ethiopia the mounds are typically dome-shaped whereas in Maki-Batu, the mounds are conical shaped. He further reported that the genus Macrotermes is represented in Ethiopia by M. subhayalinus and M. herus (Abdurahman et al., 2010) and the two species are very similar to each other and identification is notoriously difficult (Abdurahman, 1990). According to this report, the well described and severe pest species in Ethiopia, M. subhayalinus, builds closed and dome-shaped mounds and this is does not agree with different literatures. For instance, Darlington (1988) noted that M. herus and M. michaelseni build closed mounds and Haimanot (2002) reported that M. subhyalinus is morphologically very similar to M. michaelseni and their taxonomic status has not yet been fully resolved, with open mounds being attributed to M. subhyalinus and closed mounds to M. michaelseni and thus taxonomic identification of Macrotermes species in Ethiopia is very necessary (Haimanot, 2002).

Termite abundance in the land-use types: The results show that abundance (encounter rates) of the termite species differ between land-use types. The least occurrence of Macrotermes in the farmland as compared with the other areas could be attributed to agricultural activities which limit establishment of new colonies. Similarly, Mugerwa et al. (2011) noted that M. bellicosus and M. subhyalinus dominated the termite assemblage structure in the rangelands of Tororo District in Uganda. Cultivation results in the destruction of mounds of Macrotermes, Odontotermes, and Trinervitermes (Okwakol, 2001). Clearing trees and subsequent cultivation for agriculture usually destroys the nests of mound building species and those with shallow, subterranean nests, and those that depend upon trees or woody litter (Wood, 1986). In Kenya Kooyman and Onck (1987) also noted that the Microtermes species tend to increase in abundance with increasing intensity of cultivation. This increase unlike other termite species could be attributed to their deep subterranean nests and thus they are less affected by cultivation unlike termites like Macrotermes which have shallow nests (Abdurahman, 1990). Species with deep subterranean nests and with the ability to survive on living crops and crop residues remain and increase in number; the most important of these are Microtermes (Wood, 1986)

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The higher termite diversity recorded in protected lands than rangelands and farmlands implies the use of termites as bi-indicators of habitat change in the area assuming that all the sites had originally similar vegetation cover and had similar termite assemblages. Such protected areas can also be used as refuges for those termites which are sensitive to environmental disturbances. In line with this, (Jones and Eggleton, 2000) reported that termites are potentially one of the most important indicator taxa because they are at the ecological center of many tropical ecosystems. Ecological indicators are taxa that are sensitive to, and demonstrate the effects of, anthropogenic environmental stress or disturbance on biotic systems (Jones and Eggleton, 2000). In Nigeria, removal of trees and bushes followed by cultivation resulted in loss of Macrotermes, Cubitermes, Trinervitermes and shallow nests of Coptotermes. Some subterranean species such as Microtermes increase in numbers, as they could move down into the soil to avoid the effects of surface disturbance (Pearce, 1997).

Efficacy of sampling methods: The maize stalk baiting method was able to capture the entire termite species recorded in the area and thus produced the highest termite species richness than the standardized belt transect method and random collection. The termite genera encounters were also higher in the maize stalk baiting technique than the belt standardized search method. The differences indicate that the species were wood-based cellulose feeders and they were highly attracted to the maize stalk. In their assessment to test the standardized belt transect protocol, Jones and Eggleton (2000) noted that each transect they ran in three forests captured approximately 31-36% of the known local termite fauna. Thus, when termite inventories are adopted, baiting, especially maize stalk baiting, can be considered as the most suitable for assessing wood feeding termite species assemblages in savannah areas like the CRVE.

CONCLUSION AND RECOMMENDATION: Termite species belonging to one family (Termitidae), three subfamilies and seven genera (Macrotermes, Microtermes, Odontotermes, Microcerotermes, Angulitermes, Trinervitermes and Amitermes) were recorded in the area. In contrary to rangelands and farmlands, all the genera were also collected from protected lands. Likewise, Angulitermes and Trinervitermes were very rare and they were sampled only from protected lands and this shows that the

species are sensitive to anthropogenic activities and termite diversity is affected by degree of land-use. Use of maize stalk baiting was the best method in sampling maximum number of species and the entire recorded termite species in the area were captured using the method. Although, the sampling was exhaustive and covered a wide area, the termite species recorded might not represent all the available termite diversity in the area as it was carried out once only after the main rainy season. Thus, to provide a full picture of the termite diversity and species composition, further studies have to be conducted widely in different seasons throughout the year and also target at soil feeders. Taxonomic identification of Macrotermes termites to species level has to be made using molecular techniques. To date, no systematic termite inventory apparently has been carried out at national level in Ethiopia. Therefore, the result of this study can form a baseline for designing a national termite species inventory.

ACKNOWLEDGEMENTS

The authors are grateful to Adama Science and Technology University for funding the research. Special thanks go to the field assistants and volunteer farmers to do the research on their farmlands and rangelands.

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