Olwoch_45-72.indd INTRODUCTION The realization that species distributions are signifi- cantly influenced by climate has placed considerable emphasis on the need to acquire information con- cerning the ways in which the present distributions of organisms will be affected by climate change. In response to this challenge, the issue of vector dis tri- bution has recently received much attention. Nu mer- ous studies (Sutherst & Maywald 1985; Nix 1986; Perry, Lessard, Norval, Kundert & Kruska 1990; Nor- val, Perry & Young 1992; Rogers & Randolph 1993; Sutherst, Maywald & Skarratt 1995; Randolph & Rogers 1997; Estrada-Peña 1999; Rogers & Ran- dolph 2000; Randolph 2001, 2002; Erasmus, Ksha- triya, Mansell, Chown & Van Jaarsveld 2000; Eras- mus, Van Jaarsveld, Chown, Kshatriya & Wessels 2002; Olwoch, Rautenbach, Erasmus, Engelbrecht & Van Jaarsveld 2003; Van Staden, Erasmus, Roux, Wingfield & Van Jaarsveld 2004; Thomas, Cameron, Green, Bakkenes, Beaumont, Collingham, Erasmus, Ferrierra, Grainger, Hannah, Hughes, Huntley, Van Jaarsveld, Midgley, Miles, Ortega-Huerta, Peterson, 45 Onderstepoort Journal of Veterinary Research, 74:45–72 (2007) Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa J.M. OLWOCH1*, A.S. VAN JAARSVELD2, C.H. SCHOLTZ3 and I.G. HORAK4 ABSTRACT OLWOCH, J.M., VAN JAARSVELD, A.S., SCHOLTZ, C.H. & HORAK, I.G. 2007. Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa. Onderstepoort Journal of Veterinary Research, 74:45–72 The suitability of present and future climates for 30 Rhipicephalus species in Africa are predicted us- ing a simple climate envelope model as well as a Division of Atmospheric Research Limited-Area Model (DARLAM). DARLAM’s predictions are compared with the mean outcome from two global cir- culation models. East Africa and South Africa are considered the most vulnerable regions on the continent to climate-induced changes in tick distributions and tick-borne diseases. More than 50 % of the species examined show potential range expansion and more than 70 % of this range expansion is found in economically important tick species. More than 20 % of the species experienced range shifts of between 50 and 100 %. There is also an increase in tick species richness in the south-western re- gions of the sub-continent. Actual range alterations due to climate change may be even greater since factors like land degradation and human population increase have not been included in this modelling process. However, these predictions are also subject to the effect that climate change may have on the hosts of the ticks, particularly those that favour a restricted range of hosts. Where possible, the anticipated biological implications of the predicted changes are explored. Keywords: Climate change, Rhipicephalus species, sub-Saharan Africa, tick-borne disease * Author to whom correspondence is to be directed. E-mail: jane.olwoch@up.ac.za 1 Department of Geography, Geo-informatics and Meteorology, University of Pretoria, Pretoria, 0002 South Africa. 2 CIB, Department of Botany and Zoology, Stellenbosch Uni- versity, Stellenbosch, 7160 South Africa 3 Scarab Research Group, Department of Zoology and Entomol- ogy, University of Pretoria, Pretoria, 0002 South Africa 4 Department of Veterinary Tropical Diseases, Faculty of Vet- erinary Science, University of Pretoria, Onderstepoort, 0110 South Africa and Department of Zoology and Entomology, University of the Free State, Bloemfontein, 9300 South Africa Accepted for publication 28 September 2006—Editor 46 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa Philips & Williams 2004) have attempted to predict the distribution of species based on the major envi- ronmental factors that would influence this. This approach neither disregards the need for fur- ther detailed and comprehensive eco-physiological studies nor does it pretend to predict the future. What it does is define the role of climate as a factor in determining the potential for future vector estab- lishment when all other elements are excluded (Suth- erst 2003). This paper also accents the necessity of acquiring more detailed information concerning the biology and environmental sensitivities of each spe- cies in the light of likely climate change. In the ab- sence of such information, relatively straightforward statistical methods that seek correlations between environmental factors and the presence of animals or plants are likely to remain the best pragmatic ap- proach for exploring the expected future distribu- tions of large numbers of species. This study focuses on climate induced changes likely to occur in the dis tribution of some species of an economically im- portant African arthropod, namely the tick genus Rhipi cephalus. The intimate relationship between climate and the requirements of ticks for survival is well documented (Tukahirwa 1976; Rechav 1981, 1982; Short & Nor- val 1981; Minshull & Norval 1982; Norval, Walker & Colborne 1982; Dipeolu 1989; Perry et al. 1990; Nor- val et al. 1992; Okello-Onen, Tukahirwa, Perry, Row- lands, Nagda, Musisi, Heinonen, Mwayi & Opuda- Asibo 1999). This has led to several studies using climate as a means of predicting the distributions of African ticks (Rogers & Randolph 1993; Randolph 1993, 2001; Randolph & Rogers 1997; Norval, Suth- erst, Kurki, Kerr & Gibson 1997; Cumming 2000a, b; Estrada-Peña 2001; Olwoch et al. 2003). Collec- tively the results obtained from these works, some of which used data garnered mainly from interpo- lated climate databases at 25 km resolution (Perry et al. 1990) or 6 x 6 km resolutions (Cumming 2000b), suggest that accurate predictions of tick distributions at different temporal and spatial scales should be feasible. This approach would be especially useful for predicting the distribution of species in poorly sam- pled species in poorly sampled regions of Africa. The genus Rhipicephalus is the fourth largest in the Family Ixodidae (Walker, Keirans & Horak 2000), and there are 74 species currently recognized. It is essentially an African genus with approximately 63 species recorded only in the Afro-tropical region and ten species outside the region. One species, Rhipi- cephalus evertsi evertsi Dönitz, 1910, whose distri- bution was originally confined to the Afro-tropical region has now gained a foothold on the Arabian Peninsula and its distribution is expected to spread even further (Walker et al. 2000). Only 30 species are included in this paper. They are those whose ecological, life history strategies and climatic require- ments are relatively well known and, as two distribu- tion data sets have been used, the current distribu- tion of these ticks are relatively well plotted. The way in which these distributions will be influenced by cli- mate change is poorly documented and forms the essence of this paper. The use of climate-matching models to predict tick distributions One of the earliest climate matching approaches was CLIMEX, which calculates the climatic suitability of geographic regions for species using a tempera- ture-dependent growth index moderated by four growth indices: hot, cold, dry and wet (Sutherst & Maywald 1985). The use of CLIMEX in northern Australia was considered a great success for pre- dicting the distribution of the tick Boophilus micro- plus (Canestrini, 1887), and it was anticipated that this initial success could be translated into predict- ing distributions of African tick species. However, early studies tended to over-estimate the distribu- tion of B. microplus across Africa, and the predicted high incidence of Rhipicephalus appendiculatus Neu- mann, 1901 in West Africa was at complete variance with the tick’s absence in this region. Moreover, when the distributions of Amblyomma spp. in Africa were modelled using CLIMEX, these were found to be dissimilar to their known distribu- tions. These conflicting results led Norval, Perry, Melt zer, Kruska & Boothroyd (1994) to conclude that the predicted climatic suitability of regions for Amblyomma hebraeum Koch and Amblyomma vari- egatum Fabricius, 1794, is almost the opposite of the actual distribution of these ticks, both in Zim ba b- we and in the rest of Africa (Norval, Perry, Gebreab & Lessard 1991; Norval et al. 1992). BIOCLIM was the second climate-based approach employed to model tick distributions (Nix 1986; Norval et al. 1992). BIOCLIM generates 24 climatic attributes from which annual and seasonal mean conditions, extreme values and intra-year seasonal- ity are derived, for each of a selection of geographic points throughout the distribution range of a tick species. Computer-selected thresholds and limits for each of the indices are matched across a geo- graphical grid to predict potential species distribu- tion. This model generally provided a better fit be- tween the predicted and known distributions of R. 47 J.M. OLWOCH et al. appendiculatus, although at a finer scale the match in some areas of the East African highlands was unsatisfactory (Norval et al. 1992). The climate data- base used was interpolated at an increased resolu- tion (8 km), and this factor alone may explain the improved accuracy when compared to the earlier CLIMEX-based attempts. A subsequent logistic re- gression approach (Cumming 2000c), based on inter polated climate and elevation data for Africa with a resolution of 25 km (Hutchinson, Nix, Mac- Mahon & Ord 1996) achieved even better accuracy. Such an approach, however, normally requires the existence of a training data set that includes pres- ence and absence information (Estrada-Peña 2003). While it is relatively easy to ascertain where ticks have been collected, it is more difficult to confirm the reliability of surveys in which a tick species is cited as not present. Consequently the assumption that non-presence always implies absence may limit the application of this modelling approach (Estrada- Peña 2003). The use of an Advanced Very High-Resolution Radi- o meter (AVHRR) mounted on the National Oceanic and Atmospheric Administration’s (NOAA’s) meteo- rological satellites was given preference in the 1990s. This instrument allowed the direct detection of envi- ronmental factors at an 8 km resolution (Lessard et al. 1990). The main predictor in this procedure is the satellite-derived maximum mean monthly Normal- ized Difference Vegetation Index (NDVI). However, this technique was reportedly very complicated when used to predict the distribution of R. appendiculatus (Kruska & Perry 1991). There are, however, initia- tives to revive confidence in the NDVI approach as a predictive tool in research (Randolph 2002). The present study used a single species distribution modelling procedure (Erasmus et al. 2000), original- ly developed by Jeffree & Jeffree (1994, 1996), for predicting species distribution patterns and for eval- uating the relative performance of predicted future climate data sets. This model was subsequently modified to accept multivariate inputs to yield prob- ability of presence maps for species (Erasmus et al. 2000). When used to predict the contemporary po- tential distribution of African ticks (Olwoch et al. 2003) the model achieved positive predictions of more than 70 % for the four tick species tested. Climate data used for predicting African tick distributions The principal sources of climate data for predictive distribution modelling are climate surfaces, gener- ated by interpolating data sampled at varying inten- sities across a region. Consequently, differences between these climate surfaces can usually be attrib- uted to spatial and temporal evenness of the data used for interpolation. Most modern interpolation techniques are pattern based and statistically incor- porate horizontal as well as vertical (altitudinal) ad- justments (Hutchinson 1989, 1991; Hutchinson & Gessler 1994). These climate surfaces are, how ever, relatively smooth because of extensive interpolation between low-resolution point observations. Another source of climate data is Global Circulation Models (GCMs). These are coupled ocean-atmos- phere models that provide three-dimensional simu- lations of the atmosphere. To date GCMs have pro- duced climate data at a horizontal resolution that is generally too coarse for use in predictive species modelling (> 100 x 100 km grid point resolution), especially for species that are habitat specialists or that are influenced by fine-scale environmental gra- dients. Computational requirements usually preclude GCMs being run at meso-scale grid resolutions (10–100 km). The present study used a Division of Atmospheric Research Limited Area Model (DARLAM) as the main climate data set. DARLAM is a potential alter- native source of high-resolution climate data that involves the nesting of a high-resolution limited area model within a GCM over the area of interest (for review see McGregor 1997). The GCM supplies the limited area model with initial and boundary condi- tions. With a grid resolution of 10–100 km, the lim- ited area model is able to simulate some of the meso-scale properties of the circulation model. This technique provides a viable fine scale alternative to the use of observed or interpolated climate surfaces or very coarse scale GCMs climate surfaces. The resolution attained by this dynamic modelling pro- cess is essentially limited by the computing power available to the modellers. The implication of using these datasets for predicting current tick distribu- tions has been assessed by Olwoch et al. (2003). In this study, DARLAM’s future predictions are com- pared with those obtained by using mean climates from two GCMs (CGCM and Centre for Climate Sys- tem Research/National Institute for Environ mental Studies [CCSR/NIES]). MATERIALS AND METHODS Study area The study area covers sub-Saharan Africa (Fig. 1) and was divided into 3 000 grids cells of 60 x 60 km 48 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa resolution. This resolution was determined by the DARLAM climate data. Tick data Point localities of tick recoveries were obtained from Cumming (1999), who compiled the data that he used from various collections of ticks, and from re- cent collections made by one of us (I.G.H.). Com- bining data sets from different sources frequently compounds identification and distribution errors and for this reason data congruence with Walker et al. (2000), who provide well-illustrated distributions of Rhipicephalus species, was used to assess data quality in the final compiled dataset. Synonyms pro- vided by the latter authors also solve the common dataset problem of referring to one species, but us- ing different names, or referring to a group of spe- cies as a single species. The tick species selected for this study belong to the ixodid genus Rhipi ceph- alus. Species point localities were assigned to partic- ular 60 km x 60 km grid cells by means of a spatial intersect in ArcView GIS. A conservative estimate of the accuracy of these point localities is 0.2 ° (G.S. Cumming, personal communication 2003) and con- sequently this approximation is considered reason- able. The Rhipicephalus species selected are those whose distribution and life history strategies are rel- a tively well known and it is our hope that these re- sults will provide a baseline model for future model- ling of other tick species. Predictive species modelling A simple climate envelope model was used to pre- dict the future distribution of the focal species (Eras- mus et al. 2000, 2002). The input data comprised 3 000 grid cells of 60 x 60 km size populated with climate variables covering sub-Saharan Africa. Reli- able presence records of the selected tick species and the present climate values at these locations were used to construct a climate envelope, using a principal components-type approach. This climate envelope represents the range of climates within which a particular tick is