Taxic Richness Patterns and Conservation Evaluation of Madagascan Tiger Beetles (Coleoptera: Cicindelidae)

Home , Andrakata, Ankarana Reserve, Ankazobe (district), Fauna of Madagascar, Mahalevona, Mangerivola Reserve

Taxic richness patterns and conservation evaluation of Madagascan tiger beetles (Coleoptera: Cicindelidae)

Lantoniaina Andriamampianina1,∗, Claire Kremen2, Dick Vane-Wright3, David Lees4 & Vincent Razafimahatratra5,† 1Wildlife Conservation Society, BP 8500 Antananarivo 101, Madagascar 2Center for Conservation Biology and Wildlife Conservation Society, Department of Biological Sciences, Stanford University, Stanford, California 94305, U.S.A. 3Biogeography and Conservation Laboratory, Department of Entomology, 4Department of Palaeontology, The Natural History Museum, Cromwell Road, South Kensington, SW7 5BD, U.K. 5Faculté des Sciences, Université d’Antananarivo, Antananarivo 101, Madagascar ∗Author for correspondence (e-mail: [email protected]; phone/fax: 261-20-22-41174) †Deceased

Received 12 March 1999; accepted 28 February 2000

Key words: biodiversity patterns, species richness, endemism, conservation priority areas, Madagascar

Abstract

Distributional ranges of 17 genera and 172 species of Malagasy tiger beetles (Coleoptera, Cicindelidae) have been compiled to determine patterns of species richness and endemism. These patterns reveal large sampling gaps, and potential priority areas for conservation action. Northern and south-western parts of the island are richer in genera, whereas eastern and especially northern parts of the rainforest show higher species richness, due to extensive radiations within the genera Pogonostoma and Physodeutera. A set of 23 areas are identiﬁed in this study as priority foci for tiger beetle conservation, and six general regions are bioinventory priorities.

Introduction ecosystems), and species identifications were often uncertain. Except possibly for lemurs and birds, avail- The high level of biological diversity and local able data did not reflect the detailed distribution of the endemism in Madagascar reflects not only long iso- taxa (Ganzhorn et al. 1997). Since then, several works lation but the existence of a great diversity of habi- have used smaller, taxonomically better defined groups tats, climatic zones, topography and soil types (Lowry to refine the patterns of biodiversity and to set priority et al. 1997). Recently, the Malagasy government has areas in Madagascar (Emberton 1997; Lees 1997; Lees developed a national environmental action plan that et al. 1999). aims to conserve this unique fauna and flora. One of This paper concerns quantitative biogeography of the most important strategic objectives defined in this Madagascan tiger beetles (Coleoptera: Cicindelidae). plan is the development of an efficient protected areas Cicindelids have been considered to pass a range of network (Banque Mondiale et al. 1988; ONE 1997). test criteria proposed (Pearson & Cassola 1992) for a Knowledge of patterns of biodiversity is essential to promising indicator group: taxonomy stable; biology guide conservation planning. In April 1995, a work- well-known; easy to observe in the field; occurring shop was held in Madagascar that aimed to produce a in a broad range of habitat types; individual species concerted set of national biodiversity priorities. The showing tendency to be specialised within a narrow thematic groups treated during this workshop were, habitat; and finally, preliminary evidence that diversity however, large and vague (e.g. invertebrates, aquatic patterns are similar to those for other taxa. Positive 110 L. Andriamampianina et al. correlations found in the distribution of tiger bee- the taxonomy of Rivalier (1950, 1965, 1967, tles, birds and butterflies in North America, in the 1970). Malagasy tiger beetles are divided into two Indian subcontinent and in Australia at a coarse spa- subfamilies: tial scale (Pearson & Cassola 1992) have for some but – The Collyrinae, represented in Madagascar by not all relationships remained significant after being a unique genus Pogonostoma. This entirely subjected to more rigorous spatial statistics taking endemic genus comprises 81 arboreal species. into account spatial autocorrelation (Carroll & Pearson Generally found in primary forests, they are more 1998; Pearson & Carroll 1998). We describe the aggre- diverse in the evergreen eastern rainforests than gate distribution of all known tiger beetles (family in the dry forests. Adult beetles are found most Cicindelidae) in Madagascar, and consider in partic- of the time on tree trunks, pursuing small inver- ular, species richness patterns at a fine quarter degree tebrates. From time to time, they also chase their resolution and areas of geographical concentration of prey in the surrounding herbaceous strata. Their range-restricted species. In this paper we do not exam- larvae live in holes in the tree trunks. Because of ine the more general question of the relative efficiency their arboreal habit, members of this genus are of tiger beetles in representing other groups of organ- highly threatened by deforestation. isms. However, a previous study (Lees et al. 1999) – The Cicindelinae, comprising 16 other genera of using a nearly identical dataset showed that known tiger beetles with terrestrial habitats. Altogether, ranges of Malagasy cicindelids exhibit a hollow-curve they include 95 species. They are mainly ground range-size frequency distribution skewed to more nar- dwelling, although many of them are found only row ranges, which makes their richness patterns con- in forests. Their larvae construct tubes in the soil. siderably less influenced by the geometric effects of Both adults and larvae prey on small inverte- Madagascar’s boundaries on species richness than for brates. Some of the genera such as Physodeutera, some other groups such as butterflies and vertebrates. Peridexia and Calyptoglossa are common in rain- Although the overall range-size frequency distribution forests, whereas others (Chaetotaxis, Prothyma, of all Madagascar’s biota is unknown, terrestrial verte- Stenocosmia, Waltherhornia) are mainly found brates (about 1000 species) would seem highly unlikely in dry forest and arid places such as prairies, to be representative. In this paper, we evaluate the effi- savannas or the xerophytic formations of the west- ciency of the protected areas network in Madagascar ern, southern and central parts of Madagascar (encompassing 57 quarter degree grid areas) to rep- (Figure 1). Some others (notably Chaetodera, resent known species of Malagasy tiger beetles. We Lophyridia, Habrodera and Lophyra) are found identify a set of priority areas for conservation action, in abundance in sandy places along beaches and other areas where more field research is required. and rivers, mainly in the west and south of Madagascar. Finally, there are those which are Madagascan tiger beetles highly ubiquitous in habitat preference, occurring wherever suitable open substrate exists The Cicindelidae is one of Madagascar’s best-known for breeding (Hipparidium, Ambalia, Cylindera, insect groups. The Malagasy tiger beetle fauna has Cicindelina and Myriochile), such as on roads been the subject of many studies summarised in through towns, in villages, in forests or wood- Andriamampianina (1996). Although most papers are lands on shaded trails, and along rivers. on taxonomy and systematics, extensive distributional data on this group in Madagascar is available in the literature and in museum collections. Methodology The cicindelid fauna of Madagascar (n = 176) is the third richest of any country in the world (Pearson & Data collection Cassola 1992), with more than 99% endemism at the species level. Just one species, Myriochile A species checklist for the family was established melancholica Fabricius, represented in Madagascar by through literature surveys based on recent revisions the endemic subspecies M.m. trilunaris Klug, is also of Pogonostoma by Rivalier (1970), Physodeutera by found in the mainland Africa. Although some species Rivalier (1967), and Jeannel (1946) for all other genera. can be found widely across the island, most have Synonyms were also used in the search for distribu- very localised distributions. In this paper we follow tional data in the old literature and collections. Biogeography and conservation of Madagascan tiger beetles 111

