Taxic Richness Patterns and Conservation Evaluation of
Transcription
Taxic Richness Patterns and Conservation Evaluation of
Journal of Insect Conservation 4: 109–128, 2000. © 2000 Kluwer Academic Publishers. Printed in the Netherlands. Taxic richness patterns and conservation evaluation of Madagascan tiger beetles (Coleoptera: Cicindelidae) Lantoniaina Andriamampianina1,∗ , Claire Kremen2 , Dick Vane-Wright3 , David Lees4 & Vincent Razafimahatratra5,† 1 Wildlife Conservation Society, BP 8500 Antananarivo 101, Madagascar 2 Center for Conservation Biology and Wildlife Conservation Society, Department of Biological Sciences, Stanford University, Stanford, California 94305, U.S.A. 3 Biogeography and Conservation Laboratory, Department of Entomology, 4 Department of Palaeontology, The Natural History Museum, Cromwell Road, South Kensington, SW7 5BD, U.K. 5 Faculté 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 The high level of biological diversity and local endemism in Madagascar reflects not only long isolation but the existence of a great diversity of habitats, climatic zones, topography and soil types (Lowry et al. 1997). Recently, the Malagasy government has developed a national environmental action plan that aims to conserve this unique fauna and flora. One of the most important strategic objectives defined in this plan is the development of an efficient protected areas network (Banque Mondiale et al. 1988; ONE 1997). Knowledge of patterns of biodiversity is essential to guide conservation planning. In April 1995, a workshop was held in Madagascar that aimed to produce a concerted set of national biodiversity priorities. The thematic groups treated during this workshop were, however, large and vague (e.g. invertebrates, aquatic ecosystems), and species identifications were often uncertain. Except possibly for lemurs and birds, available data did not reflect the detailed distribution of the taxa (Ganzhorn et al. 1997). Since then, several works have used smaller, taxonomically better defined groups to refine the patterns of biodiversity and to set priority areas in Madagascar (Emberton 1997; Lees 1997; Lees et al. 1999). This paper concerns quantitative biogeography of Madagascan tiger beetles (Coleoptera: Cicindelidae). Cicindelids have been considered to pass a range of test criteria proposed (Pearson & Cassola 1992) for a promising indicator group: taxonomy stable; biology well-known; easy to observe in the field; occurring in a broad range of habitat types; individual species showing tendency to be specialised within a narrow habitat; and finally, preliminary evidence that diversity patterns are similar to those for other taxa. Positive 110 L. Andriamampianina et al. correlations found in the distribution of tiger beetles, birds and butterflies in North America, in the Indian subcontinent and in Australia at a coarse spatial scale (Pearson & Cassola 1992) have for some but not all relationships remained significant after being subjected to more rigorous spatial statistics taking into account spatial autocorrelation (Carroll & Pearson 1998; Pearson & Carroll 1998). We describe the aggregate distribution of all known tiger beetles (family Cicindelidae) in Madagascar, and consider in particular, species richness patterns at a fine quarter degree resolution and areas of geographical concentration of range-restricted species. In this paper we do not examine the more general question of the relative efficiency of tiger beetles in representing other groups of organisms. However, a previous study (Lees et al. 1999) using a nearly identical dataset showed that known ranges of Malagasy cicindelids exhibit a hollow-curve range-size frequency distribution skewed to more narrow ranges, which makes their richness patterns considerably less influenced by the geometric effects of Madagascar’s boundaries on species richness than for some other groups such as butterflies and vertebrates. Although the overall range-size frequency distribution of all Madagascar’s biota is unknown, terrestrial vertebrates (about 1000 species) would seem highly unlikely to be representative. In this paper, we evaluate the efficiency of the protected areas network in Madagascar (encompassing 57 quarter degree grid areas) to represent known species of Malagasy tiger beetles. We identify a set of priority areas for conservation action, and other areas where more field research is required. Madagascan tiger beetles The Cicindelidae is one of Madagascar’s best-known insect groups. The Malagasy tiger beetle fauna has been the subject of many studies summarised in Andriamampianina (1996). Although most papers are on taxonomy and systematics, extensive distributional data on this group in Madagascar is available in the literature and in museum collections. The cicindelid fauna of Madagascar (n = 176) is the third richest of any country in the world (Pearson & Cassola 1992), with more than 99% endemism at the species level. Just one species, Myriochile melancholica Fabricius, represented in Madagascar by the endemic subspecies