known to occur, and can be interpreted as the realized niche, as defined solely by climate. To arrive at a predicted distribution in a climate change scenario, the existing climate enve- lope is applied to a climate surface representing fu- ture climates, and a new geographical interpretation of distribution is derived [see Erasmus et al. (2000, 2002) and Olwoch et al. (2003) for a more detailed explanation]. This approach was used as a stand- ardized base for evaluating the relative perform- ances of the DARLAM and the mean GCM climate data sets, and allows the creation of a probability surface of climate suitability for each species mod- elled. FIG. 1 Study area National boundaries km 49 J.M. OLWOCH et al. DARLAM present and future climate data The Australian Commonwealth Scientific and Indus- trial Research Organisation (CSIRO) developed the high-resolution limited-area model DARLAM for use in both short-term meso-scale atmospheric studies and longer-term climate simulation experiments (Walsh & McGregor 1995). In the present study ten separate 30-day simulations were performed for both January and July for separate 10-year periods. The periods selected are the 1990s and 2020. The sim- ulations were performed at a horizontal grid resolu- tion of 60 x 60 km using a domain of 100 x 100 grid points that cover sub-Saharan Africa. The month ly average of the ten simulations constitutes the model climatology for the month. The CSIRO Mark 2 GCM was used to force DARLAM at its lateral bounda- ries. The GCM was integrated for the period 1960– 2100, with greenhouse gas forcing corresponding to the A2 SRES (Special Report on Emission Sce- nario, issued by the Intergovernmental Panel on Climate Change) scenario. Engelbrecht, Rautenbach, MacGregor & Katzfey (2002) illustrated that DARLAM is capable of simu- lating the regional characteristics of atmospheric variables such as near-surface temperature, low- level wind patterns and rainfall over sub-Saharan Africa with considerable detail. The model does, however, tend to overestimate total rainfall over re- gions with a steep topography. The DARLAM simula- tions were performed at the Laboratory for Research in Atmospheric Modelling (LRAM) at the University of Pretoria on a Pentium III workstation with two 550 MHz processors. GCM future climate data The GCM climate data used in this study were downloaded from the IPCC/DDC website. Through various stages in ArcView GIS, the original GCM data were processed to fit the 60 x 60 km resolution of DARLAM. The Canadian Global Coupled Model (CGCM2) was the first GCM climate used in this study. It is based on the earlier CGCM1, but with some improvements aimed at addressing shortcom- ings identified in the first version. In particular, the ocean mixing parameterization has been changed (Gent & McWilliams 1990), and following Flato & Hibler (1992) sea-ice dynamics has been included. The version of GCM2 used for control and doubled CO2 experiments has ten vertical levels with the low- est prognostic level located at approximately 200 m above the surface. A description of CGCM2 and a comparison, relative to CGCM1, of its response to increasing greenhouse-gas forcing can be found in Flato & Boer (2001). The climate change projec- tions used in this study are those from the newer IPCC SRES A2 scenario. The second GCM model used was developed by the Center for Climate System Research/National Institute for Environmental Studies, (Japan) (CCSR/ NIES) CGCM (Nozawa, Emori, Numaguti, Tsushima, Takemura, Nakajima, Abe-Ouchi & Kimoto 2001). This model is also based on Emission Scenarios (SRES) of the Intergovernmental Panel on Climate Change (IPCC). It is a Transient Coupled Ocean- Atmosphere Model, which was developed to inves- tigate the direct and indirect climate impacts of the anthropogenic sulphate and carbonaceous aerosols in future projections of climate change. The data used here are from the A2 scenarios. Direct radia- tive forcing of the carbonaceous aerosols nearly nul- lifies that of the sulphate aerosols for all scenarios. Estimated total indirect radiative forcing is about –1.3 Wm-2 for the A1, B1, and B2 scenarios, and is about –2.0 Wm-2 for the A2 scenario in the latter half of the 21st Century. Global and annual averages of the surface air temperature increase for all sce- narios because of the dominance of the radiative forcing of the increased CO2. Global warming is de- celerated with an increase in the anthropogenic sul- phate and carbonaceous aerosols, because indirect forcing due to the aerosols has a significant cooling effect. Geographical distribution of the surface warm- ing does not depend much on the scenarios. Cloud feedback becomes dominant in the latter half of the 21st Century, and this introduces further warm ing at the surface. Predicting current and future distribution of Rhipicephalus species The predicted current distributions were initially ob- tained using current climate predicted by DARLAM. This represents a useful comparison between pre- dicted distributions and known records (see Olwoch et al. 2003). To obtain predicted future distributions the grid cells are populated with future climate vari- ables. The predicted current distributions were ob- tained using the predictive species model (Erasmus et al. 2000) and six climate variables of current and future mean maximum temperature, mean minimum temperature and mean rainfall of January and July provided by DARLAM. The predicted future distribu- tions were obtained by using both DARLAM and the mean GCM climates. The predictive modelling was executed in S-Plus (S-Plus 2000), while maps of the results were drawn in ArcView GIS. The resultant potential distribution maps represent the probability values of their suitability for ticks based on climate. 