Figure 1. Major vegetation types and rivers in Madagascar, showing major towns.

Geographical information was derived principally London, England, The Museum´ National d’Histoire from Rivalier (1970), Jeannel (1946), Olsouﬁeff Naturelle (MNHN) in Paris, France, and at the (1934), Horn (1934) and Pearson (1993). All avail- Parc Botanique et Zoologique de Tsimbazaza in able locality data were also taken from labels on spec- Antananarivo, Madagascar. The private collection of imens at The Natural History Museum (BMNH) in Andre´ Peyrieras (Antananarivo, Madagascar) was also 112 L. Andriamampianina et al.

Centre (1991). See Lees et al. (1999) for additional details. Two distributional databases were created for the Malagasy tiger beetles, one at the generic level and one at the specific level. Ankarafantsika All localities were checked against the F.T.M. (1979–1985) 1 : 500,000 map series and F.T.M. (1987) Masoala 1 : 1,000,000 map series. Viette (1991), annotated with grid-cell references, was used as a principal reference to Fenerive Est standardise localities across taxa. Each locality was ref- Ivoloina Park erenced to one grid-cell, which may include other localities. When a given locality was too large to fit into one Perinet Analamazaotra Kirindy/ cell or when locality data were imprecise (e.g. ‘South CFPF Manjakatompo Antongil bay’) an appropriate grid-cell was selected as default for all similar instances. Locality data that were too imprecise (e.g. ‘E. Madagascar’) were not used. Isalo Most of the empirical field data used in this paper national park were obtained during the course of more general inven- Tulear tories that included other groups. Field samples were far from exhaustive at any one site. Furthermore, at the spatial resolution of the study there were many grid squares that could be filled neither by existing data nor Figure 2. Sites visited during recent fieldwork (1993–1998): black new field studies. To avoid severe under-estimation of points indicate study sites. The size of the black points is proportional the range occupancy of each species, some form of to the size of the site. Lines drawn between sites show itinerary along predictive mapping was required. which incidental observations were made. To correct for this uneven sampling effort across Madagascar, species ranges were estimated by geographical interpolation of the available data. To interpo- checked. In addition, based on a preliminary analy- late, range continuity was assumed between recorded sis of the literature and museum collection data, tiger limits of qualifying grid-cells. Grid-cells were deemed beetle expeditions were recently carried out across ‘qualifying’ if they contained appropriate habitat in the Madagascar (Figure 2) to fill in data at several sites elevational range of the species, based on available dis- where sampling gaps were identified. Figure 2 shows tributional data. Although many factors (biotic as well recent inventory sites. as abiotic) affect the distribution of beetles, for simplic- ity, these two factors alone were used to guide interpo- Data processing lation. For each species, elevational range and habitat requirements were either determined from sampling WORLDMAP IV was used to analyse patterns of records if available, or inferred from locality details. spatial diversity. WORLDMAP is a graphical tool for The resulting ranges of taxa contained gaps depending interactive assessment of priority areas for conserving on patchiness of habitat. Whether ranges in nature are biodiversity (Williams 1994). Madagascar was divided more or less fragmented than such fairly crude inter- into 912 grid-cells on a quarter degree grid, each cell polated models can only be determined by ground- (measuring approximately 27 km × 27 km, area about truthing, guided by such range maps, followed by more 729 km2 varying with latitude by 4%) referenced detailed statistical correlation of empirical records with by a number. Every cell was coded for habitat types geophysical datasets. However, whereas co-occurrence and elevational range present in the cell. Habitat cate- within an area as large as a quarter degree square by gories were derived from the Foibe Taosaritanin Mada- no means implies sympatry in space or time, the spatial gasikara (F.T.M.) 1979–1985, 1 : 500,000 map series resolution chosen for this analysis is still fine enough to of Madagascar, and habitat areas in the humid for- reflect biogeographic patterns implicit in the distribu- est biome were checked using recent (1985) satel- tion of empirical records, while coarser than the scales lite imagery from Green and Sussman (1990), also typical of metapopulation studies (see Lees et al. 1999, shown in the map of World Conservation Monitoring for additional details). Biogeography and conservation of Madagascan tiger beetles 113