M.m. trilunaris Klug, is also found in the mainland Africa. Although some species can be found widely across the island, most have very localised distributions. In this paper we follow the taxonomy of Rivalier (1950, 1965, 1967, 1970). Malagasy tiger beetles are divided into two subfamilies: – The Collyrinae, represented in Madagascar by a unique genus Pogonostoma. This entirely endemic genus comprises 81 arboreal species. Generally found in primary forests, they are more diverse in the evergreen eastern rainforests than in the dry forests. Adult beetles are found most of the time on tree trunks, pursuing small invertebrates. From time to time, they also chase their prey in the surrounding herbaceous strata. Their larvae live in holes in the tree trunks. Because of their arboreal habit, members of this genus are highly threatened by deforestation. – The Cicindelinae, comprising 16 other genera of tiger beetles with terrestrial habitats. Altogether, they include 95 species. They are mainly ground dwelling, although many of them are found only in forests. Their larvae construct tubes in the soil. Both adults and larvae prey on small invertebrates. Some of the genera such as Physodeutera, Peridexia and Calyptoglossa are common in rainforests, whereas others (Chaetotaxis, Prothyma, Stenocosmia, Waltherhornia) are mainly found in dry forest and arid places such as prairies, savannas or the xerophytic formations of the western, southern and central parts of Madagascar (Figure 1). Some others (notably Chaetodera, Lophyridia, Habrodera and Lophyra) are found in abundance in sandy places along beaches and rivers, mainly in the west and south of Madagascar. Finally, there are those which are highly ubiquitous in habitat preference, occurring wherever suitable open substrate exists for breeding (Hipparidium, Ambalia, Cylindera, Cicindelina and Myriochile), such as on roads through towns, in villages, in forests or woodlands on shaded trails, and along rivers. Methodology Data collection A species checklist for the family was established through literature surveys based on recent revisions of Pogonostoma by Rivalier (1970), Physodeutera by Rivalier (1967), and Jeannel (1946) for all other genera. Synonyms were also used in the search for distributional 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 from Rivalier (1970), Jeannel (1946), Olsoufieff (1934), Horn (1934) and Pearson (1993). All available locality data were also taken from labels on specimens at The Natural History Museum (BMNH) in London, England, The Muséum National d’Histoire Naturelle (MNHN) in Paris, France, and at the Parc Botanique et Zoologique de Tsimbazaza in Antananarivo, Madagascar. The private collection of André Peyrieras (Antananarivo, Madagascar) was also 112 L. Andriamampianina et al. Ankarafantsika Masoala Fenerive Est Ivoloina Park Kirindy/ CFPF Perinet Analamazaotra Manjakatompo Isalo national park Tulear Figure 2. Sites visited during recent fieldwork (1993–1998): black points indicate study sites. The size of the black points is proportional to the size of the site. Lines drawn between sites show itinerary along which incidental observations were made. checked. In addition, based on a preliminary analysis of the literature and museum collection data, tiger beetle expeditions were recently carried out across Madagascar (Figure 2) to fill in data at several sites where sampling gaps were identified. Figure 2 shows recent inventory sites. Data processing WORLDMAP IV was used to analyse patterns of spatial diversity. WORLDMAP is a graphical tool for interactive assessment of priority areas for conserving biodiversity (Williams 1994). Madagascar was divided into 912 grid-cells on a quarter degree grid, each cell (measuring approximately 27 km × 27 km, area about 729 km2 varying with latitude by ± 4%) referenced by a number. Every cell was coded for habitat types and elevational range present in the cell. Habitat categories were derived from the Foibe Taosaritanin Madagasikara (F.T.M.) 1979–1985, 1 : 500,000 map series of Madagascar, and habitat areas in the humid forest biome were checked using recent (1985) satellite imagery from Green and Sussman (1990), also shown in the map of World Conservation Monitoring 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. All localities were checked against the F.T.M. (1979–1985) 1 : 500,000 map series and F.T.M. (1987) 1 : 1,000,000 map series. Viette (1991), annotated with grid-cell references, was used as a principal reference to standardise localities across taxa. Each locality was referenced to one grid-cell, which may include other localities. When a given locality was too large to fit into one cell or when locality data were imprecise (e.g. ‘South 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. Most of the empirical field data used in this paper were obtained during the course of more general inventories 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 new field studies. To avoid severe under-estimation of the range occupancy of each species, some form