50 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa Analysis of predicted tick range changes A number of analyses were performed to compare the predicted current and future distributions of ticks. These included: (i) Analysis of species range expansion (ii) An analysis of range contraction (iii) Change in species richness pattern (iv) Species range shifts (v) An assessment of overlap between DARLAM and GCM predicted future distributions. These range changes were initially analysed for the whole study area and subsequently, in some cases, on a regional basis. In the second analysis, ticks were grouped into the following regions depend ing on their principal regional distribution, namely East Africa, Central Africa and southern Africa, and a fourth group of ticks that were termed “general” ticks. The East African tick species include Rhipicephalus aquatilis Walker, Keirans & Pegram, 1993, Rhipi ceph- alus armatus Pocock, 1900, Rhipicephalus bequaer ti Zumpt, 1949, Rhipicephalus carnivoralis Walker, 1966, Rhipicephalus humeralis Rondelli, 1926, Rhipi- cephalus kochi Dönitz, 1905, Rhipicephalus ma - culatus Neumann, 1901, Rhipicephalus muehlensi Zumpt, 1943, Rhipicephalus planus Neumann, 1907 and Rhipicephalus pulchellus Gerstäcker, 1873. The Central African species include Rhipicephalus complanatus Neumann, 1911, Rhipicephalus com- positus Neumann, 1897, Rhipicephalus dux Dönitz, 1910, Rhipicephalus longus Neumann, 1907, Rhipi- cephalus lunulatus Neumann, 1907, Rhipicephalus masseyi Nuttall & Warburton, 1908, Rhipicephalus punctatus Warburton, 1912, Rhipicephalus senega- lensis Koch, 1844, Rhipicephalus supertritus Neu- mann, 1907 and Rhipicephalus ziemanni Neumann, 1904. The southern African species include ticks of the Rhipicephalus capensis group (Rhipicephalus cap- ensis Koch, 1844; Rhipicephalus follis Dönitz, 1910 and Rhipicephalus gertrudae Feldman-Muhsam, 1960), Rhipicephalus distinctus Bedford, 1932, Rhipi- cephalus exopthalmos Keirans & Walker, 1993, Rhipicephalus oculatus Neumann, 1901, Rhipi ceph- alus zambeziensis Walker, Norval & Corwin, 1981 and the subspecies Rhipicephalus evertsi mimeti- cus Dönitz, 1910. There are also species that have wide ranging dis- tributions that overlap in various regions of Africa. These species, termed “general” ticks include R. appendiculatus, R. evertsi evertsi Neumann, 1897, Rhipicephalus pravus Dönitz, 1910 and Rhipi ceph- alus simus Koch, 1844. The above groupings are presented to facilitate interpretation of the current findings and do not imply that the ticks placed in particular geographical regions are restricted to these areas, but rather localise their distribution with extensions into neighbouring regions. Range expansion and contraction In order to obtain range changes in terms of contrac- tions or expansions, predicted current or future dis- tributions were first obtained. The predicted current or future distributions were taken as the number of grid cells in which the probability of occurrence is equal to or greater than 50 %. The difference in the number of grid cells between the predicted present distribution (DP) and predicted future distribution (DF) constitutes distribution range change (DC). These range changes may either represent contrac- tions or expansions. We initially performed this anal- ysis on a sub-Saharan scale and later on a regional scale in order to establish which regions in Africa would experience greater changes in predicted tick distribution ranges (current and future) and there- fore appear more vulnerable to climate change. We analysed the differences between the predicted dis- tributions using the Kolmogorov-Smirnoff two-sam- ple test. Furthermore, we divided the ticks into eco- nomically important and unimportant species. A com parison of range changes between the current and future predictions was performed on this latter grouping to assess which of the two groups is more vulnerable to climate change. In all the above analyses we assessed the propor- tion of species that experienced expanded or con- tracted range changes and the degree of the pre- dicted expansion/contraction. Analysis of change in species richness pattern and degree of range shifts Species richness patterns were calculated as the number of species in the predicted current or future distribution per grid cell following Erasmus et al. (2002). This analysis was performed for the whole of sub-Saharan Africa. Range shifts were calculated as the number of additional grid cells in the predict- ed future distribution as a proportion of the current predicted distribution. We used the current predict- ed distribution instead of current known records be- cause most regions in Africa are poorly sampled. 51 J.M. OLWOCH et al. Comparing predicted future distributions of ticks based on climates simulated by DARLAM and GCM The accuracy of any climate model is as good as the initial conditions that are used to configure it. Since there is no climate model that provides an ac- curate projection of the future, it seemed prudent to use the results from more than one climate model in this study. A comparison was therefore made to as- sess the differences between the predicted future climate suitability for tick species using a regional climate provided by DARLAM and a mean of two GCMs described above. The analysis was performed on a sub-Saharan scale and also on a regional scale. We assessed the degree of proportional overlap be- tween the predicted current distribution and the pre- dicted future distribution (DARLAM and GCM) by means of the proportional overlap method (Prender- gast, Quinn, Lawton, Eversham & Gibbons 1993; Reyers, Van Jaarsveld & Krüger 2000). In this case the proportional overlap was calculated as Nc/Ns where Nc is the number of common grid cells be- tween a pair of areas under comparison and Ns is the number of grid cells containing data for both groups or the maximum number of overlapping grid cells possible. RESULTS Model Validation was not performed in this study be- cause the same climate envelope model had previ- ously been subjected to rigorous evaluation using presence-absence data re sulting from a coordinat- ed and systematic survey effort. Erasmus et al. (2002) used the distribution records of 34 bird spe- cies and tested performance of the model using re- ceiver operator characteristic analyses (Fielding & Bell 1997). The model performed sig nificantly better than a random model with no discriminatory ability. It also accurately predicted the complete known dis- tributions for 24 of the 34 bird species, using a 20 % sub-sample of the known rec ords (Erasmus et al. 2002). This satisfactory documented performance of the model and the relatively good predictions that were obtained when it was used to predict the cur- rent distributions of four Afri can ticks (Olwoch et al. 2003) are sufficient reasons to consider the model adequate for the present study. Future climate—DARLAM The climatological anomalies for the 2020s vs the 1990s as predicted by DARLAM are depicted in Fig. 2. January minimum and maximum temperatures are simulated to increase by more than 2 °C over certain regions of sub-Saharan Africa. Many of the eastern regions are expected to become drier with an associated pattern of higher sea-level pressure, whilst the western subcontinent is expected to be- Maximum temperature (°C) Minimum temperature (°C) Mean sea-level pressure (hPa) Rainfall (mm/day) FIG. 2 DARLAM’s climatological anomalies for the 2020s v. 1990s 52 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa come wetter. An interesting feature of the July anom- aly fields is that parts of the central subcontinent are simulated to become cooler and wetter. Species distribution changes Broad scale range changes (Fig. 3A, B) The predicted current and future distributions of the selected Rhipicephalus spp. using DARLAM are provided in figures 10–39. On a sub-Saharan scale, the ranges of 46 % of the tick species, namely R. appendiculatus, R. capensis group, R. distinctus, R. humeralis, R. kochi, R. longus, R. masseyi, R. ocu- latus, R. planus, R. punctatus, R. senegalensis, R. simus, R. zambeziensis and R. ziemanni are pre- dicted to contract. The ranges of 54 % of the spe- cies, namely R. aquatilis, R. armatus, R. bequaerti, R. carnivoralis, R. complanatus, R. compositus, R. dux, R. evertsi evertsi, R. evertsi mimeticus, R. exop- thalmos, R. lunulatus, R. maculatus, R. muehlensi, R. pravus, R. pulchellus and R. supertritus are pre- dicted to expand over the same period (Fig. 3A, B). These results translate into an area expansion of 3 502 800 km2 (12 %) in total tick range with a total reduction of 640 800 km2 (2 %). Central African species (Fig. 4A, B, 16,17, 19, 25, 26, 28, 34, 35 and 39) Fifty-five percent of species in central Africa are pre- dicted to show range reductions (R. longus, R. mas- seyi, R. punctatus, R. senegalensis and R. zie manni) while 45 % (R. complanatus, R. compositus, R. dux and R. lunulatus) are predicted to show range ex- pansions. Although the ranges of the majority of tick species are predicted to contract, the total area of contraction is only 19 %, while the total area of expan- sion by the remaining species is 81 %. The tick spe- cies predicted to expand its range most in this region is R. lunulatus with a total expansion of 252 000 km2. East African species (Fig. 5A, B, 11, 12, 13, 15, 23, 24, 27, 29, 31 and 33) In East Africa 30 % of the species (R. humeralis, R. kochi and R. planus) are predicted to show range contractions, while 70 % (R. aquatilis, R. armatus, R. bequaerti, R. carnivoralis R. maculatus, R. mueh- lensi and R. pulchellus) are predicted to show range expansions. This translates into a total expansion of 1 760 400 km2 (91 %) with a mere 169 200 km2 (9 %) reduction in total area. The predicted greater expan- sions are mainly attributable to R. bequaerti which � �� �� ��� �� � �� � �� � �� ��� �� � �� ��� � � � �� ��� �� ��� � �� ��� �� � �� ��� � � �� �� � � �� �� ��� ��� � �� ��� � � �� ��� � � � �� � � � �� �� � �� � � �� �� ��� �� �� �� � �� ��� �� � � ������� �������� ����� ����� ��������������������� �� ����� ����� ����� ��� ��� ��� ��� � �� ��� �� � ��� �� � � ��� �� �� ��� ��� ��� � � ��� �� �� �� � � ��� �� � � �� ��� � ��� �� �� �� � � ��� �� �� � � � ��� �� � ��� �� �� ��� �� � � ��� �� �� ��� �� � � �� ��� �� �� �� �� � ��� �� ��� � ��� �� �� �� � ��� �� �� �� � ��� ��� �� ��� �� ��� ���� ��� � �� � �� � � �� �� ��� �� � �� � �� � �������� ����� ����� ������������������� �� ����� ����� ����� ��� ��� ��� ��� � ������� ������� ������ � ����� ������� ���������� ����� ����� ������������������� �� ������� ����� ������� ���������� ����� ����� ��������������������� �� ������� �� ��� � � �� �� � � �� �� ��� � � � �� � � � �� �� � �� ��� �� � � � ����� ����� ��� ��� ��� ��� � � �� �� ��� �� � �� � �� � � � �� �� ��� �� � �� � �� � ����� ��� ��� ��� ��� � ��� �� �� �� � � ��� �� �� � � � ��� �� � ��� �� ��� � ������� ������ FIG. 3 Rhipicephalus species in sub-Saharan Africa that are predicted to show (A) range size contraction and (B) range size expansion FIG. 4 Rhipicephalus species in Central Africa that are pre- dicted to show (A) range size contraction and (B) range size expansion 53 J.M. OLWOCH et al. more than doubles its present range and R. pulchel- lus, which is predicted to expand its range by some 921 600 km2 (49 %). Southern African species (Fig. 6A, B, 14, 18, 21, 22, 30 and 38) In southern Africa some 66 % (R. capensis group, R. distinctus, R. oculatus and R. zambeziensis) of the tick species are predicted to contract their rang- es. Although only 33 % (R. evertsi mimeticus and R. exopthalmos) of the ticks are predicted to expand their current ranges, the total range expansion is 439 200 km2 (23 %) while the total range reduction is only 64 800 km2 (1 %). Most of the expansion in this region is attributable to R. evertsi mimeticus. “General” tick species (Fig. 7A, B, 10, 20, 32 and 36) The “general” ticks are those Rhipicephalus species that are widely distributed with current distributions overlapping within various geographical regions of the subcontinent. This does not necessarily mean that these species are not specialists with regard to their ecological requirements, e.g. R. appen diculatus is confined to parts of eastern, central and south- eastern Africa (Walker et al. 2000). It is a species of significant economic importance in Africa because it transmits Theileria parva, the cause of East Coast fever (ECF), which is a major cause of cattle mortal- ity and also causes considerable production losses in cattle in most African countries (Okello-Onen et al. 1999). The only tick species predicted to contract its range in this group is R. appendiculatus, which is predicted to contract its range by 212 400 km2 (5 %). The ranges of 75 % of the ticks in this category, namely R. evertsi evertsi, R. pravus and R. simus are predicted to expand. The total range expansion in this region is equivalent to 864 000 km2 (7 %). This expansion is mainly associated with R. simus, which is predicted to expand its range by 601 200 km2 (70 %). Changes in species richness patterns and range shifts The future climate predicted by DARLAM will alter the species richness distribution pattern of African Rhipicephalus. Compared to the current pattern (Fig. 8A) the predicted pattern is spatially different and broader (Fig. 8B) with encroachment of ticks into new regions. These regions, which include An- FIG. 5 Rhipicephalus species in East Africa that are predicted to show (A) range size contraction and (B) range size expansion FIG. 6 Rhipicephalus species in southern Africa that are pre- dicted to show (A) range size contraction and (B) range size expansion ������� �������� ���������� ����� ����� ��������������������� �� ����� ��� ��� ��� ��� � � �� �� ��� �� � �� � �� � ��� ��� ��� �� ��� ��� �� ��� ��� � � ������� �������� ���������� ����� ����� ������������������� �� ����� ��� ��� ��� ��� � � �� �� ��� �� � �� � �� � ��� ��� ����� ��� ��� �� ��� �� ��� ��� ��� ��� � ��� ��� � � ��� ��� ��� � � ��� �� ��� � � ��� �� ��� � � ������� ������ ������� ���� ������ ���������� ����� ����� ��������������������� �� ��� ��� ��� ��� � � �� �� ��� �� � �� � �� � ���� ��� � ���� � � � � ���� ��� ���� � � ���� ���� � ������� ���� ��� ��� ��� ��� ���� ������ ���������� ����� ����� ������������������� �� ��� ��� ��� � � �� �� ��� �� � �� � �� � � ���� ��� ��� �� ������� ������ 54 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa gola, Namibia, Botswana and the Northern Cape Province of South Africa, are forecast to experience a more than 50 % increase in tick species richness (Fig. 8C). This could be related to increased rainfall in these regions, rendering the south-western re- gions of sub-Saharan Africa more suitable for ticks. The general west-east shift in species ranges re- ported by Erasmus et al. (2002) is not supported in this study on ticks, in which varying degrees of shift in different directions appears to be the emergent pattern for this taxon. Analysis of range shifts fur- ther indicates that 80 % of species show less than a 50 % range shift, while 20 % of species show a range displacement of more than a 100 %. The latter spe- cies are mainly those that populate the western arid regions of Africa (Fig. 8). Species range changes and tick-borne disease implications Thirty percent of the ticks included in this study are classified as economically important because they are vectors of diseases of domestic livestock or other animals (Table 1). Although R. appendiculatus, the principal Rhipicephalus vector of disease in Afri ca shows range contraction, the remaining vectors are responsible for 52 % of the predicted range expan- sion under future climate conditions. The non-vec- tor species are responsible for 48 % of future tick range expansions. Predicted future distributions using DARLAM and GCM climate data The predictions of future climate suitability for ticks when DARLAM is used are generally more exten- sive than those generated when using the combined GCM climates. With the exception of four species (R. compositus, R. evertsi mimeticus, R. exopthal- mos and R. oculatus), and the R. capensis group, DARLAM predicts wider ranges of climatic suitability than the combined GCM climate surfaces. DAR- LAM’s total predictions are 31 % broader than the GCMs. The average difference in the range sizes predicted for DARLAM and GCM is 511 200 km2. Stat is tically there is no significant difference between the predictions by DARLAM and GCM using a Kol- mogorov-Smirnov test (P > 0.1, n = 30). However, when the climate data simulated by DARLAM and GCM are analysed for statistical significance, there FIG. 7 Rhipicephalus species in the “General” ticks that are pre- dicted to show (A) range size contraction and (B) range expansion ������� !"���� �#��������� ����� ����� ������������������� �� ����� ����� ����� ��� ��� ��� ��� � � �� �� ��� �� � �� � �� � �� ��� �� �� �� �� � �� ��� �� � �� � � �� � ������� ������ ������� !"���� �#��������� ����� ����� ��������������������� �� ����� ����� ��� � �� �� ��� �� � �� � �� � ������� ������ � � FIG. 8 Species richness pattern of Rhipicephalus species in sub- Saharan Africa: (A) current; (B) future; and (C) areas with more than 50 % increase 0–4 5–8 9–12 13–16 17–21 National boundaries B A C More than 50 % increase in species richness 55 J.M. OLWOCH et al. TABLE 1 Rhipicephalus spp. and the diseases with which they or the toxins they produce are associated Disease and causal agents Animal affected Vectors East Coast fever (Theileria parva) Cattle R. appendiculatus R. zambeziensis Tick toxicosis Cattle R. appendiculatus Corridor disease or Buffalo disease (Theileria parva) Cattle, buffalo R. appendiculatus R. zambeziensis Gall sickness (Anaplasma marginale) Cattle R. evertsi evertsi R. simus Equine biliary fever or equine piloplasmosis (Babesia caballi, Theileria equi) Horses, mules, donkeys R. evertsi evertsi Spring lamb paralysis Lambs, calves R. evertsi evertsi Canine biliary fever or canine tick fever (Babesia canis), rickettsiosis (Ehlichia canis) Dogs R. sanguineus Paralysis Sheep, lambs, calves R. lunulatus Nairobi sheep disease (Bunyaviridae) Sheep R. pulchellus TABLE 2 Proportional overlaps between pairs of grid cells between the predicted ranges from DARLAM and the ranges predicted us- ing mean values for two GCMs “General” tick species DARLAM 2030 GCM 2030 Common grids Proportional overlap value (%) R. appendiculatus R. evertsi evertsi R. pravus R. simus 969 1 220 1 300 1 209 858 945 868 858 476 611 621 597 55 65 29 60 East Africa R. aquatilis R. armatus R. bequaerti R. carnivoralis R. humeralis R. kochi R. maculatus R. muehlensi R. planus R. pulchellus 33 153 215 205 79 797 425 699 719 841 17 116 40 111 33 491 173 678 413 217 5 29 11 67 12 326 98 315 278 179 29 25 28 60 55 29 56 67 55 25 Central Africa R. complanatus R. compositus R. dux R. longus R. lunulatus R. masseyi R. punctatus R. senegalensis R. ziemanni 626 699 201 1 020 897 450 458 803 611 595 737* 192 990 518 338 440 622 558 400 443 77 642 308 202 262 415 352 67 60 40 25 28 60 28 56 72 South Africa R. capensis group R. distinctus R. evertsi mimeticus R. exopthalmos R. oculatus R. zambeziensis 485 707 423 212 386 299 547* 352 577* 236* 434* 226 309 254 49 116 293 169 56 72 49 15 49 60 Bold* = GCM predictions wider than those of DARLAM 56 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa FIG. 9 Box plots based on the Kolmogorov-Smirnov test that compared climate predicted by DARLAM and a combined GCM ���������� �� �� �� $� $� $� $� $� �� �� �� �� �� �� �� �� � � � �� � $� �% �� �% �� % � &% &�� ���������� �� �� � �� �� � �� $% $� �% �� �% �� % � ���������� �� �� � � � �� � ����� ����� ��� ��� ��� ��� � &��� � � �� �� ���������� �� �� ���������� �� �� $� �� �� �� �� �� �� �� �� �� �� � � � � � �� �� � ���������� �� �� ����� '�� ��� (�� ��� %�� ��� $�� ��� ��� � &��� � � �� �� �%&(%�) *��* �*��� � 57 J.M. OLWOCH et al. FIG. 10 R. appendiculatus FIG. 11 R. aquatilis FIG. 10–39 Predicted probability distribution of Rhipicephalus species using DARLAM and GCM future climates: (A) DARLAM; (B) GCM 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability FIG. 12 R. armatus FIG. 13 R. bequaerti 58 Climate change and Rhipicephalus (Acari: Ixodidae) in Africa FIG. 14 R. capensis group FIG. 15 R. carnivoralis 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability A B 0.748–1.000 0.499–0.748 0.249–0.499 0.000–0.249 National boundaries Probability FIG. 16 R. complanatus FIG. 17 R. compositus FIG. 10–39 Continued