Given the potential errors implicit in the interpo- was also estimated for other selected taxa, namely lated data, some rules were adopted to minimise over- birds, lemurs, chameleons, frogs, butterflies, syntomine estimation of range-size and to increase consistency: moths and enariine scarab beetles. (1) interpolation was carried out only on taxa for which we had at least two empirical grid-cell records. Priority areas analysis (2) Interpolation relied on a minimum convex polygon Two types of priority areas were considered: priori- (convex hull) drawn around all reliable outlying grid- ties for conservation of tiger beetles, and priorities for cell records. (3) Interpolation was not carried out across research. gaps greater than or equal to three degrees latitude The 44 Malagasy protected areas cover approxi- (12 grid-cells) or two degrees longitude (8 grid-cells), mately 3% of the island’s planar surface area (ANGAP reflecting the anisotropic nature of species ranges in 1998). In the analysis of priority areas for conserva- Madagascar. This rule permitted disjunction in the dis- tion, we assumed that species found inside a reserve tribution of some taxa (and thus allowed conservatively are fully protected. We therefore excluded all squares for more fragmented ranges than in the study of Lees corresponding to the current reserve network (along et al. 1999). (4) Below the species level, the distribution with the taxa they represent) before proceeding to of subspecies was also considered. No interpolations the priority areas analysis. In other words, priority were made to join the distributions of two otherwise areas for conservation identified in this work are com- disjunct subspecies. (5) Records considered unreliable plementary to those included in current reserves. In were not used for the interpolation. order to ensure adequate conservation, only squares where reserve area overlaps more than 20% of a grid- Data analysis cell area were considered to be adequately protected (Figure 3). All data analyses were carried out on interpolated plus Thus, a set of priority areas for conservation is pro- empirical data. posed in this paper to complete the representation of all tiger beetle taxa at least once. Squares within the set Diversity patterns and measurements were selected by a stepwise analysis considering taxon Two measures, taxon richness and range-size rarity, richness and range-size rarity. Some areas, unavailable were used to describe patterns of biodiversity. Taxon for protected area status, are excluded from the analysis richness was calculated as the number of taxa recorded a priori. per grid-cell. Range-size rarity was here expressed in Priority areas for research were determined based the sense of restricted range size. This measure of mainly on the spatial pattern of data (sampling gaps). endemism is on a sliding scale, quantified for each cell Given the historical nature of much of the data used in as the sum of the range-size rarity scores of the fauna. this study and the land use changes that have occurred The score for each taxon is defined as the inverse of its throughout Madagascar (Green & Sussman 1990; occupancy (number of squares that contain this taxon; Nelson & Horning 1993), it is necessary to update maps Williams 1994). In the analysis of diversity patterns, the with new faunal and geophysical data in order to direct term ‘hotspot’ is used to refer to cells with the highest conservation planning. scores for richness or range-size rarity. Maps are shown in grey scales, with darker shades indicating the higher Results and discussion scores. Database Effectiveness of the current protected areas network In total, 176 species belonging to 17 genera have been The effectiveness of the current reserve system (now 44 listed in this study (Appendix). Of these, a total of 49 reserves, comprising 57 grid-cells, 49 of which over- species were observed during recent fieldwork, includ- lap a reserve polygon by at least 20%: Figure 3) to ing many that were known from only a few records represent tiger beetles was assessed against the near- (e.g. Chaetotaxis descarpentriesi Deuve, Stenocosmia minimum set (for at least one representation for each angusta Rivalier). For the four species described by Madagascan species) computed by WORLDMAP. In Maran (1942), no distributional data could be found addition to cicindelids, the scope of protected areas (Appendix). 114 L. Andriamampianina et al.

Figure 3. Distribution of the current protected area network in Madagascar on a quarter degree grid map. The 49 grid-cells with 20% or more overlap with a reserve are shown as light grey squares.

A total of 1572 grid-cell/species data points were grid-cells among the total of 912 cells for the whole assembled based on empirical records, and 14,038 island (25.21%, see Figure 3). The ratio of interpo- additional data entries were added by interpolation. lated to empirical records (8.9 : 1) reﬂects a large gap in The empirical records mapped on to a total of 230 sampling effort. Biogeography and conservation of Madagascan tiger beetles 115

Distribution and diversity pattern of the island (Figure 4). The main gaps are found in the tiger beetles north surrounding the high mountains of Tsaratanana and the Androna and the Analavory plateaus. Many Figure 4 shows the grid-cells for which we have areas in western Madagascar are also poorly sam- empirical data; distribution patterns resulting from pled including the Ambongo region; the Bemaraha interpolation are given in Figures 5a,b and 6a,b. plateau and the Makay massif; as are many areas in the central high-plateau (such as the Beveromay and Ankazobe plateaus), Bongolava and Itremo massifs Distribution and gaps and Ranotsara basin. Under-sampling is also iden- Tiger beetles are distributed within a wide range of tified in the extreme south around the Ivakoany habitats throughout Madagascar. There is an excess Massif and Mahafaly plateau. Even in the middle of range-restricted taxa at both specific (Lees et al. of the eastern rainforest, substantial gaps are iden- 1999) and generic levels. As with most other taxo- tified between established reserves (e.g. around the nomic groups, however, many sites are severely under- Kalambatritra massif, and in the region of Fandriana, sampled for cicindelids. Sampling gaps occur all over Figure 4).

Figure 4. Distribution of empirical records. Black points correspond to grid-cells where real data were observed/collected. Places or regions where there are sampling gaps are indicated by circles. 116 L. Andriamampianina et al.

Figure 5a.

Patterns of diversity and endemism of Tsaratanana massif, in the north-west region of At the generic level, the richest areas are in the north- Ambongo, Ankarafantsika and Maevatanana, and in west and in the south. In contrast, areas around central the extreme south around the Ivakoany massif. and eastern Madagascar show lower diversity. Areas Areas with high species richness are localised in of high-generic richness (Figure 5a) and endemism the northern portion of rainforest, with highest scores (Figure 5b) are found in the north in the vicinity recorded in the vicinity of the Masoala peninsula Biogeography and conservation of Madagascan tiger beetles 117

Figure 5b. Maps showing pattern of distribution of Madagascan tiger beetles at generic level according to (a) richness in genera; (b) aggregate endemism per grid-cell. Darker shades indicate higher values. Number of genera (interpolated species ranges) shown in a. and Maroantsetra and the mid-latitude region around The high number of species found in rainforest is due Zahamena (Figure 6a). High concentrations of range- to the pronounced radiation of the genera Pogonostoma restricted species are found in the rainforest from north and Physodeutera within the eastern rainforest biome. to south around Montagne d’Ambre, Tsaratanana, In fact, these two large genera comprise 75.7% of all Masoala and Zahamena; and in the west around Malagasy cicindelids. It seems likely that the highly Ankarafantsika (Figure 6b). dissected topography of the eastern part of the island 118 L. Andriamampianina et al.

Figure 6a. and therefore the relatively large extent of the rain- Patterns of taxic richness and range-size rarity forest biome for animals of small body size, has showed a positive correlation at both the speciﬁc level created more microhabitats than in other ecosystems (r = 0.91) and generic level (r = 0.79). There is lower and has promoted variation in species composition at correlation between the two taxonomic levels, for rich- smaller spatial scales (see Rosenzweig 1995; Fjeldsa&˚ ness (r = 0.54) and endemism (r = 0.51). These results Lovett 1997). are mainly due to the uneven distribution of species Biogeography and conservation of Madagascan tiger beetles 119

Figure 6b. Maps showing pattern of distribution of Madagascan tiger beetles at speciﬁc level according to (a) species richness; (b) aggregate endemism per grid-cell. Darker shades indicate higher values. Number of species (interpolated ranges) shown in a. amongst the 17 genera and substantial differences in The scope of the current protected area range distribution between species and/or genera. Thus, network patterns of diversity vary with taxonomic level and the measure utilised. This point should be considered for Near-minimum sets of areas any distributional analysis. In this study, all priority Considering the entire set of grid-cells, representation area analyses are carried out on the species database. of all members of the family, with at least a single 120 L. Andriamampianina et al. representation per species, requires a near-minimum that their range sizes are likely to be very small, their set of 30 grid-cells. However, only three quarter-degree habitats are unusual, or their abundance has been below grid-cells would theoretically represent half of the detection level. At present levels of knowledge, all 30 known Malagasy tiger beetle species and eight squares areas have high biological and conservation value, and would cover two-thirds (Table 1). need to be considered to ensure full protection of tiger Three important points emerge from this table. Many beetles. of the grid squares, notably those listed in steps 19–30, The 49 grid-cells occupied by current reserves would have ‘unique species’ not found in any other squares in theory protect representatives of all 17 genera of of the island. Only 6 of these 30 squares are included tiger beetles. However, only 139 species out of the in the current reserves network. Finally, many of the total 172 species (80.8%) are included, even though squares are situated in the eastern rainforest, the best- only 30 squares chosen by complementarity would be surveyed biome of Madagascar. Although there is little sufﬁcient to include all species. Thus, many of the cur- doubt that the distributions of these ‘unique’ species rent protected areas might appear to be redundant when are underestimated (if the taxa are not oversplit: Lees considering the protection of tiger beetles with at mini- et al. 1999), the fact that they escaped the eyes and traps mum a single representation. However, it is very impor- of various collectors at other localities suggests either tant to notice that a better protection strategy would

Table 1. Priority grid-cells suggested by the near-minimum set for one representation of Madagascan tiger beetles, in order of complementarity. Step Square Grid-cell Species richness Current reference name A CNS Cum% reserves 1 153 Maroantsetra 57 57 33.14 N 2 29 Montagne d’Ambre 21 78 45.35 Y 3 281 Maevatanana 15 93 54.07 N 4 688 Fianarantsoa 12 105 61.05 N 5 83 Tsaratanana Reserve 8 113 65.7 Y 6 365 Zahamena Reserve 6 119 69.19 Y 7 816 Vondrozo 5 124 72.09 N 8 563 Andranomena reserve 5 129 75 Y 9 37 Ankarana Reserve 4 133 77.33 Y 10 98 Andapa-Andrakata! 4 137 79.65 N 11 24 Diego-Suarez 4 141 81.98 N 12 961 Fort-Dauphin 3 144 83.72 N 13 253 Sitampiky; Ambongo 3 147 85.47 N 14 475 Antananarivo 3 150 87.21 N 15 861 Ianapera 3 153 88.95 N 16 80 Maromandia 2 155 90.12 N 17 343 Fenerive 2 157 91.28 N 18 948 Col Manangotry 2 159 92.44 N 19 364 Andranomalaza 2 161 93.6 N 20 292 Soanierana Ivongo 1 162 94.19 N 21 125 Antalaha 1 163 94.77 N 22 388 Mitanoka; Onibe River 1 164 95.35 N 23 623 Sakaleona R 1 165 95.93 N 24 737 Vohilava-Faraony 1 166 96.51 N 25 191 Mandritsara 1 167 97.09 N 26 57 Vohemar 1 168 97.67 N 27 183 Bebokay; Ambalabe 1 169 98.26 N 28 50 Nosy Be, Lokobe 1 170 98.84 Y 29 139 Antakotako 1 171 99.42 N 30 514 Tsiafajavona 1 172 100∗ N A = Number of added species, CNS = cumulative number of species, Cum% = cumulative percentage of species. ∗100% corresponds to 172 species because no distributional data is known for the four species of Maran (1942). Biogeography and conservation of Madagascan tiger beetles 121 consider more than a single representation of each conservation problem in Madagascar is indicated by species (Usher 1986) as well as contiguity of protected our finding that once the 139 tiger beetle species pro- cells, a conservation criterion implicit in the ‘rescue tected by the current reserve network are removed from effect’ in metapopulation dynamic theory (Brown & the analysis, an additional 23 squares would be required Kodric-Brown 1997; Hanski 1982), which directly to protect the remaining 33 (19.2%) species, based on leads to the need for habitat corridors. Moreover, the their current known distribution, and at the spatial res- consideration of 20% grid-cell overlap by reserve in olution used here for analysis. this study does not take into account the notion of The set of areas given in Figure 7 would complete long-term minimum viable area and/or population for representation of known Madagascan tiger beetle a species, nor allows for range shifts along latitudinal species, in addition to those already protected by the and elevational gradients in the event of rapid climate 44 existing reserves. change. In addition, protection of sites for all species Although it is acknowledged that (for sampling of Cicindelidae would not in any way guarantee reasons alone) different taxa will suggest different protection of the rest of the flora and fauna. orders of priority areas (Vane-Wright et al. 1994; By comparison with the 80.6% of tiger beetles Prendergast & Eversham 1997), a priority areas list covered using the 20% grid-cell overlap criterion in for conservation of tiger beetles is given here (Table 2). the current reserve system (this value increases to 84% The square 24 corresponding to Montagne des Français using a 1% criterion), about 97% of lemurs and bird is identified as the first priority to be considered for species are represented, 92% of tenrecs, as well as more additional conservation of cicindelids. This massif is than 91% of butterflies, more than 82% of chameleons surrounded by dry forests and wooded grasslands that and frogs; more than 64% of syntomine moths and over grow on a combination of various rock types includ- 55% of enariine scarab beetles, based on current knowl- ing unconsolidated sands, sandstone and some Tertiary edge of the ranges of these groups (Lees et al. 1999). and also Mesozoic limestone with marls and chalks The high proportion of protected lemurs and birds prob- (Du Puy & Moat 1996). ably results from the fact that lemurs and birds are the A few areas are included in this list of priority areas best studied groups in Madagascar, especially within that could be considered as corridors between estab- protected areas; and that they tend to have wider ranges lished reserves. Thus the Vondrozoregion (square 816) and their limits are least severely under-estimated (Lees links the three reserves of Manombo, Kalambatritra et al. 1999). Furthermore, some reserves were created and also Pic d’Ivohibe, potentially providing con- partly to protect some of these charismatic vertebrates. tinuity between these reserves. The area surround- In contrast, the other groups lack data from several ing Mitanoka (388) is likewise between Zahamena, reserves and we can expect that percentages of known Mangerivola and Betampona, three reserves situated species represented in the reserve system will increase in the Middle East. if further inventories are conducted. Nevertheless, this Furthermore, there are several areas that would ex- study indicates strongly that new protected or man- tend established reserves. These include Maroantsetra aged areas need to be created to ensure protection of (153) in the north-west of Masoala National Park; other elements of biodiversity, the majority of species Mandritsara (191) west of Makira plateau located in of which seem likely to have relatively narrow ranges, the north of Marotandrano special reserve; Ankara small body size and/or restricted mobility (Lees et al. (304) next to Kasijy special reserve; Maromandia 1999). (80) west of Manongarivo reserve; the part of Bongolava massif around Ambaravaranala (422), in Priority areas for conservation the north-east of Ambohijanahary special reserve; Ankoadava Mahabo next to Andranomena special For the identification of priority areas for tiger bee- reserve; Tsintongambarika forest next to Mandena tle conservation, we assumed optimistically that each reserve and finally the area next to Fort-Dauphin (961), of the current reserves in Madagascar furnishes ade- south of Andohahela reserve. quate protection for their included fauna and flora and Finally, there are areas located in some bioclimatic, we have not considered questions of reserve area and geological or vegetation classes that are not well rep- habitat continuity for population or ecosystem via- resented in the current protected areas network of bility, nor the effects of climate change in relation Madagascar, like Ankazotelo (878) which is part of to anthropogenic fragmentation. The urgency of the the Mahafaly spiny forest region in the south; and the 122 L. Andriamampianina et al.

Montagne des Francais

Maromandia

Marogaoma Maevatanana Antalaha

Sitampity Ambongo Maroantsetra Mandritsara

Ankara Soanierana Ivongo Fenerive Est Ambaravaranala Mitanoka

Andrangoloaka

Ankoadava Ankazomivady Sakaleona Fianarantsoa

Vohilava Faraony Vondrozo Ankazotelo

Tsitongambarika

Fort-Dauphin

Figure 7. Distribution of the 23 areas identified in this study as additional priorities for the conservation of tiger beetles. surrounding region of Maevatanana (281) in the north- Antananarivo are effectively protected because they are west that is also diverse in rock types, including alluvial treated as sacred. Also, data recorded from this local- and lakes deposits, sandstone and Mesozoic limestone ity may be unreliable because of imprecise labels, since with Marls, as well as ultrabasic, and metamorphic and many early collectors lived and acquired specimens in igneous basement rocks (Du Puy & Moat 1996); and Antananarivo. Therefore, two alternative squares were two areas among the high-plateau, including Anka- chosen instead: Andrangoloaka and Ambaravaranala. zomivady (619), east of the Itremo massif. This area Many of the priority areas correspond to ‘Classified is outstanding because of its geological characteris- Forests’ (COEFOR/CI 1993), parts of the ‘domaine tics that include metamorphic and igneous rocks, plus forestier national’ but lack legal protection. Some of quartzites and marbles (cipolin), two of the rarest still these sites are also suggested as potentially valuable vegetated geological types in Madagascar (Du Puy & foci for research and conservation because of their Moat 1996). This area presents similar geological unique geological characters, which may result in characteristics to the region north of Maroantsetra unique floral and faunal assemblages. For example, (153), another interesting area with a unique species, these include the area north of Antongil bay (north Physodeutera umbrosa. The second high-plateau area of Maroantsetra & Mahalevona), and Ankazomivady, is the Andrangoloaka forest (497), between the two east of Itremo (Du Puy & Moat 1996). Many of these lakes of Mantasoa and Tsiazompaniry. areas, and especially those that could act as corridors Although the area around Antananarivo (square between established reserves, were identified as hav- 475), the capital of Madagascar, appears to be very ing a high, very high or even exceptional biological rich, it has not been included for a number of reasons. or conservation importance in the recommendations of The few remaining semi-natural areas found near the priority-setting workshop (Ganzhorn et al. 1997). Biogeography and conservation of Madagascan tiger beetles 123

