Composition and distribution of indigenous trees and shrubs as possible criteria for indicating adapted species in semi-arid rangelands Edward K. Mengich1*, Daniel K. Too2, Joseph M. Macharia3 and Ralph Mitloehner4 1 Rift Valley Eco-regional Research Programme, Kenya Forestry Research Institute (KEFRI), PO Box 382-20203, Londiani, Kenya, 2Department of Natural Resources, Egerton University, PO Box 536-20115, Njoro, Kenya, 3Department of Botany, Egerton University, PO Box 536-20115, Njoro, Kenya and 4Institute of Silviculture, Georg-August University, Busgenweg 1, D-37077, Goettingen, Germany Abstract This study assessed the composition and natural distribution of indigenous trees and shrubs as possible criteria for selecting suitable species for rehabilitation of degraded sites in semi-arid rangelands. Study sites were identified at Nthangu, Kathonzweni and Kibwezi forests of Makueni County, Kenya using existing vegetation, agro-climatic maps and Landsat imageries. The sites had mean annual rainfalls of 974 mm, 700 mm and 616 mm, respectively, and moisture indices of 49%, 35% and 32%. Data were collected by establishing sample plots and assessing species counts and diameters at breast height (DBH). Basal area, relative dominance, relative abundance, relative frequency and important value indices (IVIs) were computed for individual families and species at each site. The number of families, genera and species declined from Nthangu (33, 60, 77) through Kibwezi (30, 48, 70) to Kathonzweni (28, 42, 69). Corresponding mean basal areas were 16.7 m2 ha1, 76.8 m2 ha1 and 19.3 m2 ha1. The families Combretaceae, Burseraceae and Mimosaceae were the most important and widely distributed. Based on ecological importance values, candidate species for rehabilitation of degraded sites at Nthangu, Kathonzweni and Kibwezi were Combretum molle and Acacia hockii; Combretum collinum, Commiphora campestris and Acacia tortilis; and Commiphora africana and A. tortilis, respectively. possibles pour la selection d’especes appropriees pour rehabiliter des sites degrades dans des espaces semi arides. Des sites d’etudes furent identifies dans les for^ets de Nthangu, Kathonzweni et Kibwezi, dans le Makueni County, au Kenya, en se basant sur la vegetation existante, sur des cartes agro-climatiques et l’imagerie Landsat. Ces sites ont une pluviosite annuelle moyenne de 974 mm, 700 mm et 616 mm respectivement et des indices d’humidite de 49%, 35% et 32%. Les donnees furent collectees en etablissant des parcelles echantillons et en relevant le nombre d’especes et leur DBH. La surface basale, la dominance relative, l’abondance relative, la frequence relative et l’indice d’importance des especes (IVI) furent calcules pour chaque famille et pour chaque espece sur chaque site. Le nombre de familles, de genres et d’especes Kibwezi allait en decroissant de Nthangu (33, 60, 77) a (30, 48, 70) et enfin Kathonzweni (28, 42, 69). Les surfaces basales moyennes correspondantes etaient de 16.7 m2 ha1, 76.8 m2 ha1 et 19.3 m2 ha1. Les familles des Combretaceae, des Burseraceae et des Mimosaceae etaient les plus importantes et etaient largement distribuees. En se basant sur les indices d’importance ecologiques, les especes candidates pour la rehabilitation Nthangu, Kathonzweni et Kibwezi de sites degrades a etaient: Combretum molle et Acacia hockii; Combretum collinum, Commiphora campestris et Acacia tortilis; et Commiphora africana et A. tortilis, respectivement. Key words: important value index, indigenous, Kenya, rangelands, semi-arid, trees Resume Introduction Cette etude a evalue la composition et la distribution naturelle d’arbres et d’arbustes indigenes comme criteres Woody vegetation cover in the semi-arid rangelands of Kenya has declined over time because of over-exploitation due to pressure from increasing human populations and accompanying increases in demand for various tree *Correspondence: E-mail: emengich3@hotmail.com © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 3 4 Edward K. Mengich et al. products and services. In addition, the number of trees and shrubs that can tolerate drought stress has diminished because of other factors such as frequent outbreaks of bushfires, and climate change, which have resulted in harsher and drier conditions in rangelands (Ensor & Berger, 2009; Omambia, Shemsanga & Li, 2009). These factors have resulted in moderate to severe land degradation and desertification that are manifested in forms of impoverishment, depletion of vegetation cover, and deterioration of physical, chemical and biological soil properties. There have been concerted efforts to reverse the impacts of degradation through tree planting in the semi-arid rangelands of Kenya (Kigomo, 2003). However, rehabilitation