of predictive mapping was required. To correct for this uneven sampling effort across Madagascar, species ranges were estimated by geographical interpolation of the available data. To interpolate, range continuity was assumed between recorded limits of qualifying grid-cells. Grid-cells were deemed ‘qualifying’ if they contained appropriate habitat in the elevational range of the species, based on available distributional data. Although many factors (biotic as well as abiotic) affect the distribution of beetles, for simplicity, these two factors alone were used to guide interpolation. For each species, elevational range and habitat requirements were either determined from sampling records if available, or inferred from locality details. The resulting ranges of taxa contained gaps depending on patchiness of habitat. Whether ranges in nature are more or less fragmented than such fairly crude interpolated models can only be determined by groundtruthing, guided by such range maps, followed by more detailed statistical correlation of empirical records with geophysical datasets. However, whereas co-occurrence within an area as large as a quarter degree square by no means implies sympatry in space or time, the spatial resolution chosen for this analysis is still fine enough to reflect biogeographic patterns implicit in the distribution of empirical records, while coarser than the scales typical of metapopulation studies (see Lees et al. 1999, for additional details). Biogeography and conservation of Madagascan tiger beetles Given the potential errors implicit in the interpolated data, some rules were adopted to minimise overestimation of range-size and to increase consistency: (1) interpolation was carried out only on taxa for which we had at least two empirical grid-cell records. (2) Interpolation relied on a minimum convex polygon (convex hull) drawn around all reliable outlying gridcell records. (3) Interpolation was not carried out across gaps greater than or equal to three degrees latitude (12 grid-cells) or two degrees longitude (8 grid-cells), reflecting the anisotropic nature of species ranges in Madagascar. This rule permitted disjunction in the distribution of some taxa (and thus allowed conservatively for more fragmented ranges than in the study of Lees et al. 1999). (4) Below the species level, the distribution of subspecies was also considered. No interpolations were made to join the distributions of two otherwise disjunct subspecies. (5) Records considered unreliable were not used for the interpolation. Data analysis All data analyses were carried out on interpolated plus empirical data. Diversity patterns and measurements Two measures, taxon richness and range-size rarity, were used to describe patterns of biodiversity. Taxon richness was calculated as the number of taxa recorded per grid-cell. Range-size rarity was here expressed in the sense of restricted range size. This measure of endemism is on a sliding scale, quantified for each cell as the sum of the range-size rarity scores of the fauna. The score for each taxon is defined as the inverse of its occupancy (number of squares that contain this taxon; Williams 1994). In the analysis of diversity patterns, the term ‘hotspot’ is used to refer to cells with the highest scores for richness or range-size rarity. Maps are shown in grey scales, with darker shades indicating the higher scores. 113 was also estimated for other selected taxa, namely birds, lemurs, chameleons, frogs, butterflies, syntomine moths and enariine scarab beetles. Priority areas analysis Two types of priority areas were considered: priorities for conservation of tiger beetles, and priorities for research. The 44 Malagasy protected areas cover approximately 3% of the island’s planar surface area (ANGAP 1998). In the analysis of priority areas for conservation, we assumed that species found inside a reserve are fully protected. We therefore excluded all squares corresponding to the current reserve network (along with the taxa they represent) before proceeding to the priority areas analysis. In other words, priority areas for conservation identified in this work are complementary to those included in current reserves. In order to ensure adequate conservation, only squares where reserve area overlaps more than 20% of a gridcell area were considered to be adequately protected (Figure 3). Thus, a set of priority areas for conservation is proposed in this paper to complete the representation of all tiger beetle taxa at least once. Squares within the set were selected by a stepwise analysis considering taxon richness and range-size rarity. Some areas, unavailable for protected area status, are excluded from the analysis a priori. Priority areas for research were determined based mainly on the spatial pattern of data (sampling gaps). Given the historical nature of much of the data used in this study and the land use changes that have occurred throughout Madagascar (Green & Sussman 1990; Nelson & Horning 1993), it is necessary to update maps with new faunal and geophysical