Table 2. Results of the priority areas analysis for conservation showing the 23 chosen squares and their contribution to the total biodiversity. Step Square Square Species richness FC PSW reference name SP A C 0 Current Reserves 139 139 139 1 24 Montagne des Franc¸ais 36 4 143 Y Y 2 816 Vondrozo 23 4 147 Y N 3 281 Maevatanana 28 3 150 Y N 4 563 Ankoadava (Mahabo) 15 2 152 Y Y 5 497 Andrangoloaka 31 1 153 N N 6 422 Ambaravaranala 11 1 154 N Y 7 878 Ankazotelo 13 2 156 N Y 8 80 Maromandia 27 1 157 Y Y 9 388 Mitanoka; Namolazana 46 1 158 Y Y 10 191 Mandritsara 28 1 159 Y Y 11 153 Maroantsetra 57 1 160 N Y 12 253 Sitampiky; Ambongo 27 1 161 Y N 13 304 Ankara; Menavava river 23 1 162 N N 14 619 Ankazomivady 23 1 163 Y Y 15 737 Vohilava-Faraony 16 1 164 Y Y 16 87 Marogaoma 30 1 165 Y Y 17 623 Sakaleona 21 1 166 Y Y 18 948 Tsitongambarika 21 1 167 Y Y 19 961 Fort Dauphin 17 1 168 N Y 20 343 Fenerive 27 1 169 Y Y 21 292 Soanierana Ivongo 16 1 170 Y Y 22 125 Antalaha 13 1 171 N N 23 688 Fianarantsoa 29 1 172 N N SP = Number of species, A = number of added species, C = cumulative number of species. The last two columns indicate whether the area corresponds to a ‘classiﬁed forests’ (FC) and whether it is listed in the recommendations from the priority setting workshop (PSW).

Priority areas for research surrounds of Kirindy/CFPF and Ankoadava (Cassola & Andriamampianina 1998), and many more species are A map showing priority areas for research is given in likely to be discovered in western vegetation types. Figure 8. The central region around Antananarivo is Additional vegetationally-distinct sampling gaps are one area of potential interest. In recent times, the region the south and south-west around the Mikea forests, and around Antananarivo has been neglected by biologists the southern spiny thorn forests around the Mahafaly since most of the natural vegetation has been destroyed. and the Ivakoany massif. Furthermore, tiger beetles However, relicts of primary vegetation include the 12 are almost unknown for some habitats such as lime- sacred hills around Antananarivo. Other little known stone karst areas (the Tsingy), and lower montane areas of the central region that could be of interest and montane shrublands such as exist at Tsaratanana, include Ankazobe (Ambohitantely), the Bongolava Anjanaharibe Sud and Marojejy. All those areas should massif (incl. Ambohijanahary), Itremo massif, and be surveyed because many beetles and members of forests along the cliffs of Angavo such as Anjozorobe other groups now risk extinction before even we know and Andrangoloaka. A census of the current condition of their existence. of these areas guided by recent satellite maps could yield surprising data. Other poorly sampled western areas of Madagascar Conclusion are yielding promising results, for example the Sakalava region. Two new species of cicindelids A total of 176 species of Malagasy cicindelid beetles were recently discovered in the middle-west in the are listed here, including all described species, but this 124 L. Andriamampianina et al.

Figure 8. General areas identified as priorities for research work, indicated by circles. list is certainly not comprehensive, and more fieldwork example is the species Stenocosmia angusta, thought to and taxonomic research are required. be confined to the region of Ampijoroa, Ankarafantsika Most field studies to date have been carried out (square 210) since its discovery in 1965, but collected within reserves in Madagascar, both for cicindelids and 480 km away, from the Kirindy/CFPF forest (square other taxa. Therefore, reserves are in general better 563) in 1997. sampled than unprotected areas. This is probably the This study reveals that, for those areas that are rela- reason for the fairly high level of overall biodiversity tively well known, the eastern rainforest, its western apparently covered by the current reserves. Two species extension (the Sambirano), and the extreme north have just been discovered and a number of little known of Madagascar (which comprises transitional forest species have been rediscovered, outside reserves. How- between the humid rainforest and the dry forest) have ever, the distributions of many species are still poorly the richest biodiversity for tiger beetles, a pattern found known (for example, the four species described by in many other groups (Lees et al. 1999). Some areas in Maran (1942) are entirely unknown in terms of the centre, the west and the south appear depauperate. distribution). Whether this reflects reality or an information gap The ranges of many locally endemic species are can only be answered by further fieldwork. Areas of particularly susceptible to underestimation, because of high aggregate endemism occur more patchily all over sampling gaps and also because of their rarity. An the island. The extreme north, the north-west and the Biogeography and conservation of Madagascan tiger beetles 125 extreme south appear most outstanding at the generic Fabio Cassola was particularly helpful with the identifi- level, whereas the north eastern rainforest is remark- cation of the tiger beetles specimens recently collected. able at the specific level. This contrast shown here for a The Ministry of Water and Forests and ANGAP are Madagascan group is interesting in light of the sugges- thanked for their collaboration with research permits. tion that higher taxon richness may be a good surrogate for species richness (Williams & Gaston 1994). Many of the tiger beetle species are covered in the References current protected areas network (85%). Creation of more reserves would, however, be needed to protect all Andriamampianina, L. (1996) Biogeography of enariine (Melolon- of the tiger beetles. A set of 23 squares was identified in thidae) and cicindelid beetles in Madagascar. Unpublished M.Sc. Thesis, University of Kent at Canterbury. this study as potential priority areas or foci for conser- ANGAP (1998) Programme Environmental II. Composante Aires vation action. Rather surprisingly, the high-plateau area Protegees´ et Ecotourisme: rapport annuel des activities´ 1998, 23 p. surrounding Antananarivo (which may still contain an Banque Mondiale, USAID, Cooperation´ Suisse, UNESCO, UNDP unexpected number of minute forest relics of value for and WWF (1988) Madagascar: Plan d’action Environnemental. I: some taxonomic groups) and remaining natural habi- Document de synthese` gen´ erale´ et propositions d’orientations. Brown, J.H. and Kodric-Brown, A. (1977) Turnover rates in insular tats of this high-plateau was identified as important for biogeography: effect of immigration on extinction. Ecology 58, more research, which would check poorly-documented 775–87. historical records for cicindelids in this area. In addi- Carroll, S.S. and Pearson, D.L. (1998) Spatial modeling of butterfly tion, many areas in the south, west and some high species richness using tiger beetles (Cicindelidae) as a bioindicator mountains would be a priority for future surveys taxon. Ecol. Appl. 8, 531–43. Cassola, F. and Andriamampianina, L. (1998) Deux especes` nou- and bioinventory work. This also applies to many velles de Cicindelides´ de Madagascar (Coleoptera Cicindelidae) unprotected areas between established reserves. Boll. Soc. Entomol. Ital. 130, 47–54. The choice of protected areas depends on many COEFOR/CI (1993) Repertoire´ et Carte de Distribution: Domaine factors beyond raw biodiversity scores for selected Forestier de Madagascar. Direction des Eaux et Forets,ˆ Service taxa. Nevertheless, the present study is a contribu- des Ressources Forestieres,` Projet COEFOR (contribution a` l’etude´ des Foretsˆ Classees),´ et Conservation International, tion towards improving the information base relevant 20 pp. + 1 map. to conservation in Madagascar. Added to information Du Puy, D.J. and Moat, J. (1996) A refined classification of the gathered from other studies, it should help in mak- primary vegetation of Madagascar based on the underlying geol- ing better decisions regarding the design, planning and ogy: using GIS to map its distribution and to assess its con- implementation of conservation strategies throughout servation status. In Proceedings of the International Symposium on the Biogeography of Madagascar (W.R. Lourenço, ed), Madagascar. pp. 205–18, +3 maps. Paris: Editions de l’ORSTOM. Emberton, K.C. (1997) Diversities and distributions of 80 land-snail species in southeastern-most Madagascan rainforests, with a report Acknowledgements that lowlands are richer than highlands in endemic and rare species. Biodivers. Conserv. 6, 1137–54. Fjeldsa,˚ J. and Lovett, J.C. (1997) Biodiversity and environmental Andriamampianina’s field work and museum visits stability. Biodivers. Conserv. 6, 315–23. were supported by grants from the Darwin Initiative Foibe Taosaritanin Madagasikara (F.T.M.) (1979–1985) U.K., the Wildlife Conservation Society and the Durrell 1 : 500,000 map series. Foibe Taosaritanin Madagasikara (F.T.M.) (1987) Carte Interna- Institute of Conservation and Ecology, U.K. In par- ◦ tionale du Monde 1 : 1,000,000. Cartes n SDa ` SG-38-39. ticular, Wildlife Conservation Society (Madagascar) Ganzhorn, J.U., Rakotosamimanana, B., Hannah, L., Hough, J., is thanked for the financial and logistical support Iyer, L., Olivieri, S., Rajaobelina, S., Rodstrom, C. and Tilkin, G. for this study. Other authors were supported by (1997) Priorities for biodiversity conservation in Madagascar. BBSRC, Leverhulme Foundation and NERC (David Prim. Report 48(1). Lees). The Natural History Museum in London, Green, G.M. and Sussman, R.W. (1990) Deforestation history of the eastern rain forests of Madagascar from satellite images. Science the Museum´ National d’Histoire Naturelle in Paris, 248, 212–15. the Parc Botanique et Zoologique de Tsimbazaza Hanski, I. (1982) Dynamics of regional distribution: the core and in Antananarivo, M. Andre´ Peyrieras and M. Fabio satellite hypothesis. Oikos 38, 210–21. Cassola are thanked for making their tiger beetle Horn, W. (1934) Les Cicindelides´ de Madagascar. Premiere` partie: Catalogue Bibliographique et Synonymique. Mem. Acad. collection available. We wish to thank Paul Williams Malgache 20, 7–28. for providing the computer program WORLDMAP and Jeannel, R. (1946) Coleopt´ eres` Carabiques de la Region´ Malgache assisting with establishment of the Madagascar grid. (Premiere` partie). Faune de l’ Empire Français 6, 104–206. 126 L. Andriamampianina et al.

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Genus 8: Cicindelina Jeannel, 1946 P. pokornyi Maran, 1942 (?) C. oculata Chaudoir, 1843∗ P. megalommo¨ıdes Walter Horn, 1896 P. belalonensis Deuve, 1987 Genus 9: Chaetotaxis Jeannel, 1946 P. flagellicorne Walter Horn, 1897∗ C. rugicollis Fairmaire, 1871∗ P. pseudo-trimaculata Walter Horn, 1934 C. cicindeloides Walter Horn, 1905 P. andriai Rivalier, 1965 C. descarpentriesi Deuve, 1987∗ P. dorri Fleutiaux, 1899 C. soalalae Fairmaire, 1903 ∗ P. rectipenis Walter Horn, 1934 C. ankarahitrae Jeannel, 1946 P. centropunctata Walter Horn, 1934 C. serieguttata Walter Horn, 1934 P. biguttula Fairmaire, 1903 C. macropus Chaudoir, 1865 P. dubia Maran, 1942 (?) C. grandidieri Kunckel, 1887∗ P. marginemaculata Walter Horn, 1934 C. semi-confluens Rivalier, 1965 P. peyrierasi Rivalier, 1967 C. leptographa Rivalier, 1965 P. rectolabialis Walter Horn, 1913 Genus 10: Ambalia Jeannel, 1946 P. punctipenne Fairmaire, 1903 A. aberrans Fairmaire, 1871∗ P. virgulata Fairmaire, 1904 A. satura Rivalier, 1965 P. bucephala Walter Horn, 1900 P. sikorae Walter Horn, 1896∗ Genus 11: Calyptoglossa Jeannel, 1946 P. subtilivelutina Walter Horn, 1934b C. frontalis Audouin et Brulle,´ 1839∗ P. viridi-cyanea Audouin et brulle,´ 1839∗ Genus 12: Peridexia Chaudoir, 1860 P. maximum Fleutiaux, 1899 P. fulvipes Dejean, 1831∗ P. janthina Fairmaire, 1903 P. hilaris Fairmaire, 1883∗ P. natalia Walter Horn, 1934 P. catalai Jeannel, 1946 Genus 13: Prothyma Hope, 1838 P. alluaudi Fleutiaux, 1903∗ P. radama Kunckel, 1887∗ P. tricolorata Walter Horn, 1934 Genus 14: Physodeutera Lacordaire, 1843 P. perrieri Rivalier, 1967 P. adonis Castelnau, 1835 Genus 15: Walterhornia Olsoufieff, 1934 P. intermedia Rivalier, 1967 W. speculifera Walter Horn, 1934∗ P. parcepunctata Jeannel, 1946 P. uncifera Jeannel, 1946 Genus 16: Stenocosmia Rivalier, 1965 P. lateralis Olsoufieff, 1934 S. angusta Rivalier, 1965∗ P. bellula Fleutiaux, 1886 S. tenuicollis Fairmaire, 1904 P. trimaculata Fleutiaux, 1889∗ P. sobrina Rivalier, 1967 Subfamily 2: Collyrinae Csiki, 1906 P. debilis Rivalier, 1967 P. uniguttata Fairmaire, 1871 Genus 17: Pogonostoma Klug, 1835 P. komorousi Maran, 1942 (?) P. cylindricum Fleutiaux, 1899∗ P. fairmairei Walter Horn, 1899∗ P. brevicorne Walter Horn, 1898 P. hajni Maran, 1942 (?) P. vestitum Fairmaire, 1900∗ P. rufosignata Audouin et Brulle,´ 1839∗ P. violaceum Fleutiaux, 1902 P. mocquerysi Fleutiaux, 1899∗ P. septentrionale Fleutiaux, 1903 P. punctum Rivalier, 1951∗ P. cyanescens Klug, 1835∗ P. umbrosa Rivalier, 1967 P. caerulea Castelnau & Gory, 1837∗ P. cyanea Audouin et Brulle,´ 1839 P. violaceolevigata Walter Horn, 1927 P. minima Walter Horn, 1893 P. sambiranense Rivalier, 1965 P. madari Maran, 1942 (?) P. affine Horn, 1893 P. vadoni Rivalier, 1951 P. chalybaeum Klug, 1835∗ P. rubescens Jeannel, 1946 P. spinipennis Castelnau et Gory, 1837 P. pseudorubescens Deuve, 1987 P. rugosoglabra Walter Horn, 1923 P. perroti Rivalier, 1967 P. propinquum Rivalier, 1970∗ P. gigantea Walter Horn, 1913 128 L. Andriamampianina et al.

P. globicolle Rivalier, 1965 P. sericeum Klug, 1835 P. perroti Rivalier, 1970 P. perrieri Fairmaire, 1900 P. malleatum Rivalier, 1970 P. kraatzi Walter Horn, 1894 P. peyrierasi Rivalier, 1970 P. ovicolle Walter Horn, 1893 P. densepunctatum Rivalier, 1970 P. microtuberculatum Walter Horn, 1934 P. subtilis Walter Horn, 1904 P. longicolle Jeannel, 1946 P. hamulipenis Walter Horn, 1934 P. anthracina Castelnau et Gory, 1837 P. tortipenis Walter Horn, 1934 P. maculicorne Walter Horn, 1934 P. impressum Rivalier, 1970 P. humbloti Rivalier, 1970 P. externespinosa Walter Horn, 1934 P. excisoclavipenis Walter Horn, 1934 P. elegans Brulle,´ 1834∗ P. basidilatatum Walter Horn, 1909 P. abadiei Rivalier, 1965 P. pusilla Castelnau et Gory, 1937 P. brullei Castelnau & Gory, 1837 P. laportei Walter Horn, 1900∗ P. alluaudi Walter Horn, 1898 P. inerme Jeannel, 1946 P. rufidens Rivalier, 1970 P. flavomaculatum Walter Horn, 1892 P. atrorotundata Walter Horn, 1934 P. beananae Rivalier, 1963 P. sudiferum Rivalier, 1965 P. moestum Rivalier, 1970 P. subtiligrossa Walter Horn, 1934∗ P. schaumi Walter Horn, 1893∗ P. litigiosum Rivalier, 1970 P. mathiauxi Jeannel, 1946 P. ankaranense Deuve, 1986 P. fleutiauxi Walter Horn, 1905∗ P. srnkai Walter Horn, 1893 P. basale Fleutiaux, 1899 P. gibbosum Rivalier, 1970 P. globulithorax Jeannel, 1946 P. meridionale Fleutiaux, 1899 P. parallelum Walter Horn, 1909 P. sikorae Walter Horn, 1894∗ P. geniculatum Jeannel, 1946 P. flavomaxillaris Cassola & P. horni Fleutiaux, 1899 Andriamampianina, 1998∗ P. rugosiceps Rivalier, 1970 P. pseudo-minimum Walter Horn, 1934 P. nigricans Klug, 1835 P. flavopalpale Jeannel, 1946 P. vadoni Jeannel, 1946 P. infimum Rivalier, 1970 P. pallipes Rivalier, 1970 P. gladiator Walter Horn, 1934 P. mocquerysi Fleutiaux, 1899 P. phalangio¨ıde Rivalier, 1970 P. delphinense Jeannel, 1946 P. levigatum Walter Horn, 1908 P. minimum Fleutiaux, 1899 P. comptum Rivalier, 1970 P. sicardi Walter Horn, 1927 P. surdum Rivalier, 1970 P. parvulum Rivalier, 1970