of these sites has severely been constrained by a narrow species selection base, lack of information to guide selection of suitable species and use of exotic species that are not adapted to local conditions (Reubens et al., 2011). Planting of appropriate tree species in respective localities is key to successful rehabilitation of degraded sites (Derbel & Chaieb, 2013). Provenance trials have been used to identify suitable species to plant in the semi-arid rangelands of Kenya. However, the trials have largely tested suitability of exotic species (Milimo, Dick & Munro, 1994). Due to the long period that provenance trials take to generate information and therefore, guide on appropriate species to plant, use of faster-growing exotic species that are not adapted to local conditions is common. The use of indigenous species for rehabilitation has not been widely practised in Kenya, but the potential for their use is widely acknowledged. Indigenous species have a wide range of advantages over exotic species as they are adapted to local conditions. When properly matched with planting sites, they are easier to establish, more likely to survive and environmentally friendly. There are also minimal risks of indigenous tree species becoming invasive (Greening, Landscape & Tree Management Section Development Bureau, 2010). Nevertheless, rehabilitation using indigenous species in Kenya faces challenges as the country lacks guidelines on species–site matching. Published guidelines (KEFRI, 1990; Braun, Albrecht & Kamondo, 1993) are based on results of years of range-wide reconnaissance surveys that recorded occurrence of species in given localities. Braun, Albrecht & Kamondo (1993) superimposed the results of the reconnaissance surveys on the forest seed zones of Kenya as a further improvement to reduce risk of moving tree seed far beyond its right ecological conditions. The publications provide general guidelines at a national scope due to the nature of data used in their development. However, rehabilitation programmes for specific sites are still faced with the challenge of correct species choices for prevailing physiognomic characteristics. In this study, we assessed the composition and natural distribution of indigenous trees and shrubs and derived basal areas (BAs) and important value indices (IVIs) as possible dynamic criteria for selecting suitable tree and shrub species for the rehabilitation of degraded sites for specific locations in semi-arid rangelands. Materials and methods Study area The study was conducted in Makueni County, Kenya (Fig. 1). The county lies between 1°35′S and 2°35′S, and 37°10′E and 38°30′E and covers an area of 7,965.8 km2 (GoK, 1997). A large proportion of the county falls within the semi-humid to arid regions (agro-climatic zones IV–VI) with moisture indices of less than 50% and a mean annual rainfall of much less than 1100 mm (Sombroek, Braun & Van Der Pour, 1982). Agro-climatic zones IV–VII are referred to as rangelands. Agro-climatic zones are delineated according to mean annual rainfall and temperatures (Jaetzold and Schmidt, 1983). Makueni County is generally low-lying with altitude ranging between 600 m above sea level (a.s.l) at Tsavo and 1900 m a.s.l on Kilungu hills. It is characterized by arid and semi-arid climates (Pratt, Greenway & Gwynne, 1966; Maundu & Tengnas, 2005; Van den Abeele, Ngatia & Macharia, 2005) with low and unreliable annual rainfall ranging from 200 mm to 900 mm in the low-lying areas and from 800 mm to 1200 mm in the hills (GoK, 1997). The rainfall is bimodally distributed and occurs in March/April and November/ December. The November/December rains are usually more reliable both in amount and distribution and accounts for 58% of the annual total precipitation (Musembi, 1986). Study sites Three study sites were randomly selected using agroclimatic maps and Landsat imageries and on the basis of existing vegetation (Lambrecht, 1989; Shiver & Borders, 1996; Fasona & Omojola, 2009; Fig. 1). The maps and satellite pictures were obtained from the Survey of Kenya and the Department of Resource Surveys and Remote Sensing (DRSRS; GoK, 2006). Selection of sites was complemented by a tour of the county and a survey of the vegetation and local weather facilities. © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Indigenous trees and shrubs as adaptation criteria 5 Fig 1 Location and agro-ecological zones of Makueni County, Kenya Selected sites were minimally disturbed natural forests, woodlands and bushlands located in Nthangu (1575 m a.s.l.; site I), Kathonzweni (1250 m a.s.l.; site II) and Kibwezi (900 m a.s.l.; site III; Fig. 1). Mean annual rainfall and moisture indices for Nthangu, Kathonzweni and Kibwezi were 974 mm and 49%, 700 mm and 35%, and 616 mm and 32%, respectively. As there were no climatic data recording stations within the selected sites, the climatic data were those of nearby weather stations within the same zone, that is, Katende forest station, KARI Kampi ya Mawe and Makindu meteorological station, for the three sites, respectively. Data collection Indigenous and shrubs were identified, and data were collected by establishing sample plots at each study site. © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Establishment of sample plots. Sample plots were established according to Lambrecht (1989) and Shiver & Borders (1996). At Nthangu, the sample plots were established along a 15-km earth road running from the forester’s office in the north to Mitubu hill overlooking Wote township in the south. The plots were established at Mbenza, Kyamboo, Kangaea and Mitubu points of the forest. At Kathonzweni, sample plots were established along a 30-km earth road connecting Kambi ya Mawe market along Kathonzweni–Wote Road and Miangeni market near Athi River. The plots were established at Kampi ya mawe, Mathemba A, Mathemba B and Miangeni. At Kibwezi, plots were established along the earth road connecting Kisayani market on the Kibwezi–Athi River road, and Kyanginywa market on the Kibwezi River. The plots were established within the forest owned by the University of Nairobi’s Institute of Drylands Research. 6 Edward K. Mengich et al. Along each road, four benchmarks were established at 4–5-km intervals. One to 3 km away from each benchmark in alternate directions (to the right or left), a 50-m by 50-m (2500 m2) plot was established (Lambrecht, 1989). In sloping areas, sample plots were aligned along the slope. In flat areas, the alignment was random. Identification and inventory of trees and shrubs. With assistance from a forester and a parataxonomist, all trees and shrubs in each plot were identified by their local names, botanical names and families, using available literature (Dale & Greenway, 1961; Beentje, 1994; Maundu & Tengnas, 2005). Trees and shrubs were counted and recorded by species. For ease of assessment and to avoid repetition, each sample plot was partitioned into five equal units, and resultant data were aggregated for analysis. A tree or shrub would be counted as occurring whether it was a seedling, sapling or mature individual. Assessment of basal areas (BAs). Basal areas were only assessed for trees above 1.30 m in height. Diameters at breast height (DBH; 1.30 m above ground) were measured using diameter tapes. Where trees were below this height, DBH was not taken. Basal areas were derived as follows: BA ¼ p ðDBH=2Þ2 where BA = basal area; p = 3.14; DBH = diameter at beast height. The BAs were reported on a per-hectare basis. Statistical analysis Data were compiled and entered into MS-Excel, versions 2003 and 2007, where data management techniques such as data checking for errors, exploration and general pattern of the data were performed. Basal area (BA), abundance (A), frequency (F %), relative dominance (RD %), relative frequency (RF %), relative abundance (RA) and IVIs of individual species and families were computed for each study site. In the case of BA, data were analysed using GenStat v10 (VSI International, Hemel Hempstead, UK), a general statistical software. Analysis of variance (ANOVA) was performed to compare the basal areas of members of the same species growing in at least two study sites. Data were log-transformed to satisfy the normality assumption of ANOVA. Whenever the normal distribution of the data was not observed, nonparametric tests (Mann–Whitney U and Kruskal–Wallis) were used. Mann–Whitney U-test was used for comparing the BA measurements of the species that occurred in only two study sites. For these nonparametric tests, mean ranks of the response variate for comparison of independent variables were used, and statistical differences were declared using Z-score and chi-square tests for Mann–Whitney U and Kruskal–Wallis, respectively. All statistically significant differences were declared at P < 0.05. Results Species characterizing various sites Determination of important value indices (IVI). As BA is a factor in IVI determination, IVIs were only determined for tree species whose BA could be calculated, that is, trees with a minimum height of 1.30 m above ground. Count and BA measurements were used to calculate IVIs for both tree species and families. The IVIs were derived following Misra (1968): IVI ¼ relative abundance (RA) þ relative dominance (RD) þ relative frequency (RF) where: abundance (A) = number of individuals per hectare (N ha1); RA = abundance of a given species/total abundance of all species 9 100; dominance (D) = basal area (m2 ha1); RD = basal area of a given species/total basal area of all species 9 100; frequency (F) = percentage (%) of plots in which the species is represented; RF = frequency of a given species/total frequency of all species 9 100. Vegetation at Nthangu forest consisted of 77 different tree and shrub species that may be categorized into 33 families and 60 genera (Table 1). The most common genera in order of frequency were Combretum (5), Acacia (5), Grewia (2), Cassia (2), Searsia (2) and Terminalia (2). The most common families were Combretaceae (6), Mimosaceae (6), Papilionaceae (5), Euphorbiaceae (5), Anacadiaceae (4) and Caesalpiniaceae (3). Vegetation at Kathonzweni consists of 69 different tree and shrub species that may be categorized into 28 families and 42 genera (Table 2). The most common genera in order of frequency were Commiphora (8), Acacia (6), Combretum (4), Boscia (3), Grewia (2) and Terminalia (2). The most common families were Mimosaceae (9), Burseraceae (7), Combretaceae (6), Capparaceae (4) and Papilionaceae (3). © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Indigenous trees and shrubs as adaptation criteria 7 Table 1 The tree and shrub species of Nthangu forest, Makueni County No. Species local name Botanical name Family name Type IVI 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. Kaluma Mukinyai Kiama Mutheu Mukongoo Munyua Mukaati Mukukuma Muuku Mutotoo Mwaalika Mukomoa Mutoo muka Musemei Muuwa nzuki Kiba Munoa mathoka Mukengeka Mpingo Mwaanzia Utithi Mung’uthe Mulawa muka Itithyo Muketa Kitolousuu Mwaa Mulawa isamba Kivuti Muvau Mutula Mukokola Kithongoi Muvuavoi Muthia Kithauna Muthulu Mulului Musensili Mutuva Mwinthongoi Musuusuu Mutote Mutote Mutongu Muvuu Kithunzi Kiaa Lunguyu Pithosporum viridiflorum Euclea divinorum Combretum molle Searsia spp. Diospyrus mespiliformis Acacia hockii Faurea saligna Uvaria scheffleri Terminalia brownii Pachystigma schumannianum Heeria reticulata Vangueria infausta Azanza garckeana Acacia nilotica Combretum apiculatum Pappea capensis Dicrostachys cinerea Cassia sengueana Dalbergia melanoxylon Bridelia taitensis Combretum collinum Lonchocarpus eriocalyx Grewia bicolour Combretum zeyheri Myrsine africana Ziziphus abyssinica Acacia tortilis Grewia spp. Erythrina abyssinica Dombeya kirkii Ximenia americana Combretum exalatum Dodonea angustifolia Steganotaenia araliaceae Acacia mellifera Lannea schimperi Croton megalocarpus Balanites aegyptiaca Gnidia latifolia Grewia tembensis Pavetha gardeniifolia Crotalaria spp. Carissa edulis Carissa edulis Solanum incanum Grewia villosa Maytenus heterophylla Euphorbia candelabrum Indigofera spp. Pithosporaceae Ebenaceae Combretaceae Anacardiaceae Ebenaceae Mimosaceae Proteaceae Annonaceae Combretaceae Rubiaceae Anacardiaceae Rubiaceae Malvaceae Mimosaceae Combretaceae Sapindacea Mimosaceae Caesalpiniaceae Papilionoideae Euphorbiaceae Combretaceae Papilionaceae Tiliaceae Combretaceae Myrsinaceae Rhamnaceae Mimosoidiae Tiliaceae Papilionaceae Sterculiaceae Olacaceae Combretaceae Sapindaceae Umbelliferae Mimosaceae Anacardiaceae Euphorbiacea Simaraubaceae Thymelaeaceae Tiliaceae Rubiaceae Papilionaceae Apocynaceae Apocynaceae Solanaceae Tiliaceae Celastraceae Euphobiacea Papilionaceae T, S T T S,T T S, T S, T T, T S,T T S, T T T T S, T T, S T S,T S, T S,T S, T S, T S, T S, T T, S T T T S,T T,S T S, T T S, T T T T S S S S S S S S S, T T S 21.6 21.3 20.2 19.8 18.6 16.9 11.7 11.4 10.4 8.9 8.7 7.6 6.6 6.3 6.1 5.8 4.7 4.6 4.4 4.0 3.9 3.9 3.5 3.3 3.1 3.1 2.8 2.0 1.9 1.9 1.8 1.8 1.6 1.6 1.1 1.1 0.9 0.8 – – – – – – – – – – – © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 8 Edward K. Mengich et al. Table 1 (continued) No. Species local name Botanical name Family name Type IVI 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. Mwindenguwe Mukakaa Mukala Mukandu Mukayau Mukenea Mukiliuli Mukuluu Mukulwa Mukubu Mungendia nthenge Mongolli Munyunga-mai Musomolo Musovi Mutandi Muthika Muthingii Muthumula Muti Mutoo Mutumbuu Mutungate Mutungu Mutunguu Muua Muvatha Mwalula Triumfetta flavescens Premna resinosa Antidesma venosum Ocimum suave Salvadora spp. Zanthoxylum chalybeum Harrisonia abyssinica Securinega virosa Acalypha fruticosa Craibia brownii Kleinia squarrosa Acacia senegal Cassia didymobotrya Lantana camara Hoslundia opposita Ochna inermis Indigofera spp Ormocarpus kirkii Tamarindus indica Aspilia mossambisensis Terminalia prunoides Scutia myrtina Commiphora habessinica Commiphora spp. Thylachium thomasii Sclerocarya birrea Vernonia lesiopus Croton dichogamus Tiliaceae Verbenacea Euphorbiacea Labiatae Salvadoraceae Rutaceae Simaraubaceae Euphorbiaceae Euphorbiacea Papilionoidiae Compositae Mimosaceae Caesalpiniaceae Verbenaceae Labiatae Ochnaceae Papilionaceae Papilionaceae Caesalpiniaceae Compositae Combretaceae Rhamnaceae Burseraceae Burseraceae Capparaceae Anacardiaceae Compositae Euphorbiacea S S T S T, S S, T S, T S S T S S, T S, T S S T S T T S T S, T S, T T S, T T H,S T – – – – – – – – – – – – – – – – – – – – – – – – – – – – T, tree; S, shrub; B, bush; C, climber; H, herb; L, liana; IVI, important value index. NB: Species indicated as ‘tree’ (T) but not having a value for IVI was <1.30 m in height. Vegetation at Kibwezi consisted of 70 different tree and shrub species that may be categorized into 30 families and 48 genera (Table 3). The most common genera in order of frequency were Acacia (8), Commiphora (7), Grewia (3), Boscia (3), Combretum (2) and Lannea (2). The most common families were Mimosaceae (11), Burseraceae (7), Capparaceae (4), Tiliaceae (3), Euphorbiaceae (3), Combretaceae (3) and Anacardiaceae (3). Basal areas of tree species Overall mean BAs of trees at Nthangu, Kathonzweni and Kibwezi were 16.7 m2 ha1, 19.3 m2 ha1 and 76.8 m2 ha1, respectively. The nine top tree species at each site accounted for 70.7%, 86.5% and 78.3% of the overall mean BA, respectively. Among three species, each occurring in at least two of the study sites, mean BAs differed between sites (Table 4). For instance, the BA of Terminalia brownii was higher at Nthangu (0.17 m2 ha1) than at Kathonzweni (0.07 m2 ha1), although the differences were not significant (t1,19 = 1.26, P > 0.05), while those of Acacia tortilis (t1,252 = 3.83, P < 0.05) and Acacia nilotica (t1,71 = 1.96, P < 0.05) were both significantly higher at Kathonzweni (0.09 m2 ha1, 0.04 m2 ha1, respectively) than at Kibwezi (0.04 m2 ha1, 0.02 m2 ha1, respectively). At Kathonzweni, A. tortilis had the highest BA (0.09 m2 ha1), followed by T. brownii (0.07 m2 ha1) and A. nilotica (0.04 m2 ha1) in that order. The former and the latter were significantly different (t1,252 = 3.83, P < 0.05). At Kibwezi, A. tortilis had a BA (0.04 m2 ha1) © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Indigenous trees and shrubs as adaptation criteria 9 Table 2 The tree and shrub species of Kathonzweni forest, Makueni County No. Local name Botanical name Family name Type IVI 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. Utithi Muuwa nzuki Mwaa Musemei Muthaalwa Muuku Iulu Mulawa isamba Ikuu Itula Kiusia Muthia Mukuswi Kiluli Mwaanzia Kyoa kikaa Mutandi Ithityo Mwai Kyundua Munyua Muvuavoi Mutoo Mwalanthate Iliva Kyoa isamba Musovi Mukakaa Musensili Lunguyu Muthika Muvuluvulu Isivu Itumbukyamuu Mupopotwe Kiongwa Kithea Kithunzi Kiva Kyenzenze Kiongwa Kyuasi Mutungu Mutunguu Mwindenguwe Mukayau Mukigeka Mukiliuli Mukokola Mukuluu Combretum collinum Combretum apiculatum Acacia tortilis Acacia nilotica Lannea triphylla Terminalia brownii Commiphora campestris Grewia spp. Commiphora africana Commiphora baluensis Sterculia africana Acacia mellifera Acacia brevispica Boscia angustifolia Bridelia taitensis Commiphora ovalifolia Ochna inermis Combretum zeyheri Platycelyphium voense Albizia amara Acacia hockii Steganotaenia araliacea Terminalia prunoides Cassia abbreviata Commiphora rostrata Albizia anthelmintica Hoslundia opposita Premna resinosa Gnidia latifolia Indigofera spp. Indigofera spp. Opilia celtidifolia Boscia coriacea Faurea saligna Maerua kirkii Combretum paniculatum Cordia monoica Maytenus heterophylla Pappea capensis Boscia spp. Combretum paniculatum Lannea stuhlmanii Solanum incanum Thylachium thomasii Triumfetta macrophyla Salvadora persica Cassia sengueana Harrisonia abysinica Combretum exalatum Securinega virosa Combretaceae Combretaceae Mimosoideae Mimosaceae Anacardiaceae Combretaceae Burseraceae Tiliaceae Burseraceae Burseraceae Sterculiaceae Mimosoideae Mimosaceae Capparaceae Euphorbiaceae Burseraceae Ochnaceae Combretaceae Papilionaceae Mimosoideae Mimosaceae Umbelliferae Combretaceae Caesalpinioideae Burseraceae Mimosaceae Labiatae Verbenaceae Thymealaeaceae Papilionaceae Papilionaceae Opiliaceae Capparaceae Proteaceae Capparidaceae Combretaceae Boraginaceae Celastraceae Sapindaceae Capparaceae Combretaceae Anacardiaceae Solanaceae Capparaceae Tiliaceae Salvadoraceae Caesalpiniaceae Simaraubaceae Combretaceae Euphorbiaceae S,T T T T S,T T T T T T T S,T S,T S,T S,T S,T T T T T S,T T S,T T S,T T, B S S S S S, L S, T S, T S, T S, L S, T S, T T S, T S, L S, T S S, T S T, S T S, T T S 44.4 29.5 27.6 16.8 15.9 12.6 12.4 11.4 11.0 9.4 7.6 7.4 6.2 6.2 5.7 4.6 4.4 3.3 3.2 2.7 2.5 2.4 2.4 2.2 1.8 1.1 – – – – – – – – – – – – – – – – – – – – – – – – © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 10 Edward K. Mengich et al. Table 2 (continued) No. Local name Botanical name Family name Type 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. Mukulwa Mukume Mulawa muka Mongolli Munoa mathoka Musomolo Mutaavesi Muthuigi Muthito Mutongu Mutoo muka Mutotoo Mutungate Mututi Mutuba Mpingo Muvuu Mwaitha Yongwa Acalypha fruticosa Haplocoelum foliolosum Grewia bicolour Acacia senegal Dicrostachys cinerea Lantana camara Lantana camara Ormocarpus kirkii Cadaba farinosa Solanum incanum Azanza garckeana P. schumannianum Commiphora habessinica Thunbergia holstii Grewia tembensis Dalbergia melanoxylon Grewia villosa Entada leptostachya Commiphora riparia Euphorbiaceae Sapindaceae Tiliaceae Mimosaceae Mimosaceae Verbenaceae Verbenaceae Papilionaceae Capparidaceae Solanaceae Malvaceae Rubiaceae Burseraceae Acanthaceae Tiliaceae Papilionaceae Tiliaceae Mimosaceae Burseraceae S S, S, S, S, S S T S, S T S T S S S, S C T T T T T T T IVI – – – – – – – – – – – – – – – – – – – T, tree; S, shrub; B, bush; C, climber; L, liana; IVI, important value index. NB: Species indicated as tree (T) but not having a value for IVI was <1.30 m in height. twice higher than that of A. nilotica (0.02 m2 ha1). However, the differences were not statistically significant (t1,71 = 1.96, P > 0.05). Ecologically important families and trees The most important families according to descending order of their IVIs at Nthangu were Combretaceae (45.7), Ebenaceae (39.9), Mimosaceae (31.8), Anacardiaceae (29.6) and Pittosporaceae (21.6). At Kathonzweni, they were Combretaceae (92.2), Mimosaceae (64.3), Burseraceae (39.2), Anacardiaceae (15.9) and Tiliaceae (11.4). At Kibwezi, the most important families were Mimosaceae (77.3), Burseraceae (40.6), Bombacaceae (38.3), Caesalpiniaceae (30.4) and Capparaceae (17.5). The other families had overall IVIs of 41.6, 31.7 and 46.8 for the three sites, respectively. At Nthangu, ecologically important tree species according to descending order of their IVIs could be determined for 38 species (Table 1). The most important species were Pithosporum viridiflorum (21.6), Euclea divinorum (21.3), Combretum molle (20.2), Searsia spp. (19.8) and D. mispiliformis (18.6). At Kathonzweni, ecological importance of trees by their IVIs could be determined for 26 of the species in descending order (Table 2). The most important species were Combretum collinum (44.4), Combretum apiculatum (29.5), A. tortilis (27.6), A. nilotica (16.8) and Lannea triphylla (15.9). Ecologically important tree species at Kibwezi are listed in Table 3 according to descending order of IVIs. Only 42 tree species could have their IVIs determined. The most important species were Adansonia digitata (38.3), D. elata (25.7), A. tortilis (20.8), Commiphora africana (17.4) and Acacia mellifera (14.3). Commonly occurring tree species Predominant species that occurred commonly in all the three sites and exhibited the highest IVIs were A. nilotica (6.3) and A. tortilis at Kathonzweni (27.6) and Kibwezi (20.8; Table 5). Discussion The number of species identified in this study (139) represents less than a tenth of the trees and shrubs that may be encountered somewhere in the ASALs of Kenya (Ogolla & Mugabe, 1997; Maundu & Tengnas, 2005). In a similar study covering an area comprising four counties of © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Indigenous trees and shrubs as adaptation criteria 11 Table 3 The tree and shrub species of Kibwezi forest, Makueni County No. Species local name Botanical name Family name Species type IVI 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. Mwambo Mwangi Mwaa Ikuu Muthia Munina Kyoa isamba Mulawa muka Itula Mutunguu Musemei Mulawa isamba Mwalula Kiluli Kisaya Kyenzenze Kyoa kikaa Mwalanthate Mutoo Mupopotwe Kiaa Mongolli Munoa mathoka Iliva Iulu Muvau Mulela Muthaalwa Muvuavoi Mukangakanywa Kyaakyusi Kiusia Mutungate Yongoa Kikaiki Muthuigi Kyuasi Mukaiao Yumbu Ikindu Mutiligo Mukami Mutongu Mugea Kiongwa Muvuu Muthika Musomolo Isivu Adansonia digitata Delonix elata Acacia tortilis Commiphora africana Acacia mellifera Acacia elatior Albizia anthelmintica Grewia bicolour Commiphora baluensis Thylachium africanum Acacia nilotica Grewia spp. Croton dichocamus Boscia angustifolia Bechermia discolour Boscia spp. Commiphora ovalifolia Cassia abbreviata Terminalia prunoides Maerua kirkii Euphorbia spp. Acacia senegal Dicrostachys cinerea Commiphora rostrata C. campestris Dombeya kirkii Acacia xanthophloea Lannea triphylla Steganotaenia eraliacea Garcinia livingstonii Combretum schumannii Sterculia africana Commiphora habessinica Commiphora hildbraedii Acacia thomasii Ormocarpus kirkii Lannea schumanii Salvadora persica Ficus spp. Phoenix reclinata Lawsonia inermis Neutonia hildbrandii Solanum incanum Anisotes ukambensis C. paniculatum Grewia villosa Indigofera spp. Lantana camara Boscia coriacea Bombacaceae Caesalpinioideae Mimosoidiae Burseraceae Mimosoideae Mimosoideae Mimosaceae Tiliaceae Burseraceae Capparaceae Mimosoideae Tiliaceae Euphorbiaceae Capparaceae Rhamnaceae Capparaceae Burseraceae Caesalpinioidiae Combretaceae Capparidaceae Euphorbiaceae Mimosoideae Mimosoideae Burseraceae Burseraceae Sterculiaceae Mimosoideae Anacardiaceae Umbelliferae Gittiferae Combretaceae Sterculiaceae Burseraceae Burseraceae Mimosaceae Papilionaceae Anacardiaceae Salvadoraceae Moraceae Palmae Lythraceae Mimosaceae Solanaceae Acanthaceae Combretaceae Tiliaceae Papilionaceae Verbenaceae Capparaceae T T T T S,T T T S,T T S,T T T S,T T T,S S,T S,T T S,T S,T T S,T T S,T T S,T T S,T T S,T T T T T T T S, T T,S T T S,T T S S S,L S S,H S S,T 38.3 25.7 20.8 17.4 14.3 14.1 9.1 7.4 6.8 6.4 5.8 5.8 5.7 5.7 5.4 5.4 5.1 4.7 4.5 4.4 3.8 3.7 3.5 3.4 3.3 3.2 3.2 2.9 2.7 2.6 2.5 2.4 2.4 2.2 2.0 1.5 1.2 1.2 1.0 1.0 0.8 0.8 – – – – – – – © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 12 Edward K. Mengich et al. Table 3 (continued) No. Species local name Botanical name Family name Species type 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. Kinatha Kitanda mboo Mwaanzia Mwindenguwe Mukakaa Mukokola Mukulwa Mukuswi Mulalambila Mukulwa Mung’uthe Musilingu Musovi Mutandi Mutungu Mututi Mutuba Muvatha Muvuluvulu Mwaitha Maerua decumbens Capparis tomentosa Bridelia taitensis Triumfetta macrophylla Premna resinosa Combretum exalatum Acalypha fruticosa Acacia brevispica Hibiscus spp. Acalypha fruticosa Lonchocarpus eriocalyx Grewia fallax Hoslundia opposita Ochna inermis Lannea alata Thunbergia holstii Grewia tembensis Vernonia spp. Opilia celtidifolia Entada leptostachya Capparaceae Capparaceae Euphorbiaceae Tiliaceae Verbenaceae Combretaceae Euphorbiaceae Mimosoideae Malvaceae Euphorbiaceae Papilionaceae Tiliaceae Labiatae Ochnaceae Anacardiaceae Acanthaceae Tiliaceae Compositae Opiliaceae Mimosaceae S S,C S,T S S T S S,T S S S,T S,T S T S,T S S S L C IVI – – – – – – – – – – – – – – – – – – – – T, tree; S, shrub; B, bush; C, climber; L, liana; IVI, important value index. NB: Species indicated as tree (T) but not having a value for IVI was <1.30 m in height. Table 4 Mean basal areas (BAs) of the most important tree species at Nthangu, Kathonzweni and Kibwezi forests of Makueni County Study site Species Mean BA (m2 ha1) Max Min Range S.E n Nthangu Pithosporum viridiflorum Euclea divinorum Combretum molle Searsia spp. Diospyrus mespiliformis Acacia hockii Faurea saligna Terminalia brownii Combretum collinum Combretum apiculatum Acacia tortilis Acacia nilotica Lannea triphylla T. brownii Commiphora campestris Adansonia digitata Delonix elata A. tortilis Commiphora africana Acacia mellifera Acacia elatior 0.13 0.07 0.13 0.02 0.04 0.04 0.13 0.17 0.02 0.004 0.09 0.04 0.02 0.07 0.04 8.47 0.86 0.04 0.05 0.02 0.09 0.363 0.349 0.608 0.102 0.385 0.212 0.385 0.407 0.332 0.132 0.369 0.138 0.053 0.266 0.224 20.79 2.684 0.342 0.363 0.302 0.622 0.006 0.008 0.001 0.001 0.004 0.005 0.014 0.014 0.001 0.002 0.004 0.004 0.003 0.006 0.004 2.290 0.001 0.001 0.001 0.001 0.001 0.358 0.341 0.607 0.101 0.381 0.207 0.371 0.393 0.332 0.130 0.365 0.135 0.050 0.256 0.220 18.50 2.683 0.341 0.362 0.301 0.622 0.004 0.005 0.014 0.000 0.004 0.001 0.008 0.007 0.000 0.000 0.024 0.000 0.000 0.002 0.001 27.09 0.300 0.001 0.001 0.001 0.016 15 15 21 67 32 24 7 5 257 65 47 33 47 14 27 4 8 205 75 123 24 Kathonzweni Kibwezi © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 Indigenous trees and shrubs as adaptation criteria Table 5 Important value indices (IVIs) of trees commonly occurring in Nthangu, Kathonzweni and Kibwezi forests of Makueni County Important value index (IVI) Species Nthangu Kathonzweni Kibwezi Acacia tortilis Acacia nilotica Acacia mellifera Grewia spp. Steganotaenia erialiacea Total 2.8 6.3 1.1 2.0 1.6 13.8 27.6 16.8 7.4 11.4 2.4 65.6 20.8 5.8 14.3 5.8 2.7 49.4 Kenya’s Eastern Province – Makueni, Machakos, Kitui and Mwingi, DRSRS identified a total of 300 tree and shrub species which were used to describe the local vegetation (Ojiambo et al., 2001). In the nearby semi-arid Miombo woodland area of eastern Tanzania, a total of 86 tree species were identified (Back’eus et al., 2006). The low species numbers observed in our study indicate that at some point in time, there may have been a phenomenon, which resulted in a drastic reduction in tree species diversity. Studies elsewhere have attributed similar reductions in tree species diversity to (i) intensified human activities that may have led to elimination of some of the tree and shrub species through wood harvesting, fires, livestock browsing/grazing and agricultural activities (Back’eus et al., 2006; Suleiman, Buchroithner & Elhag, 2012; Van Der Hoek et al., 2013); (ii) natural phenomena such as frequent droughts and wildlife damage (Birkett & Stevens-Wood, 2005) that may have led to loss of some of the more susceptible tree and shrub species; and (iii) salinity of soils, which can only support a limited number of hardy and adapted species (Macharia, 1981). This study did not, however, gather information to support or reject these propositions. Nthangu had the largest number of species, families and genera (77, 33, 60) followed by Kibwezi (70, 30, 48) and Kathonzweni (69, 28, 42). Similar gradients have been documented within the study area and its immediate surroundings by Ego, Kiptot & Ochieng (2001) and Van den Abeele, Ngatia & Macharia (2005). It is suggested that the gradients observed in our study could be mainly a function of the long-term effects of higher rainfall at Nthangu compared with the other two sites. Climate provides one of the strongest controls on the species composition and characteristics of vegetation, and Kenya is thought to owe its high biological diversity to the enormous variation in climate and topography, which © 2013 John Wiley & Sons Ltd, Afr. J. Ecol., 53, 3–15 13 result in a great range of habitats (Coughenour & Ellis, 1993; Back’eus et al., 2006; Yimer, Ledin & Abdelkadir, 2006). Our findings further agree with those obtained in a study conducted in Turkana County of Kenya. In the Turkana study, the regional herbaceous vegetation biomass reflected a gradient in which the highest levels of biomass occurred where rainfall was >600 mm, and the lowest levels occurred where rainfall was estimated at 150 mm–200 mm (Coughenour & Ellis, 1993). Similar gradients have been observed at the Nairobi National Park in southern Kenya (Helsa, Tieszen & Boutton, 1985) and in a geologically homogeneous area in Australia (Myers & Neales, 1984). According to the results of our study and the physiognomic classifications described in the literature (Pratt & Gwynne, 1977; Chidumayo & Gumbo, 2010), vegetation at Nthangu, Kathonzweni and Kibwezi forests may be categorized as Euclea-Combretum-Acacia thicket, Combretum-Terminalia-Acacia woodland, and Acacia-Commmiphora woodland, respectively. In a vegetation survey conducted at sites located at various points within Makueni County, DRSRS identified different vegetation associations dominated by different plant species and genera as shown in this study (Ojiambo et al., 2001). In a separate study, it was difficult to delineate the dominant species in Kikumbulyu location of Kibwezi division due to severe clearing and high population density (Ngoda & Obwoyere, 2001). Based on our study, the families Combretaceae, Burseraceae and Mimosaceae were the most important and widely distributed in the area. Combretaceae was widely represented at Nthangu and Kathonzweni, where it was ecologically the most important at both the sites. Combretum was the most common genus in this family. Burseraceae was the third most important at Kathonzweni and the second most important at Kibwezi. Commiphora was the most common genus in this family. Mimosaceae was ecologically the most important at Kibwezi, second most important at Kathonzweni and third most important at Nthangu forest. Acacia was the most common genus in this family. The observation on Acacia is consistent with results obtained in arid northern Kenya, where the genus Acacia was common and dominant in 29 of 30 sites sampled for a vegetation survey (Coughenour & Ellis, 1993). The ecological importance of these families and genera indicate that they could provide candidate tree species for rehabilitation of respective sites. This is because these species and vegetation categories grew naturally and dominated the respective sites, indicating that conditions prevailing at the sites were favourable for the survival and growth of these 14 Edward K. Mengich et al. species. Based on IVIs in the genus combretum, C. molle would be a priority candidate for rehabilitation at Nthangu followed by T. brownii, while C. collinum would be the right choice at Kathonzweni followed by C. apiculatum. In the genus Burseraceae, C. africana would be preferred at Kibwezi, followed by C. baluensis, while at Kathonzweni, Commiphora campestris would be preferred over C. africana. In the genus Acacia, A. tortilis would be preferred at Kibwezi and Kathonzweni, followed by A. mellifera and A. nilotica, respectively. At Nthangu, Acacia hockii would be the most suitable. Results of the basal area, which showed that these species contributed significantly to the total BA in respective sites, render credence to their use for rehabilitation purposes. Although the genera Pithosporum and Adansonia had high IVIs at Nthangu and Kibwezi, respectively, they would not be suitable candidates for rehabilitation due to their limited spread. 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