data in order to direct conservation planning. Results and discussion Database Effectiveness of the current protected areas network The effectiveness of the current reserve system (now 44 reserves, comprising 57 grid-cells, 49 of which overlap a reserve polygon by at least 20%: Figure 3) to represent tiger beetles was assessed against the nearminimum set (for at least one representation for each Madagascan species) computed by WORLDMAP. In addition to cicindelids, the scope of protected areas In total, 176 species belonging to 17 genera have been listed in this study (Appendix). Of these, a total of 49 species were observed during recent fieldwork, including many that were known from only a few records (e.g. Chaetotaxis descarpentriesi Deuve, Stenocosmia angusta Rivalier). For the four species described by Maran (1942), no distributional data could be found (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 assembled based on empirical records, and 14,038 additional data entries were added by interpolation. The empirical records mapped on to a total of 230 grid-cells among the total of 912 cells for the whole island (25.21%, see Figure 3). The ratio of interpolated to empirical records (8.9 : 1) reflects a large gap in sampling effort. Biogeography and conservation of Madagascan tiger beetles Distribution and diversity pattern of tiger beetles Figure 4 shows the grid-cells for which we have empirical data; distribution patterns resulting from interpolation are given in Figures 5a,b and 6a,b. Distribution and gaps Tiger beetles are distributed within a wide range of habitats throughout Madagascar. There is an excess of range-restricted taxa at both specific (Lees et al. 1999) and generic levels. As with most other taxonomic groups, however, many sites are severely undersampled for cicindelids. Sampling gaps occur all over 115 the island (Figure 4). The main gaps are found in the north surrounding the high mountains of Tsaratanana and the Androna and the Analavory plateaus. Many areas in western Madagascar are also poorly sampled including the Ambongo region; the Bemaraha 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 and Ranotsara basin. Under-sampling is also identified in the extreme south around the Ivakoany Massif and Mahafaly plateau. Even in the middle of the eastern rainforest, substantial gaps are identified between established reserves (e.g. around the Kalambatritra massif, and in the region of Fandriana, 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 At the generic level, the richest areas are in the northwest and in the south. In contrast, areas around central and eastern Madagascar show lower diversity. Areas of high-generic richness (Figure 5a) and endemism (Figure 5b) are found in the north in the vicinity of Tsaratanana massif, in the north-west region of Ambongo, Ankarafantsika and Maevatanana, and in the extreme south around the Ivakoany massif. Areas with high species richness are localised in the northern portion of rainforest, with highest scores 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 Zahamena (Figure 6a). High concentrations of rangerestricted species are found in the rainforest from north to south around Montagne d’Ambre, Tsaratanana, Masoala and Zahamena; and in the west around Ankarafantsika (Figure 6b). The high number of species found in rainforest is due to the pronounced radiation of the genera Pogonostoma and Physodeutera within the eastern rainforest biome. In fact, these two large genera comprise 75.7% of all Malagasy cicindelids. It seems likely that the highly dissected topography of the eastern part of the island 118 L. Andriamampianina et al. Figure 6a. and therefore the relatively large extent of the rainforest biome for animals of small body size, has created more microhabitats than in other ecosystems and has promoted variation in species composition at smaller spatial scales (see Rosenzweig 1995; Fjeldså & Lovett 1997). Patterns of taxic richness and range-size rarity showed a positive correlation at both the specific level (r = 0.91) and generic level (r = 0.79). There is lower correlation between the two taxonomic levels, for richness (r = 0.54) and endemism (r = 0.51). These results 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 range distribution between species and/or genera. Thus, patterns of diversity vary with taxonomic level and the measure utilised. This point should be considered for any distributional analysis. In this study, all priority area analyses are carried out on the species database. The scope of the current protected area network Near-minimum sets of areas Considering the entire set of grid-cells, representation of all members of the family, with at least a single 120 L. Andriamampianina et al. representation per species, requires a near-minimum set of 30 grid-cells. However, only three quarter-degree grid-cells would theoretically represent half of the known Malagasy tiger beetle species and eight squares would cover two-thirds (Table 1). Three important points emerge from this table. Many of the grid squares, notably those listed in steps 19–30, have ‘unique species’ not found in any other squares of the island. Only 6 of these 30 squares are included in the current reserves network. Finally, many of the squares are situated in the eastern rainforest, the bestsurveyed biome of Madagascar. Although there is little doubt that the distributions of these ‘unique’ species are underestimated (if the taxa are not oversplit: Lees et al. 1999), the fact that they escaped the eyes and traps of various collectors at other localities suggests either that their range sizes are likely to be very small, their habitats are unusual, or their abundance has been below detection level. At present levels of knowledge, all 30 areas have high biological and conservation value, and need to be considered to ensure full protection of tiger beetles. The 49 grid-cells occupied by current reserves would in theory protect representatives of all 17 genera of tiger beetles. However, only 139 species out of the total 172 species (80.8%) are included, even though only 30 squares chosen by complementarity would be sufficient to include all species. Thus, many of the current protected areas might appear to be redundant when considering the protection of tiger beetles with at minimum a single representation. However, it is very important 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 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Square reference Grid-cell name Species richness A CNS Cum% Current reserves 153 29 281 688 83 365 816 563 37 98 24 961 253 475 861 80 343 948 364 292 125 388 623 737 191 57 183 50 139 514 Maroantsetra Montagne d’Ambre Maevatanana Fianarantsoa Tsaratanana Reserve Zahamena Reserve Vondrozo Andranomena reserve Ankarana Reserve Andapa-Andrakata! Diego-Suarez Fort-Dauphin Sitampiky; Ambongo Antananarivo Ianapera Maromandia Fenerive Col Manangotry Andranomalaza Soanierana Ivongo Antalaha Mitanoka; Onibe River Sakaleona R Vohilava-Faraony Mandritsara Vohemar Bebokay; Ambalabe Nosy Be, Lokobe Antakotako Tsiafajavona 57 21 15 12 8 6 5 5 4 4 4 3 3 3 3 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 57 78 93 105 113 119 124 129 133 137 141 144 147 150 153 155 157 159 161 162 163 164 165 166 167 168 169 170 171 172 33.14 45.35 54.07 61.05 65.7 69.19 72.09 75 77.33 79.65 81.98 83.72 85.47 87.21 88.95 90.12 91.28 92.44 93.6 94.19 94.77 95.35 95.93 96.51 97.09 97.67 98.26 98.84 99.42 100∗ N Y N N Y Y N Y Y N N N N N N N N N N N N N N N N N N Y N 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 consider more than a single representation of each species (Usher 1986) as well as contiguity of protected cells, a conservation criterion implicit in the ‘rescue effect’ in metapopulation dynamic theory (Brown & Kodric-Brown 1997; Hanski 1982), which directly leads to the need for habitat corridors. Moreover, the consideration of 20% grid-cell overlap by reserve in this study does not take into account the notion of long-term minimum viable area and/or population for a species, nor allows for range shifts along latitudinal and elevational gradients in the event of rapid climate change. In addition, protection of sites for all species of Cicindelidae would not in any way guarantee protection of the rest of the flora and fauna. By comparison with the 80.6% of tiger beetles covered using the 20% grid-cell overlap criterion in the current reserve system (this value increases to 84% using a 1% criterion), about 97% of lemurs and bird species are represented, 92% of tenrecs, as well as more than 91% of butterflies, more than 82% of chameleons and frogs; more than 64% of syntomine moths and over 55% of enariine scarab beetles, based on current knowledge of the ranges of these groups (Lees et al. 1999). The high proportion of protected lemurs and birds probably results from the fact that lemurs and birds are the best studied groups in Madagascar, especially within protected areas; and that they tend to have wider ranges and their limits are least severely under-estimated (Lees et al. 1999). Furthermore, some reserves were created partly to protect some of these charismatic vertebrates. In contrast, the other groups lack data from several reserves and we can expect that percentages of known species represented in the reserve system will increase if further inventories are conducted. Nevertheless, this study indicates strongly that new protected or managed areas need to be created to ensure protection of other elements of biodiversity, the majority of species of which seem likely to have relatively narrow ranges, small body size and/or restricted mobility (Lees et al. 1999). Priority areas for conservation For the identification of priority areas for tiger beetle conservation, we assumed optimistically that each of the current reserves in Madagascar furnishes adequate protection for their included fauna and flora and we have not considered questions of reserve area and habitat continuity for population or ecosystem viability, nor the effects of climate change in relation to anthropogenic fragmentation. The urgency of the 121 conservation problem in Madagascar is indicated by our finding that once the 139 tiger beetle species protected by the current reserve network are removed from the analysis, an additional 23 squares would be required to protect the remaining 33 (19.2%) species, based on their current known distribution, and at the spatial resolution used here for analysis. The set of areas given in Figure 7 would complete representation of known Madagascan tiger beetle species, in addition to those already protected by the 44 existing reserves. Although it is acknowledged that (for sampling reasons alone) different taxa will suggest different orders of priority areas (Vane-Wright et al. 1994; Prendergast & Eversham 1997), a priority areas list for conservation of tiger beetles is given here (Table 2). The square 24 corresponding to Montagne des Français is identified as the first priority to be considered for additional conservation of cicindelids. This massif is surrounded by dry forests and wooded grasslands that grow on a combination of various rock types including unconsolidated sands, sandstone and some Tertiary and also Mesozoic limestone with marls and chalks (Du Puy & Moat 1996). A few areas are included in this list of priority areas that could be considered as corridors between established reserves. Thus the Vondrozo region (square 816) links the three reserves of Manombo, Kalambatritra and also Pic d’Ivohibe, potentially providing continuity between these reserves. The area surrounding Mitanoka (388) is likewise between Zahamena, Mangerivola and Betampona, three reserves situated in the Middle East. Furthermore, there are several areas that would extend established reserves. These include Maroantsetra (153) in the north-west of Masoala National Park; Mandritsara (191) west of Makira plateau located in the north of Marotandrano special reserve; Ankara (304) next to Kasijy special reserve; Maromandia (80) west of Manongarivo reserve; the part of Bongolava massif around Ambaravaranala (422), in the north-east of Ambohijanahary special reserve; Ankoadava Mahabo next to Andranomena special reserve; Tsintongambarika forest next to Mandena reserve and finally the area next to Fort-Dauphin (961), south of Andohahela reserve. Finally, there are areas located in some bioclimatic, geological or vegetation classes that are not well represented in the current protected areas network of Madagascar, like Ankazotelo (878) which is part of the Mahafaly spiny forest region in the south; and the 122 L. Andriamampianina et al. Montagne des Francais Maromandia Marogaoma Maevatanana Antalaha Maroantsetra Mandritsara Sitampity Ambongo Soanierana Ivongo Ankara Fenerive Est Ambaravaranala Mitanoka Andrangoloaka Ankoadava Ankazomivady Sakaleona Fianarantsoa Vohilava Faraony Ankazotelo Vondrozo 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 northwest that is also diverse in rock types, including alluvial and lakes deposits, sandstone and Mesozoic limestone with Marls, as well as ultrabasic, and metamorphic and igneous basement rocks (Du Puy & Moat 1996); and two areas among the high-plateau, including Ankazomivady (619), east of the Itremo massif. This area is outstanding because of its geological characteristics that include metamorphic and igneous rocks, plus quartzites and marbles (cipolin), two of the rarest still vegetated geological types in Madagascar (Du Puy & Moat 1996). This area presents similar geological characteristics to the region north of Maroantsetra (153), another interesting area with a unique species, Physodeutera umbrosa. The second high-plateau area is the Andrangoloaka forest (497), between the two lakes of Mantasoa and Tsiazompaniry. Although the area around Antananarivo (square 475), the capital of Madagascar, appears to be very rich, it has not been included for a number of reasons. The few remaining semi-natural areas found near Antananarivo are effectively protected because they are treated as sacred. Also, data recorded from this locality may be unreliable because of imprecise labels, since many early collectors lived and acquired specimens in Antananarivo. Therefore, two alternative squares were chosen instead: Andrangoloaka and Ambaravaranala. Many of the priority areas correspond to ‘Classified Forests’ (COEFOR/CI 1993), parts of the ‘domaine forestier national’ but lack legal protection. Some of these sites are also suggested as potentially valuable foci for research and conservation because of their unique geological characters, which may result in unique floral and faunal assemblages. For example, these include the area north of Antongil bay (north of Maroantsetra & Mahalevona), and Ankazomivady, east of Itremo (Du Puy & Moat 1996). Many of these areas, and especially those that could act as corridors between established reserves, were identified as having a high, very high or even exceptional biological or conservation importance in the recommendations of 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 reference Square name Species richness SP A C 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 Current 24 816 281 563 497 422 878 80 388 191 153 253 304 619 737 87 623 948 961 343 292 125 688 Reserves Montagne des Français Vondrozo Maevatanana Ankoadava (Mahabo) Andrangoloaka Ambaravaranala Ankazotelo Maromandia Mitanoka; Namolazana Mandritsara Maroantsetra Sitampiky; Ambongo Ankara; Menavava river Ankazomivady Vohilava-Faraony Marogaoma Sakaleona Tsitongambarika Fort Dauphin Fenerive Soanierana Ivongo Antalaha Fianarantsoa 139 36 23 28 15 31 11 13 27 46 28 57 27 23 23 16 30 21 21 17 27 16 13 29 139 4 4 3 2 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 139 143 147 150 152 153 154 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 FC PSW Y Y Y Y N N N Y Y Y N Y N Y Y Y Y Y N Y Y N N Y N N Y N Y Y Y Y Y Y N N Y Y Y Y Y Y Y Y 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 A map showing priority areas for research is given in Figure 8. The central region around Antananarivo is one area of potential interest. In recent times, the region around Antananarivo has been neglected by biologists since most of the natural vegetation has been destroyed. However, relicts of primary vegetation include the 12 sacred hills around Antananarivo. Other little known areas of the central region that could be of interest include Ankazobe (Ambohitantely), the Bongolava massif (incl. Ambohijanahary), Itremo massif, and forests along the cliffs of Angavo such as Anjozorobe and Andrangoloaka. A census of the current condition of these areas guided by recent satellite maps could yield surprising data. Other poorly sampled western areas of Madagascar are yielding promising results, for example the Sakalava region. Two new species of cicindelids were recently discovered in the middle-west in the surrounds of Kirindy/CFPF and Ankoadava (Cassola & Andriamampianina 1998), and many more species are likely to be discovered in western vegetation types. Additional vegetationally-distinct sampling gaps are the south and south-west around the Mikea forests, and the southern spiny thorn forests around the Mahafaly and the Ivakoany massif. Furthermore, tiger beetles are almost unknown for some habitats such as limestone karst areas (the Tsingy), and lower montane and montane shrublands such as exist at Tsaratanana, Anjanaharibe Sud and Marojejy. All those areas should be surveyed because many beetles and members of other groups now risk extinction before even we know of their existence. Conclusion A total of 176 species of Malagasy cicindelid beetles 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 and taxonomic research are required. Most field studies to date have been carried out within reserves in Madagascar, both for cicindelids and other taxa. Therefore, reserves are in general better sampled than unprotected areas. This is probably the reason for the fairly high level of overall biodiversity apparently covered by the current reserves. Two species have just been discovered and a number of little known species have been rediscovered, outside reserves. However, the distributions of many species are still poorly known (for example, the four species described by Maran (1942) are entirely unknown in terms of distribution). The ranges of many locally endemic species are particularly susceptible to underestimation, because of sampling gaps and also because of their rarity. An example is the species Stenocosmia angusta, thought to be confined to the region of Ampijoroa, Ankarafantsika (square 210) since its discovery in 1965, but collected 480 km away, from the Kirindy/CFPF forest (square 563) in 1997. This study reveals that, for those areas that are relatively well known, the eastern rainforest, its western extension (the Sambirano), and the extreme north of Madagascar (which comprises transitional forest between the humid rainforest and the dry forest) have the richest biodiversity for tiger beetles, a pattern found in many other groups (Lees et al. 1999). Some areas in the centre, the west and the south appear depauperate. Whether this reflects reality or an information gap can only be answered by further fieldwork. Areas of high aggregate endemism occur more patchily all over the island. The extreme north, the north-west and the Biogeography and conservation of Madagascan tiger beetles extreme south appear most outstanding at the generic level, whereas the north eastern rainforest is remarkable at the specific level. This contrast shown here for a Madagascan group is interesting in light of the suggestion 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 current protected areas network (85%). Creation of more reserves would, however, be needed to protect all of the tiger beetles. A set of 23 squares was identified in this study as potential priority areas or foci for conservation action. Rather surprisingly, the high-plateau area surrounding Antananarivo (which may still contain an unexpected number of minute forest relics of value for some taxonomic groups) and remaining natural habitats of this high-plateau was identified as important for more research, which would check poorly-documented historical records for cicindelids in this area. In addition, many areas in the south, west and some high mountains would be a priority for future surveys and bioinventory work. This also applies to many unprotected areas between established reserves. The choice of protected areas depends on many factors beyond raw biodiversity scores for selected taxa. Nevertheless, the present study is a contribution towards improving the information base relevant to conservation in Madagascar. Added to information gathered from other studies, it should help in making better decisions regarding the design, planning and implementation of conservation strategies throughout Madagascar. Acknowledgements Andriamampianina’s field work and museum visits were supported by grants from the Darwin Initiative U.K., the Wildlife Conservation Society and the Durrell Institute of Conservation and Ecology, U.K. In particular, Wildlife Conservation Society (Madagascar) is thanked for the financial and logistical support for this study. Other authors were supported by BBSRC, Leverhulme Foundation and NERC (David Lees). The Natural History Museum in London, the Muséum National d’Histoire Naturelle in Paris, the Parc Botanique et Zoologique de Tsimbazaza in Antananarivo, M. André Peyrieras and M. Fabio Cassola are thanked for making their tiger beetle collection available. We wish to thank Paul Williams for providing the computer program WORLDMAP and assisting with establishment of the Madagascar grid. 125 Fabio Cassola was particularly helpful with the identification of the tiger beetles specimens recently collected. The Ministry of Water and Forests and ANGAP are thanked for their collaboration with research permits. References Andriamampianina, L. (1996) Biogeography of enariine (Melolonthidae) and cicindelid beetles in Madagascar. Unpublished M.Sc. Thesis, University of Kent at Canterbury. ANGAP (1998) Programme Environmental II. Composante Aires Protegées et Ecotourisme: rapport annuel des activitiés 1998, 23 p. Banque Mondiale, USAID, Coopération Suisse, UNESCO, UNDP and WWF (1988) Madagascar: Plan d’action Environnemental. I: Document de synthèse générale et propositions d’orientations. Brown, J.H. and Kodric-Brown, A. (1977) Turnover rates in insular biogeography: effect of immigration on extinction. Ecology 58, 775–87. Carroll, S.S. and Pearson, D.L. (1998) Spatial modeling of butterfly species richness using tiger beetles (Cicindelidae) as a bioindicator taxon. Ecol. Appl. 8, 531–43. Cassola, F. and Andriamampianina, L. (1998) Deux espèces nouvelles de Cicindélides de Madagascar (Coleoptera Cicindelidae) Boll. Soc. Entomol. Ital. 130, 47–54. COEFOR/CI (1993) Répertoire et Carte de Distribution: Domaine Forestier de Madagascar. Direction des Eaux et Forêts, Service des Ressources Forestières, Projet COEFOR (contribution à l’étude des Forêts Classées), et Conservation International, 20 pp. + 1 map. Du Puy, D.J. and Moat, J. (1996) A refined classification of the primary vegetation of Madagascar based on the underlying geology: using GIS to map its distribution and to assess its conservation status. In Proceedings of the International Symposium on the Biogeography of Madagascar (W.R. Lourenço, ed), 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 that lowlands are richer than highlands in endemic and rare species. Biodivers. Conserv. 6, 1137–54. Fjeldså, J. and Lovett, J.C. (1997) Biodiversity and environmental stability. Biodivers. Conserv. 6, 315–23. Foibe Taosaritanin Madagasikara (F.T.M.) (1979–1985) 1 : 500,000 map series. Foibe Taosaritanin Madagasikara (F.T.M.) (1987) Carte Internationale du Monde 1 : 1,000,000. Cartes n◦ SD à SG-38-39. Ganzhorn, J.U., Rakotosamimanana, B., Hannah, L., Hough, J., Iyer, L., Olivieri, S., Rajaobelina, S., Rodstrom, C. and Tilkin, G. (1997) Priorities for biodiversity conservation in Madagascar. Prim. Report 48(1). Green, G.M. and Sussman, R.W. (1990) Deforestation history of the eastern rain forests of Madagascar from satellite images. Science 248, 212–15. Hanski, I. (1982) Dynamics of regional distribution: the core and satellite hypothesis. Oikos 38, 210–21. Horn, W. (1934) Les Cicindélides de Madagascar. Première partie: Catalogue Bibliographique et Synonymique. Mem. Acad. Malgache 20, 7–28. Jeannel, R. (1946) Coléoptères Carabiques de la Région Malgache (Première partie). Faune de l’ Empire Français 6, 104–206. 126 L. Andriamampianina et al. Lees, D.C. (1997) Systematics and biogeography of Madagascan mycalesine butterflies (Lepidoptera: Satyrinae). Ph.D. Thesis, University of London. Lees, D.C., Kremen, C. and Andriampianina, L. (1999) A null model for species richness gradients: bounded range overlap of mycalesine butterflies (Lepidoptera: Satyrinae) and other rainforest endemics in Madagascar. Biol. J. Linn. Soc. 67, 529–84. Lowry, P.P., Schatz, G.E and Phillipson, P.B. (1997) The classification of natural and anthropogenic vegetation in Madagascar. In Natural change and human impact in Madagascar (S.M. