Doctoral theses at NTNU, 2017:238

Emmanuel Clamsen Mmas Emmanuel Clamsen Mmassy

Ecology and Conservation Challenges of the Kori

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ISBN 978-82-326-2548-2 (printed ver.) ISBN 978-82-326-2549-9 (electronic ver.) ISSN 1503-8181 Doctoral theses at NTNU, 2017:238 NTNU Philosophiae Doctor Philosophiae Department of Biology Thesis for the Degree of the Degree Thesis for Faculty of Natural Sciences of Natural Faculty sity of Science and Technology sity of Science Norwegian Univer Norwegian Emmanuel Clamsen Mmassy

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Thesis for the Degree of Philosophiae Doctor

Trondheim, September 2017

Norwegian University of Science and Technology Faculty of Natural Sciences Department of Biology NTNU Norwegian University of Science and Technology

Thesis for the Degree of Philosophiae Doctor

Faculty of Natural Sciences Department of Biology

© Emmanuel Clamsen Mmassy

ISBN 978-82-326-2548-2 (printed ver.) ISBN 978-82-326-2549-9 (electronic ver.) ISSN 1503-8181

Doctoral theses at NTNU, 2017:238

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Contents Ecology and Conservation Challenges of the in the Serengeti National Park ...... 1 Preamble ...... 3 List of Papers ...... 7 Declaration of Contribution ...... 7 Summary ...... 9 Introduction ...... 11 Conservation Challenges ...... 11 Biodiversity Conservation Approach ...... 12 Threats to Kori bustard ...... 13 Ecology and Breeding of the Kori bustard ...... 14 Home range, movement and resource selection ...... 16 Thesis Aims...... 17 Methods ...... 17 Study Area ...... 17 Study ...... 19 Data Collection ...... 21 Local Ecological Knowledge and Threat ...... 21 Occurrence in the Grass Plains ...... 22 Breeding Ecology ...... 22 Migration and Resource Selection ...... 22 Major Findings ...... 23 Paper I: Local Ecological Knowledge and Perceived Threats ...... 23 Paper II: Density and Occurrence ...... 24 Paper III: Breeding Ecology ...... 25 Paper IV: Seasonal Migration and Resource Selection ...... 26 Discussion...... 27 Local Ecological Knowledge and Perceived Threats ...... 27 Density and Occurrence ...... 28 Breeding Ecology ...... 29 Seasonal Migration and Resource Selection ...... 30 Prospect for Future Studies ...... 31

5

Conclusions ...... 32 References ...... 33

                 

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List of Papers   3DSHU, 0PDVV\(& 5¡VNDIW(  )DFWRUVDIIHFWLQJORFDOHFRORJLFDONQRZOHGJHDQGSHUFHLYHG WKUHDWVWRWKH.RULEXVWDUG(Ardeotis kori struthiunculus)LQWKH6HUHQJHWL(FRV\VWHP1RUWKHUQ 7DQ]DQLD,QWHUQDWLRQDO-RXUQDORI%LRGLYHUVLW\DQG&RQVHUYDWLRQ@  3DSHU,, 0PDVV\(&)\XPDJZD5'-DFNVRQ&5%HYDQJHU. 5¡VNDIW(  .RULEXVWDUG (Ardeotis kori struthiunculus) RFFXUUHQFH LQ WKH 6HUHQJHWL JUDVV SODLQV QRUWKHUQ 7DQ]DQLD $IULFDQ-RXUQDORI(FRORJ\'2,DMH  3DSHU,,, 0PDVV\(&)\XPDJZD5'%HYDQJHU. 5¡VNDIW( VXEPLWWHG %UHHGLQJHFRORJ\RI.RUL EXVWDUGV Ardeotis kori strunthiunculus)LQWKH6HUHQJHWL1DWLRQDO3DUN   3DSHU,9 0PDVV\(&0D\5-DFNVRQ&5.OHYHQ21\JnUG7%HYDQJHU. 5¡VNDIW( VXEPLWWHG  6HDVRQDOPLJUDWLRQDQGUHVRXUFHVHOHFWLRQLQWKH.RULEXVWDUGLQWKH6HUHQJHWLHFRV\VWHP 

Declaration of Contribution  ,GHFODUHWKDWWKLVWKHVLVLVP\RULJLQDOUHVHDUFKZRUNDQGDOOFRDXWKRUVFRQWULEXWHGWKHLULGHDV GXULQJPDQXVFULSWSUHSDUDWLRQ3KLORVRSKLHVDQGUHVHDUFKGHVLJQGDWDDQDO\VLVDQGPDQXVFULSW SUHSDUDWLRQZHUHFRQGXFWHGZLWKKHOSIURPP\VXSHUYLVRUV3URIHVVRU(LYLQ5¡VNDIWDQG'U .MHWLO%HYDQJHU0DQXVFULSWZULWLQJDQGGDWDDQDO\VLV SDSHU,9 ZHUHSHUIRUPHGZLWKKHOS IURP 'U 5RHO 0D\ DQG 'U 2GGPXQG .OHYHQ 'U &UDLJ -DFNVRQ FRQWULEXWHG LGHDV LQ PDQXVFULSWZULWLQJDQG$UF*,6PDSVGUDZLQJV'U5REHUW)\XPDJZDDQG'U7RUJHLU1\JnUG FRQWULEXWHGWKHLUFRPPHQWVLQ3DSHUV,,DQG,,,DQG3DSHU,9UHVSHFWLYHO\  

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10 Introduction

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11

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13

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Ecology and Breeding of the Kori bustard  7KH.RULEXVWDUGRFFXUVDWORZGHQVLWLHVDQGLVZLGHVSUHDGLQ$IULFD,QVRXWKHUQDQGHDVWHUQ $IULFDWKHVSHFLHVLVIRXQGLQWKHVDYDQQDKDQGRSHQEXVKODQGVDUHDVDQGUDUHO\REVHUYHGLQ WKHZRRGODQG &ROODU %UHHGLQJKDELWDWVRXWVLGH3$VDUHGLIILFXOWWRILQGEHFDXVHRIUDSLG SRSXODWLRQJURZWKDQGVXLWDEOH.RULEXVWDUGKDELWDWVEHLQJFKDQJHGIRUDJULFXOWXUDOSXUSRVHV ,QWKH6HUHQJHWLWKHVSHFLHVLVPRVWO\VHHQLQWKHVKRUWJUDVVSODLQ JUDVVOHQJWKUDQJHVFP WRPHWUH RIWKHVRXWKHDVWHUQSDUWVRIWKHHFRV\VWHP 6LQFODLU 

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14



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15

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16

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Thesis Aims

The main objective of this study has been to learn about the overall ecology of Kori bustard, with four sub aims;  6WXG\IDFWRUVDIIHFWLQJORFDOHFRORJLFDONQRZOHGJHDQGSHUFHLYHGWKUHDWVWRWKH.RULEXVWDUG LQWKHQRUWKHUQ6HUHQJHWLFRPPXQLWLHV 3DSHU,   6WXG\.RULEXVWDUGGHQVLWLHVLQWKH6HUHQJHWLJUDVVSODLQV 3DSHU,,  6WXG\EUHHGLQJHFRORJ\RI.RULEXVWDUGVLQWKH6HUHQJHWL1DWLRQDO3DUN 3DSHU,,,    6WXG\ VHDVRQDO PLJUDWLRQ DQG UHVRXUFH VHOHFWLRQ RI WKH .RUL EXVWDUG LQ WKH 6HUHQJHWL HFRV\VWHP 3DSHU,9 

Methods

Study Area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

17

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18

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Study Species

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Major Findings

Paper I emphases the ability to identify Kori and understand the local ecological knowledge and factors leading to the decline of the species among local people close to Serengeti National Park. Paper II focuses on the density and occurrence of the species in relation to the season, area, grass height and colour. Paper III gives highlights on the breeding signs, season, area, habitat and number of nests and eggs. Paper IV focuses on movement of the species during different seasons and types of habitat utilized by the species in different seasons.

Paper I: Local Ecological Knowledge and Perceived Threats

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Paper II: Density and Occurrence

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Paper III: Breeding Ecology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Paper IV: Seasonal Migration and Resource Selection

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Paper I    Vol. 6(6), pp. 459-467, June 2014 DOI: 10.5897/IJBC2014.0719 Article Number: C40FFE345313 International Journal of Biodiversity ISSN 2141-243X Copyright © 2014 and Conservation Author(s) retain the copyright of this article http://www.academicjournals.org/IJBC

Full Length Research Paper Factors affecting local ecological knowledge and perceived threat to the kori bustard (Ardeotis kori struthiunculus) in the Serengeti Ecosystem, Northern Tanzania

Emmanuel Clamsen Mmassy1 and Eivin Røskaft2*

1Tanzania Wildlife Research Institute (TAWIRI), PO Box 661, Arusha, Tanzania. 2Department of Biology, Norwegian University of Science and Technology (NTNU), N-7491, Trondheim, Norway.

Received 21 April, 2014; Accepted 23 May, 2014

This study examines local tribal knowledge regarding the ecology of the kori bustard (Ardeotis kori struthiunculus) and assessed threats to this species in Northern Serengeti communities. A picture of an indigenous kori bustard was presented to survey participants in villages in the study area. General knowledge on the kori bustard was tested in relation to the ’s general habitat, nesting habitat, food and number of individuals in groups. Of the survey respondents, 56.7% knew the name of the kori bustard and were therefore included in further analyses. The Maasai tribe showed the greatest knowledge of the species, with 98% of individuals identifying the species correctly. Additionally, male survey participants were generally more knowledgeable than females. No differences among age groups or individuals with different education levels were found, suggesting that there is a local knowledge transfer of the species to all age groups regardless of educational level of respondents and that education is not an obstacle to the local knowledge. The study concludes that nature of activities e.g. nomadic and social life, gender and tribes were contributing factors to the knowledge of the kori bustard in the northern Serengeti.

Key words: Local knowledge, kori bustards, Serengeti ecosystem.

INTRODUCTION

The study of local ecological knowledge (LEK) has been resources (Berkes, 1999; Colding, 1998; Johannes, growing as a scientific field in recent years, partly due to 1998). LEK is the informal, often explicit knowledge held the recognition that such knowledge can contribute to the by a specific group of people about their local management of various ecosystems and ecological ecosystems and includes the interplay between processes, the conservation of biodiversity and rare and organisms and their environment (Olsson and Folke, threatened species, and the sustainable use of natural 2001). LEK differs from ‘traditional’ ecological knowledge

*Corresponding author. E-mail: [email protected].

Author(s) agree that this article remain permanently open access under the terms of the Creative Commons Attribution License 4.0 International License 460 Int. J. Biodivers. Conserv.

(TEK) in the sense that the former has been derived from and education through conservation experiments and/or more recent human-environment interactions (e.g. those natural resource management in a specific project area occurring within a few generations) rather than being that is related to local ecosystem conservation (Drew and embedded in deeper cultural practices (Ohmagari and Henne, 2006). Indigenous people are frequently experts Berkes, 1997). on their local ecosystems and can have a broad Analyses of many LEK systems have identified various spectrum of indigenous knowledge. Rapid local language components of LEK, including a component related to shift to the global language might however, make them local observational knowledge of species and other lose this intellectual knowledge. Many cultures have environmental phenomena, a practical component related disappeared as a result of indigenous people exhausting to resource use activities, and a belief component related the environment’s ability to sustain their population to how people fit into or relate to ecosystems (Berkes et (Mazzocchi, 2006). Further-more, such disappearance of al., 2000). The understanding of LEK plays a major role indigenous culture has been exacerbated by European in aiding and promoting the improvement of scientific colonization, which has eroded and destroyed much research and the management of ecosystems (Berkes et traditional knowledge by replacing it with Western al., 2000). educational and cultural systems (Mazzocchi, 2006); The study of LEK began with species identification and economic development and the transition to market classification (ethnobiology) and evolved to include the economies (Godoy et al., 2005); loss of access to study of people’s understanding of ecological processes traditional resources due to conservation policies; and and their relationships with the environment (Berkes, more generally, the forces of industrialization and 1999; Williams and Baines, 1993). Not all traditional globalization (Turner and Turner, 2008). Local and practices and belief systems are ecologically adaptive, traditional ecological knowledge is increasingly and they can change with time. Acknowledging the recognized as an important component of scientific importance of LEK is commonly assumed, incorrectly, research, conservation, and resource management, that indigenous peoples sustainably conserve natural especially where LEK and TEK fill gaps in the scientific resources (Redford and Sanderson, 2000). Local literature or offer a critical source of basic environmental traditional knowledge has showed an intergenerational data (Thornton and Scheer, 2012). decline and may vary with age and gender (Gómez- Gathering scientific information regarding local Baggethun et al., 2010; Law et al., 2010; Quinlan and ecological knowledge of kori bustard has never been Quinlan, 2007). carried out in the Serengeti ecosystem. In this study we Local ecological knowledge about medicinal plants and found that 1) illegal hunting among local people for food the identification of traditional foods appears to decrease exists, 2) kori bustards are rarely observed indicating a with increasing formal education (Giovannini et al., 2011; low population size, and 3) that their nesting habitat Wester and Youngvanit, 1995). Thus, formal education outside the national park has been turned into agricultural may contribute to the loss of local traditional knowledge land and human settlements. Thus it is obvious that the because it reduces the time and number of opportunities species is in unimaginable threat. The outcome of our that children have to attain local traditional knowledge study has provided us an image that the species needs to and skills from their elders (Heckler, 2002; Luoga et al., be closely monitored and that local communities/tribes 2000). who are more knowledgeable on kori bustards are to be Local knowledge and its applications seem to be fully involved in conservation of the species to provide declining owing to a combination of factors, including loss their ideas to the management authority on how to of local language, land use change, inaccessibility to conserve this species in the future. traditional resources due to conservation programmes, industrialization and globalization, and the transition to market economies (Benz et al., 2000; Kingsbury, 2001; Study species Turner and Turner, 2008). Because this loss of know- ledge affects local communities in developing countries, Although kori bustards are used by tribes in the Serengeti the remaining local ecological knowledge must be ecosystem as a source of food in household diets safeguarded through a number of methods, including (Magige et al., 2009), there is no clear evidence that the environmental policies designed to protect its pools species is used in other social cultural, or economic (Montes, 2010). contexts. In Tanzania, the current range of the kori Local ecological knowledge can be maintained in bustard is restricted to the northern plains of the traditional communities by controlling and monitoring Serengeti and the Tarangire-Manyara ecosystems. The access to certain sites and resource use. This may result range of this sub-species where it was historically found in improvements of the knowledge base, which can then (in parts of East , , and ) has be used to respond adaptively to change in certain been shrinking, such that its current range is much environments. Through the use of particular local know- smaller than its historical range (Hallager and Boylan, ledge, a community may benefit by securing employment 2004). Mmassy and Røskaft 461

The kori bustard is listed by CITES (Appendix II) as (Maddock, 1979) of which during the northern migration, the near threatened (Birdlife-International, 2009; 2013) and wildebeests often travel out of the protected area and enter faces many threats. The main threats to the kori bustard unprotected areas with comparatively high human population densities. During the migration season, kori bustards are are human-induced, including habitat destruction through associated with the migrating and may be vulnerable to agricultural development, as well as shrub encroachment snares used by poachers to catch large animals. The villages found caused by overgrazing and subsistence hunting (Magige along the Serengeti ecosystem are fairly typical of the region, in et al., 2009; Senyatso et al., 2013). The poison used to that, hunting of wildlife for food is still a common practice (Holmern control locusts is toxic to and may also be affecting et al., 2007). the kori bustard populations (Barnes, 2000). The kori bustard is an omnivorous species, and its food sources Study communities are expected to be quite diverse. However, according to Arlott (1996), kori bustards consume mostly and The Northern Serengeti is highly diverse in terms of ethnicity. Over plant material in their grassland habitat. The population 20 tribes live in the area. We were able to interview individuals from status of this species and its population trends remain the major and the largest tribes, including migrants. Individuals from the Meru, Chaga, Iraq, Mbulu and Sonjo tribes are immigrants to unknown, both in Tanzania and the entire . the northern Serengeti. Agro-pastoralism plays a major role in the Thus, local understanding of ecology and threat of the livelihoods of these tribes. Although we interviewed people from 18 kori bustard may provide inputs that management tribes in the selected villages, we used only the data from the three authorities can apply for sustainable management of largest tribes (Maasai, Kurya and Ikoma) in our analyses because areas where the small range of species occurs. these tribes had sufficient number of survey respondents. There are Since people in the northern Serengeti live a traditional no inter-tribal conflicts among the interviewed tribes; however, the health of the ecosystem suffers due to conflicts between life style that includes hunting wildlife for food and as a conservationists and local communities. The conflict stems from the source of income, we hypothesised that people in our conservation and management authority prohibition of access to study area might have LEK of the kori bustard (Ardeotis natural resources for many tribes living in the area (Kideghesho et kori struthiunculus). We also hypothesised that there al., 2007). The Maasai are nomadic, herding their livestock to green would be differences in local knowledge of this species’ pastures and water. The Ikoma have traditionally hunted wild animals as a source of income and food, while the Kurya have ecology among different tribes. We further hypothesised traditionally been primarily farmers, but practice hunting activities to that the threats faced by the kori bustard are well some degree. understood by the people of the northern Serengeti Generally, the people in the region are poor (US $150-200 per because they live in the villages adjacent to the Serengeti annum). Their annual incomes, which are generally earned through National Park where the species is found. Scientific agro-pastoralism, are far lower than Tanzania’s average per capita information about this species’ ecology and the factors income of US $280 (WB, 2003). To compensate for the hardship of earning a low income, many communities in the area engage in influencing the major threats to the species is limited. The illegal hunting and charcoal burning (Kideghesho, 2010). With the goal of our study was to provide information on whether increasing rates of human and livestock population growth in the the species needs to be monitored, which includes the region, these activities have caused ecosystem fragmentation, possibility of local community-based monitoring because natural resource depletion and habitat destruction (Kideghesho, the species utilizes habitat found on communal lands. 2010), including the extinction of bird species of conservation The outcomes of the study could also help management concern. authorities consider impacts to this species when developing various conservation programs in the Data collection using questionnaires communities surrounding protected areas. Local knowledge could provide broad insights into kori bustard The study site was selected as an example of the type of socio- conservation that could be used by management ecological context that occurs where communal areas are adjacent to protected areas. However, the site was also selected because of authorities when developing management plans. the conflict between local communities and management authorities towards access to wildlife resources. Fieldwork was conducted in 13 villages of the northern Serengeti METHODOLOGY that were selected prior to beginning of the survey in October, 2011. The villages were selected based on their proximity to Study area protected areas (Loliondo Game Controlled Area, Ikorongo Game Reserve, Ikona Open Area, and Serengeti National Park), where The mean temperatures in the study area range between 15 and we expected the kori bustard to be frequently observed. 27°C, while the mean annual seasonal rainfall varies from 1,050 to The closed-ended question interviews (Newell, 1993) were 1200 mm (Sinclair et al., 2000). According to United Republic of conducted in Kiswahili with the aid of well-trained local translators Tanzania (URT) (2003, 2013), the study area has a total population who translated Kiswahili into the local tribal languages because of approximately 40,000 people. most elders knew only their tribal language. The use of local The Serengeti ecosystem covers an area of 25,000 km2 and translators increased the number of people who could be straddles the border of northern Tanzania and southern . The interviewed, because the translators could communicate with ecosystem comprises several different conservation areas (Figure respondents in their local language. Furthermore, because the 1) that harbour a variety of animals such as wildebeests respondents live adjacent to the protected areas and illegal (Connochaetes taurinus), antelopes, carnivores and birds. A yearly subsistence hunting of wildlife still occurs, they were initially migration of animals including wildebeests occurs in the area concerned about talking with us because they thought we were 462 Int. J. Biodivers. Conserv.

Figure 1. Map of the Serengeti ecosystem showing the locations of surveyed villages in the northern Serengeti.

wildlife officers who wanted to investigate poaching including the kori bustard. The current proposed road construction from Mto wa Mbu to Musoma town (Fyumagwa et al., 2013) was another barrier to recruiting survey participants, as some people thought we were environmentalists or conservationists who wanted to convince the local people to oppose the proposed road construction. This problem was resolved by the village leaders who convened a village meeting where he expressed our aim to interview people in the area. During the survey, a picture of the indigenous kori bustard (Figure 2) was shown to the participants. All respondents were between 18 and 85 years of age and were able to recognise the picture as a bird. However, not all respondents were able to identify the species. Face-to-face interviews were conducted in which the respondents were asked different questions after identifying the picture of the bird as kori bustard (Mmassy and Røskaft, 2013). A total of 330 individuals were randomly picked from households in the selected villages, of which 197 individuals (Table 1) were able to identify the kori bustard by looking at the picture. We stratified our sample of interview respondents based on tribe, gender, age and education level to test for species knowledge. In our analyses, we included only the three most common tribes (Maasai, Ikoma and Kurya) and 25 individuals from a combination of other smaller tribes (Luo, Chaga, Jita, Iraq, Mbulu, Meru, Natta, Figure 2. Kori bustard on the Serengeti plains (photo: E. Maragoti, Mungurumi, Zanaki, Ikizu, Mwira, Sukuma, Isenye, Sonjo) Røskaft). who identified the kori bustard correctly. Mmassy and Røskaft 463

Table 1. Knowledge variables related to kori bustard biology derived from the responses of interview participants and the validation of their knowledge using literature.

Variable Correct knowledge Incorrect knowledge No. of respondents How often do you see the Rarely: 80.6% Frequently: 19.4% 191 species Population size Small: 91.0% Large: 9.0% 162 Group size 1-2 individuals: 66.8% >3 individuals: 33.2% 129 Nesting habitat Open grasslands 90.5% Mixed woodlands 9.5% 137 insects, reptiles, small insects, reptiles, small Food , , and mammals, seeds, and 190 90% roots 10% Other, such as accidents, Threats Illegal hunting 77.6% 98 22.4% Sources: Birdlife-International, 2008; Hallager, 2012; Harrison et al., 1997. Respondents who provided a correct answer were designated as having correct knowledge.

Table 2. Analysis of the general knowledge of the kori bustard among people of the northern Serengeti in relation to gender, tribe, educational level and age groups.

Dependent variable/general knowledge (%) Constant/independent Poor Week Average Good Very good Total ² =; p = Male 35.7 7.1 22.0 24.7 10.6 255 Gender 19.17; 0.002 Female 62.7 4.0 14.7 14.7 4.0 75

Kurya 83.5 5.8 4.9 4.9 1.0 103 Maasai 2.8 5.6 27.1 38.3 26.2 107 Tribe 184.1; 0.000 Ikoma 36.8 8.4 31.6 23.2 0.0 95 Others 56.0 4.0 12.0 24.0 4.0 25

No or primary 40.7 7.0 20.5 22.3 9.5 273 education Education 2.298; 0.807 Secondary 47.4 3.5 19.3 22.8 7.0 57 education

14-45 years 42.9 5.1 19.8 23.0 9.2 217 Age Above 45 6.118; 0.295 39.8 8.8 21.2 21.2 8.8 113 years

The respondents were asked various questions about the two categories: observed daily (at least once per week or more) species, such as how often the bird was observed, the habitats in and observed rarely (less than once per week). Information on food which it was most frequently observed, the species’ population size, availability, safety from predators and the use of different habitats why the bird preferred to live in the habitat where it is observed, was separated into two categories: food availability and safety from how many individuals were observed at a time per group, where it non-human predators and hunters. Species occurrence was placed its nests, why it uses a particular breeding habitat, the food categorised as a single individual, two individuals, or greater than source of the species, and threats to the species. two (3-20) individuals. Local people in the Serengeti ecosystem could identify the The information provided by the respondents was species to the level (bustard). The identification to the genus assessed/analyzed and compared to information from various level, that is, bustard, was done following Balmford et al. (2002), literature sources (Table 1) to validate the respondents’ knowledge since mammals and birds need genus level identification (e.g., as correct or incorrect (Table 1). Respondents who provided correct ‘rabbit’). We grouped some items of knowledge that varied among answers were designated having correct knowledge and those who only a small number of respondents. For example, and could not provide correct answer were designated providing other small mammals listed as food sources were recorded as incorrect knowledge (Table 1). The six knowledge variables were mammals, while all types of reptiles ( and ) were thereafter pooled and varied between 0 and 6. Individuals with recorded as reptiles. Seeds, , , fruits and roots were knowledge for 0 or 1 of the variables were given a value of 1 (poor), recorded in a single category as plant materials. All woodland, followed by those who had knowledge of 2 variables (weak), those forest and shrub land habitats were combined and designated as who had knowledge of 3 (average), those who had knowledge of 4 mixed woodlands, while open grassland was retained as a separate (good), and those who had knowledge of 5-6 variables (very good) category. Frequency of observation of the species was recorded as (Table 2). 464 Int. J. Biodivers. Conserv.

Statistical analysis DISCUSSION

The information included in the analyses was collected through The majority of the interview participants from tribes in questionnaire surveys that were coded and entered into a the northern Serengeti knew the kori bustard and showed computer. Statistical Package for the Social Sciences (SPSS), version 16, manufactured by SPSS Inc., was used for the analyses. good knowledge of the bird’s general ecology. The Chi square analyses were primarily used to calculate the overall good knowledge of members of the Maasai and percentages of respondents at a significance level of p = 0.05. Ikoma tribes on the biology of and threats to the kori bustard may be due to the nature of the activities (hunting and nomadic migration) that have been practiced by RESULTS these tribes since historical times. Given that the Maasai tribe is pastoralist and nomadic in nature, its members Of the individuals interviewed, 59.7% (N = 330) correctly are likely to encounter this species frequently (Mmassy identified the picture as a kori bustard, while 40.3% (N = and Røskaft, 2013). The knowledge and understanding of 133) were unable to identify the species. The Maasai kori bustard ecology within the Ikoma tribe may be due to tribe was the tribe with the highest level of knowledge of their hunting practices and the inheritance of hunting the species, with 97% of individuals identifying the knowledge from their ancestors, because hunting has species correctly (N = 107). However, only 66% of the been an integral part of life in Serengeti for thousands of interviewed individuals from the Ikoma tribe (N = 95) and years (Holmern, 2010). The Ikoma have traditionally been 18% from the Kurya tribe (N = 103) were able to identify a hunting tribe that collects eggs from game birds the kori bustard from the picture. Among the other (Magige et al., 2009) and hunts wild animals for food and surveyed tribes, 44% of the individuals surveyed were income (Barnett, 2000). This type of practice could have able to identify the kori bustard correctly (N = 25). The increased their knowledge of a species like the kori difference among tribes in identifying the kori bustard was bustard. highly significant (² = 139.7, df = 3, P< 0.001). There Male respondents generally showed better knowledge was a highly significant difference between the genders, of kori bustard biology and were better able to identify the with males correctly identifying the bird more often than bird from a picture. This result may reflect the division of females (² = 17.88, df = 1, P< 0.000). In contrast, there household tasks in the villages in this study. Males were no significant differences in ability to identify the bird conduct outdoor activities, while females practice more among age groups or individuals with different education indoor activities (Mmassy and Røskaft, 2013; Røskaft et levels (² = .848, df= 1, P = .357; ² = .477, df = 1, P = al., 2004). According to Pfeiffer and Butz (2005), Maasai .490, respectively). women are responsible for taking care of domestic Generally, most of the interviewed individuals knew the animals in small confined areas around their homes, kori bustard well, and they displayed a high level of while males graze herds of cattle across large knowledge of the species with regards to frequency of geographical areas. observation, population size, group size, nesting habitat, Females are frequently less knowledgeable about food sources and threats to the species (Table 1). The wildlife, including birds around the world (Kideghesho et Maasai tribe showed a high level of knowledge of the al., 2007; Mmassy and Røskaft, 2013; Røskaft et al., biology of the kori bustard overall, with 74.5% of the 2007). However, ethnobiological research suggests that respondents showing good to very good knowledge of women are more knowledgeable about medicinal plants the bird. The second most knowledgeable tribe was the and small mammals because they are more engaged in Ikoma, where 54.8% of individuals exhibited good to very healing practices and the collection of food (Letsela et al., good knowledge. The Kurya tribe was the least 2003). This type of knowledge can, on the other hand, knowledgeable, with only 5.9% of the respondents having vary with geographical location, social status, ethnicity, good to very good knowledge (Table 2; 2 = 46.8, df = 12, occupation and experience (Heckler, 2002). Therefore, P< 0.000). Men had significantly more knowledge than we suggest that the different societal roles of the two women (Table 2), but there were no detected differences genders in our study may have resulted in greater in knowledge with regards to the age or education level of knowledge of birds among men (Røskaft et al., 2004). the respondent (Table 2). Formal education is not necessarily a prerequisite for A linear regression analysis with general knowledge as having local knowledge of a certain place. In fact, the dependent variable and gender and tribe (the two research in the fields of ethnobiology, natural food variables that were significant in the analyses described identification and ornithology has revealed variation in above) as independent variables explained 8.8% of the local knowledge between people with a formal education variation in knowledge (r² = 0.088). The results showed and those without education. In this respect, educated that, in general, tribe (Beta = 0.194, T = 3.611, P< people might be less knowledgeable than uneducated 0.0001) and gender (Beta = 0.191, t = -3.558, P <.0001) ones (Giovannini et al., 2011; Mmassy and Røskaft, independently explained some of the variation in 2013). In our study, we found that the ability of knowledge of kori bustard ecology (ANOVA; F = 15.685, respondents to identify the kori bustard was similar P< 0.000). without considering the level of education. This suggests Mmassy and Røskaft 465

that LEK of kori bustard has been shared among the The most serious threat to the kori bustard in the indigenous people of all age groups and education levels northern Serengeti, according to the respondents in this in the study area. Again if the kori bustard has been used study, is illegal hunting. This result may act as a as food source in the study area it is possible for almost challenge to management authorities to take appropriate every individual to acquire some knowledge of the action because the management authority has never species. The yearly association of the kori bustard with been informed about the presence of illegal hunting of the migration (wildebeest) may also have kori bustards. The threats to the kori bustard may be contributed to the increased identification knowledge of more serious than previously thought, given that its the inhabitants. population size is not known and, according to this study, Most respondents displayed correct knowledge (accor- the bird is now rarely seen in the Serengeti ecosystem. ding to our comparison with literature sources) of the kori The respondents claimed that the greatest threats to the bustard by stating that it was rarely observed in the kori bustard are from the hunting of these birds for food. Serengeti ecosystem. These infrequent observations of Bird conservation has been strongly dependent on the the species may indicate a declining population or low available biological information (Filion, 1987). The LEK population density or both. According to respondents, the presented by respondents will largely help in the rarity of this species might be due to habitat loss as also development of conservation priority strategies and stated by Hallanger and Boylan (2004) and/or being implementation of the species conservation program. The hunted for food (Magige et al., 2009). According to outcome of a study of the species will give information Hallager and Boylan (2004), both subspecies of kori whether the species needs to be monitored, including bustards are facing an uncertain future and birds are local community-based monitoring as the species utilize absent in areas where they were previously found. communal lands. It might also help the management However, a population trend for this species was not authority to take into account this important species when available at the time of this study because the species’ developing various conservation programs on population has not yet monitored in the study area. The communities surrounding protected areas. Such local range of the kori bustard is much smaller than it was a knowledge might provide broad insight into management few years ago due to the fact that the kori bustard is authorities when developing aspects of the species presently declining throughout its range (Hallager and management plan. LEK of a species matters as different Boylan, 2004). This trend is occurring in the northern local knowledge can be shared between stakeholders Serengeti, where people reported that the species is which might lead to an extensive knowledge of a species rarely observed, as there is an increase in human in its surroundings population and the demand of resources in the area. As has been reported in other parts of southern Africa (Hallager, 2012), the rarity of the species in the study Conclusion area may probably be due to rapid human population growth and activities associated with development. Local ecological knowledge can help traditional Given that the gathering of eggs from most large community-based systems control and monitor access to grassland birds, including kori bustards, is tradition sites and resources, thereby protecting natural resources. among tribes living close to Serengeti National Park To conserve A. kori struthiunculus for future generations, (Magige et al., 2009), it would be possible for the tribes to we recommend additional research on this species. In develop accurate knowledge of the decline of the particular, information is needed on population status, species, even though there are no existing data for the threats (apart from hunting for food) and ecological Serengeti population. requirements. Additionally, public education would help In our study area, there was a high level of knowledge prevent degradation of this species’ habitat and minimize on the nesting and feeding habitats of the kori bustard. illegal hunting, and could be effective measures for Local knowledge was consistent with the literature, which preventing further population declines. shows that the kori bustard nests in grasslands (Harrison et al., 1997). It is clear that LEK can strengthen conser- Conflict of Interests vation programs by providing information on a given species (Drew, 2005). Some studies have suggested that The author(s) have not declared any conflict of interests. local people’s knowledge have been used as input for designing and applying management plans for sus- tainable development, especially in protected areas Acknowledgements (Agrawal, 2000; Papageorgiou and Vogiatzakis, 2006). In the case of the kori bustard in the Serengeti, We express our sincere thanks to the Government of management authorities must work with local people in Norway, Ministry of Foreign Affairs for their financial to achieve successful conservation of the kori support through an IPBES (Intergovernmental Platform of bustard (Ghimire and Pimbert, 1997). Biodiversity and Ecosystem Services) grant, without which 466 Int. J. Biodivers. Conserv.

this study would not have been possible. We are very history of ecosystem services in economic theory and practice: From grateful to Dr. Robert Fyumagwa, Director of the early notions to markets and payment schemes. Ecol. Econ. 69: 1209-1218. Serengeti Wildlife Research Centre, for logistic support Hallager S (2012). Kori Bustard, Biology and natural history. Retrieved and to Mr. Noel Massawe for field assistance during the 30.05.2012, 2012, from questionnaire survey. Thanks are also due to the Director http://nationalzoo.si.edu/SCBI/TropicalEcosystems/KoriBustards/biolo General of TAWIRI, Dr. Simon Mduma, and to the gy.cfm. Hallager S, Boylan J, Eds (2004). Kori bustard species survival plan Commission for Science ant Technology, Tanzania (Ardeotis kori) husbandry manual. Washington DC, National (COSTECH) for granting permission to conduct such a Zoological Park. unique type of study. 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watershed, Sweden. Ecosysytems 4:85-104. Thornton TF, Scheer AM (2012). Collaborative engagement of local and Papageorgiou K, Vogiatzakis IN (2006). Nature protection in Greece: An traditional knowledge and science in marine environments: A review. appraisal of the factors shaping integrative conservation and policy Ecol. Soc. 17:3. effectiveness. Environ. Sci. Policy 9:476-486. Turner NJ, Turner KL (2008). Where our women used to get the food: Pfeiffer JM, Butz RJ (2005). Assessing cultural and ecological variation cumulative effects and loss of ethnobotanical knowledge and in ethnobiological research: the importance of gender. J. Ethnobiol. practice; case studies from coastal British Columbia. Botany 25(2):240-278. 86(1):103-115. Quinlan RJ, Quinlan MB (2007). Parenting and cultures of risk: A URT (2003). Population and housing censuses. Bureau of Statistics comparative analysis of infidelity, aggression & witchcraft. Am. PsOPC. Dar es Salaam, Tanzania, United Republic of Tanzania, Anthropol. 109:164-179. Government Printer. Redford KH, Sanderson SE (2000). Extracting humans from nature. URT (2013). Population and housing census counts 2012. Conserv. Biol. 14(5):1362-1364. http://www.nbs.go.tz, Tanzania, National Bureau of Statistics. Røskaft E, Hagen ML, Hagen TL, Moksnes A (2004). Patterns of WB (2003). World Development Indicators. from outdoor recreation activities among Norwegians: an evolutionary http://devdata.worldbank.org. approach. Ann. Zool. Fennici. 41(5):609-618. Wester L, Youngvanit S (1995). Biological diversity and community lore Røskaft E, Händel B, Bjerke T, Kaltenborn BP (2007). Human attitudes in northeastern Thailand. J. Ethnobiol. 15(1):71-87. towards large carnivores in Norway. Wildl. Biol. 13(2):172-185. Williams NM, Baines G, Eds (1993). Traditional ecological knowledge: Senyatso KJ, Collar NJ, Dolman PM (2013). Assessing-range wide wisdom for sustainable development. Canberra, , Centre for change in an unmonitered widespread African Resource and Environmental Studies, Australian National University. bird species. Divers. Distrib. 19(2):106-119. Sinclair ARE, Mduma SAR, Arcese P (2000). What determines phenology and synchrony of ungulate breeding in Serengeti? Ecology 81(8):2100-2111. Paper II Kori bustard (Ardeotis kori struthiunculus) occurrence in the Serengeti grass plains, northern Tanzania

Emmanuel C. Mmassy1,2*, Robert D. Fyumagwa1, Craig R. Jackson3, Kjetil Bevanger3 and Eivin Røskaft2 1Tanzania Wildlife Research Institute, P. O. Box 661, Arusha, Tanzania, 2Department of Biology, Norwegian University of Science and Technology (NTNU), N-7491 Trondheim, Norway and 3Norwegian Institute for Nature Research, Høgskoleringen 9, N-7491 Trondheim, Norway

Abstract la hauteur de l’herbe, de la saison et de la localisation The kori bustard (Ardeotis kori struthiunculus) is indigenous sur la densite/la presence de cette espece dans les plaines to grasslands and lightly wooded savannahs of southern herbeuses du Serengeti, en Tanzanie. Nous avons collecte and eastern Africa. The species is categorized as near des donnees entre janvier 2014 et juin 2015 en faisant threatened in its entire range due to anthropogenic factors des comptages sur transects pendant quatre saisons : (i) and low reproductive rates. The aim of this study was to petite saison seche, (ii) grande saison des pluies, (iii) analyse the impact of grass colour, grass height, season grande saison seche et (iv) petite saison des pluies. La and location on the density/occurrence of this bird species densite moyenne d’outardes kori dans les plaines her- Æ ² in the Serengeti grass plains, Tanzania. Data were collected beuses etait de 0.25 1.01/0.2 km avec des differences from January 2014 to June 2015 using transect counts in quasi significatives entre les sites etudies. La frequence four seasons: (i) short dry, (ii) long rain, (iii) long dry and des outardes kori etait elevee dans les herbes de hauteur – (iv) short rain seasons, respectively. The mean density of moyenne (11 30 cm) pendant la longue saison des kori bustard in the grass plains was 0.25 Æ 1.01 per 0.2 pluies et la petite saison seche. La densite d’outardes km2 with near-significant differences among the study kori est relative faible, et leur distribution varie avec la sites. The occurrence of kori bustard was high in the hauteur de l’herbe et les saisons. Nous suggerons medium height (11–30 cm) during the long rain and short d’orienter les efforts de recherche vers la prevention de dry seasons. The kori bustard density is relatively low, and leur extinction locale, en protegeant l’habitat contre ^ the distribution varies with grass height and season. We l’exces d’activites humaines, tels le paturage du betail et suggest that conservation efforts should be directed at les prelevements illegaux. preventing its local extinction by protecting the habitat from excessive human activities, such as livestock grazing and illegal offtake. Introduction Key words: Kori bustard, northern Tanzania, occurrence, season, Serengeti plains The future survival of wildlife is threatened by increasing human pressure due to high human population increases in many countries in sub-Saharan Africa (Abbitt, Scott & Resume Wilcove, 2000; Ceballos & Ehrlich, 2002; Balmford, Green L’outarde kori (Ardeotis kori struthiunculus)frequente les & Jenkins, 2003; Gaston, Blackburn & Goldewijk, 2003; prairies et les savanes peu boisees d’Afrique de l’Est et du Harcourt & Park, 2003). For a significant period of time, Sud. L’espece est classee « quasi menacee » dans la habitat fragmentation and loss has caused displacement totalite de son aire de repartition a cause de facteurs and extinction of threatened bird species in various anthropiques et d’un faible taux de reproduction. Le but ecosystems (Alroy, 2001; McKee et al., 2004; Senyatso, de cette etude etait d’analyser l’impact de la couleur et de Collar & Dolman, 2013). The largest kori subspecies (Ardeotis kori) is indigenous *Correspondance: E-mail: [email protected] to the grasslands and lightly wooded savannahs of

© 2016 John Wiley & Sons Ltd, Afr. J. Ecol. 1 2 Emmanuel C. Mmassy et al. southern and eastern Africa. The nominate subspecies Hallager & Lichtenberg, 2007; Lichtenberg & Hallager, Ardeotis kori kori occurs in , , , 2008). southern , South Africa and (Johns- Birds’ density is often positively correlated with habitat gard, 1991), while Ardeotis kori struthiunculus occurs in suitability and a sign of minimum anthropogenic distur- Ethiopia, Kenya and Tanzania. Seemingly, the species has bance (Huhta, Mappes & Jokimaki, 1996; Martin, 1998; undergone considerable population decline in all its range Clark & Shutler, 1999; Robertson & Hutto, 2006). states, except in and Angola. In East Africa, the Serengeti National Park holds the largest natural grass- population of Ardeotis kori struthiunculus had declined by lands in the world and is famous for its large aggregates of 21% by the late 19th century (Senyatso, Collar & Dolman, mammals and birds. However, the bird community struc- 2013). The species is listed in Appendix II of CITES and ture of these grasslands has been poorly studied categorized as near threatened by the IUCN (BirdLife- (Gottschalk, Ekschmitt & Bairlein, 2007). Maintaining International, 2013). the Serengeti National Park grasslands will provide critical Kori bustard (Ardeotis kori spp.) populations are dramat- refuges for grassland birds, including kori bustards, ically declining in both open and protected areas that are Karamoja apalis (Apalis karamojae), Francolins, secretary close to human settlements due to illegal hunting, habitat bird (Sagittarius serpentarius) and all vulture species, all of loss, low reproductive rates, the disappearance of large which have shown disturbing declines in recent decades mammals and collision with power lines (Dale, 1990; (Herremanns, 1998; Sinclair, Mduma & Arcese, 2002). Collar, 1996; Lawes, Fly & Piper, 2006; Thiollay, 2006; Therefore, for effective management and conservation Manu, Peach & Cresswell, 2007; Magige et al., 2009). action of free-ranging kori bustard in the Serengeti plains, Fifteen years ago, the kori bustard population in South a reliable population estimate is required (Braun, 2005; Africa was estimated to number 2000–5000 birds (Ander- Marques et al., 2013). The aim of this study was to identify son, 2000), although the numbers are thought to be vital environmental factors that influence the occurrence decreasing throughout southern Africa (Senyatso, Collar & and density of kori bustard within four study sites in the Dolman, 2013). Shaw (2013) found that the annual Serengeti grass plains. mortality of kori bustard in the Nama district alone, In this study, we predicted that (i) because we performed due to collision with power lines, was approximately one- our fieldwork in similar habitats, the density of kori seventh of the South African population, that is 720 bustard will be similar within the four study areas with no individuals. In the early 1970s, the kori bustards in Africa seasonal differences, and (ii) kori bustard will be equally were one of the most characteristic elements of the distributed, regardless of differences in grass height and northern Sahelian avifauna, except in the heart of Tenere; colour. however, by 2004, no single bustard was observed (Thiollay, 2006). Increased human population adjacent Materials and methods to protected areas has shown to cause pressure on wildlife through illegal hunting (Herremanns, 1998; Loibooki Study area et al., 2002; Kideghesho, 2006). In Burkina Faso, studies have revealed that the main factors correlated with large This study was conducted in the Serengeti grass plains in bird extinction or decline in protected areas were hunting, an area located on the eastern Serengeti National Park and habitat degradation and fragmentation due to intensive western Ngorongoro Conservation Area grass plains, cattle grazing, wood cutting and farming (Thiollay, 2006). Tanzania, between 1o and 3o30’ S and 34o and 36o E A similar situation was observed in the Serengeti ecosys- (Figure 1). Serengeti National Park (14,763 km2) is char- tem where the kori bustard population is declining due to acterized by highland savannahs dominated by thorny habitat degradation and hunting (Mmassy & Røskaft, woodland trees (, Commiphora, Ficus, Combretum and 2014). Podocarpus) and extensive grass plains (Herlocker, 1976). Behaviour and density are basic aspects of a species’ However, the study sites in the eastern Serengeti and ecology, but remain poorly known for the free-ranging kori western Ngorongoro plains are dominated by grasslands. bustard and little data exist on wild birds (Mwangi, 1988; The climate is warm and dry with mean annual temper- Osborne & Osborne, 1998; Hallager & Boylan, 2004; atures between 15°C and 27°C (Sinclair, 1995).

© 2016 John Wiley & Sons Ltd, Afr. J. Ecol. Kori bustard occurrence in the Serengeti 3

Fig 1 Map of Serengeti National Park showing the Kori bustard study area (indicated in red)

Rainfall on the Serengeti plains is bimodal, and the long Area in which human activities and settlements are rain season occurs between March and May and the short allowed. The Ikoma Open Area allows human settlement rain season between November and January (Norton- and some human activities, including livestock keeping, Griffiths, Herlocker & Pennycuick, 1975). Rainfall is hunting by residents, farming and firewood collection. The generally lower in the south and east (500 mm yearÀ1) neighbouring Ikorongo and Grumeti Game Reserves are of the ecosystem than in the north and west (950– partially protected and only allow trophy hunting. 1150 mm yearÀ1). Vegetation cover in Serengeti National Park is influenced mainly by soil type and rainfall and can Data collection be broadly classed into eastern grass plains, central acacia woodlands and northern broadleaf forests (Sinclair, 1995). The data were collected at four sites including Naabi– The Ngorongoro Conservation Area (8094 km2) spans Ngorongoro–Ndutu South (south-west, SW), Naabi– vast expanses of highland plains, savannah, savannah Ngorongoro–Ndutu North (north-west, NW), Maasai– woodlands and forests, from the plains of the Serengeti Barafu–Golkopjes North (north-east, NE) and Maasai– National Park in the north-west, to the eastern arm of the Barafu–Golkopjes South (south-east, SE) (Figure 1). The Great Rift Valley. The southern and eastern slopes of the study areas are predominantly treeless and dominated by highlands receive high rainfall, but the western plains lie in short grass with a maximum height of 30 cm, along with the mountains’ rains shadow and receive less precipitation. some small trees and shrubs in a few kopje areas. Kori In the Serengeti National Park, human activities are bustard observations were recorded on a monthly basis prohibited and contrast that of Ngorongoro Conservation from January 2014 to June 2015. The study was divided

© 2016 John Wiley & Sons Ltd, Afr. J. Ecol. 4 Emmanuel C. Mmassy et al. into four seasons: (i) short dry season (January – March), variance was used to compare the mean kori bustard (ii) long rain season (April – June), (iii) long dry period density in relation to grass colour, grass height and season (July – September) and 4) short rain season (October – in the four study areas. Chi square (v²) tests were used to December). The study sites were selected because of compare variations of kori bustard observation with previous anecdotal observations of large numbers of kori season and habitat, that is grass height and colour. Linear bustards during specific periods. regression analyses were used to relate the density of kori We collected data by driving a vehicle slowly (20 km per bustard to the four study sites, grass height and colour at h) in randomly selected 1-km-long transects (n = 1897). different seasons. All birds within 100 m of each side of the observation vehicle were counted; thus, each transect covered 0.2 km². Results When a 1-km transect was completed, another 1 km was driven without making any recordings. All kori bustards The kori bustards were regularly seen in the same area, observed within each transect line were recorded, includ- and most observations were of single birds (70.2%). The ing those that flew away but were within transects at the mean density of kori bustard in the Serengeti grass plains start of the observation. Kori observations within transects was 0.25 individuals per 0.2 km² (Æ1.01, SD, n = 1897, were performed by visual scanning of the immediate not log-transformed data). The density was 0.12 individ- surroundings, and Zeiss binoculars (10x40, field 7.10) ual/0.2 km² (Æ0.48, SD, n = 314) in the NE area, 0.24 when required, while a Leica CRF 900 (200 m) range- individual/0.2 km² (Æ0.86 SD, n = 264) in the SW area, finder was used to determine the distance from the vehicle. 0.28 individual/0.2 km² (Æ1.28 SD, n = 514) in the SE The censuses were conducted in all four study sections area and 0.28 individual/0.2 km² (Æ1.01 SD, n = 805) in (NW, SW, NE and SE). A minimum of 25 transects were the NW area. There was no statistically significant conducted daily for four days, starting at the beginning of difference in kori bustard densities among the four study every month and including a minimum of 100 transects areas (log-transformed data: ANOVA; F = 2.423, df = 3 per month. Kori bustards were also recorded outside the and 1893, P = 0.064). transects, but these observations were not included in the Although the most common colour of the grass was analysis. Grass height was initially measured by tape green when it was longer than 10 cm, a significant measure (100 cm) followed by visual estimates of grass difference in the frequencies of green colour existed in height over the entire area of observation. different grass height groups (v²=165.5, df = 4, P < 0.001; Table 1). A significant difference in kori bustard density existed in transects with different grass Data analyses heights (log-transformed data: ANOVA; F = 7.527, df = 2 Because kori bustard density data were not normally and 1894, P < 0.001), with the highest densities in short distributed, we log-transformed the data to make them grass habitats ranging from 11 to 30 cm. Statistically normally distributed. Before log transformation, we added significant variations in densities also occurred with grass the integer 1 to all observations to avoid transformation of colour (log-transformed data: ANOVA; F = 5.319, df = 2 zeros. Therefore, all statistical tests are conducted with log- and 1896, P = 0.005). transformed data. We determined the density of kori bustards in relation to the following variables: grass colour (green = 100% green, Table 1 Relation between grass colour and height where kori greenish ˃50% green, brown = 0% green); height of grass bustards were observed was described as very short (less than 10 cm), medium height (11–30 cm) and tall (>30 cm); season and study Grass height Number of site (SW, SE, NW and NE). These four areas were the result Grass colour 0-10 cm 11-30 cm >30 cm observations of dividing the study area of Figure 1 into four equal sites. Green 238 346 4 588 We calculated the total density of kori bustard per Greenish 244 158 11 413 ² 0.2 km (i.e. all koris observed on both sides of the vehicle Brown 649 242 5 896 during the 1-km transect) and compared the mean density N total 1131 746 20 1897 among the study sites. Furthermore, an analysis of

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Kori bustard densities differed highly significantly kori bustards have sufficient species-specific resources. A among the four different seasons (log-transformed data: high kori bustard density in green habitats might be ANOVA; F = 12.382, df = 3 and 1893, P < 0.001). The influenced by an abundant food supply, such as insects. density was highest during the long rain season (April– The presence of large herds of wild animals in green June: 0.33 individual/0.2 km² Æ1.40 SD, n = 610) and habitats may be attractive to kori bustards because they the short dry season (January–March: 0.30 individual/ may offer protection from predators, ultimately increasing 0.2 km² Æ0.92 SD, n = 677), while the density was the the time available to individual birds for finding and lowest during the long dry season (July–September: 0.01 consuming food (Roberts, 1996; Fernandez-Juricic, Siller & individual/0.2 km² Æ0.12 SD, n = 208) and the short rain Kacelnik, 2004). Thus, the combination of food and offer season (October–December: 0.14 individual/0.2 km², of protection might influence kori bustard to congregate in Æ0.58, SD, n = 402). these green habitats. The presence of fresh animal drop- A linear regression analysis with kori bustard density pings may also attract beetles and other invertebrates, (log-transformed) as the dependent variable and area, which are a source of food for the bustard. season, grass height and grass colour as independent Kori bustard density was high during the short dry season variables was significant (r2 = 0.017; ANOVA F = 7.989, (January–March) and the long rain season (April–June). df = 4 and 1892, P < 0.001). However, only season These high densities may be related to breeding season (B = À0.013, t = À3.645, P < 0.001) and area preparations as the male kori bustard courtship display was (B = 0.006, t = 1.961, P = 0.050) were significant con- observed from late October throughout the breeding season tributors in explaining the density variation. Grass length during field data collection. The declining number of kori (P = 0.409) and grass colour (P = 0.220) did not con- bustards at the study site after the breeding season indicated tribute significantly in explaining this variation. that the species migrate to more favourable habitats, for example with respect to food availability, during the long dry season (June–September) and migrate back to the plains Discussion before the rainy season. The results indicate that kori bustard density in the Although not significantly different, the greater number Serengeti grass plains is approximately 0.25 individual per of kori bustard observations in NW and SE, as well as in high 0.2 km2, or approximately 1.3 individuals per km2. The grass habitats, that is medium height (11–30 cm), could be mean density did not differ among the four different study related to the kori bustard predator avoidance behaviour sites. According to studies by Frith (1973) and Isakov from sites with grass that is too long or too short. Given that (1974) in Australia and , intensive land-use practices a different habitat type varies strongly with respect to have been part of Ardeotis australis’ decline in parts of composition and spatial distribution of food resources, the Australia, as well as the decline of Otis tarda in Europe. The habitats used by kori bustards are expected to have sufficient few individuals observed in the Serengeti grass plains may resources for the individual’s daily demands. be connected to intensive land-use practices in adjacent communities, as well as hunting (Mmassy & Røskaft, Conclusions and recommendations 2014). We must emphasize, however, that there is no previous information available about kori bustard densities In conclusion, kori bustard density varied between the four in the Serengeti grass plains or other areas in Tanzania study sites, but was statistically insignificant; however, with which to compare. Thus, repeating this study will be seasonal variations in density were significantly different. necessary after a period of 5–10 years. The density was high during the long rain and short dry Kori bustard density was highest in green grass habitats seasons. The study also confirmed that the distribution of with medium height (11–30 cm), probably because of food kori bustard in the study sites was influenced more by availability and topographic advantages for vigilance grass height (11–30 cm) than by grass colour. Our against predators. Habitat structure is considered to be predictions corresponded better with the season represent- important in determining habitat use by birds (Block & ing the main contributing factor for kori bustard density. Brennan, 1993). Given that different habitat types vary Due to a lack of data and research, kori bustard density with respect to composition and spatial distribution of food recorded in our study in the Serengeti grass plains cannot resources, it could be expected that the habitats used by be categorized as low or high. The species is listed as near

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threatened by the IUCN, and it is consequently important Balmford, A., Green, R.E. & Jenkins, M. (2003) Measuring the that the management authorities take appropriate actions changing state of nature. Trends Ecol. Evol. 18, 326–330. to ensure the survival of the species. Protection of suitable Birdlife-International (2013) Ardeotis kori. IUCN. Block, W.M. & Brennan, L.A. (1993) The habitat concept in habitats of the species within the Serengeti Ecosystem ornithology: theory and applications. Curr. Ornithol. 11,35–91. together with public education is imperative. The increas- Braun, C.E. (2005) Techniques for Wildlife Investigations and ing illegal exploitation of natural resources by local people Management, 6th edn. The Wildlife Society, Bethesda, MD. from communities alongside the Serengeti National Park Ceballos, G. & Ehrlich, P.R. (2002) population losses and makes community participation in the conservation and the extinction crisis. Science 296, 904–907. management of the species important. The large seasonal Clark, R.G. & Shutler, D. (1999) Avian habitat selection: variations in density indicate that the population may pattern from process in nest-site use by ducks? Ecology 80, – utilize large parts of the ecosystem, thereby increasing the 272 287. Collar, N. J.(1996) Otididae (Bustards). In: Handbook of Birds of the likelihood of encountering detrimental anthropogenic World. Hoatzin to auks (Eds J. DEL HOYO,A.ELLIOTT and J. activities. Based on an earlier study of illegal offtake of SARGATAl). Lynx Editions, Barcelona. grassland birds and eggs for home consumption (Magige Dale, J. (1990) The Kori bustard in central Zimbabwe. Honeyguide et al., 2009), we advise protected area managers to address 36, 123–128. such threats posed by illegal offtake. Fernandez-Juricic, E., Siller, S. & Kacelnik, A. (2004) Flock density, social foraging, and scanning: an experiment with starlings. Behav. Ecol. 15, 371–379. Acknowledgements Frith, H. J. (1973) Wildlife Conservation. Angus and Robertson, This project was supported by the Intergovernmental Sydney. Gaston, K.J., Blackburn, T.M. & Goldewijk, K.K. (2003) Habitat Policy Platform on Biodiversity and Ecosystem Services conversion and global avian biodiversity loss. Proc. R. Soc. Lond. (IPBES) grant from the Ministry of Foreign Affairs through Ser. B-Biol. Sci. 270, 1293–1300. the Norwegian Environment Agency to the Norwegian Gottschalk, T.K., Ekschmitt, K. & Bairlein, F. (2007) Relationship University of Science and Technology (NTNU) and between vegetation and bird community composition in TAWIRI. This study was further supported by a grant grasslands of the Serengeti. Afr. J. Ecol. 45, 557–565. from the kori bustard Species Survival Plan (SSP) and HALLAGER,S.&BOYLAN, J. (eds.) (2004) Kori Bustard Species Survival Jacksonville Zoo in the USA for financial support. In Plan (Ardeotis kori) Husbandry Manual. National Zoological Park, particular, we thank Sara Hallager, Smithsonian National Washington, DC. Hallager, S. & Lichtenberg, E.M. (2007) New display behavior in Zoological Park and chair of the kori bustard SSP, and male Kori Bustard (Ardeotis kori struthiunculus). Wilson J. Katie Bagley, and vice chair of the SSP, for Ornithol. 119, 750–755. providing funding for this study through the SSP. We also Harcourt, A.H. & Park, S.A. (2003) Threatened primates thank TAWIRI, NCAA and TANAPA for allowing us to experience high human densities: adding an index of threat conduct kori bustard research in the Ngorongoro Conser- to the IUCN Red list criteria. Biol. Conserv. 109, vation Area and Serengeti National Park. Furthermore, we 137–149. Herlocker, D. thank the Serengeti Wildlife Research Centre (SWRC) for (1976) Woody Vegetation of the Serengeti National Park. Texas A&M University, College Station, Texas. logistical support and Juma Mkwizu (driver) and the entire Herremanns, M. 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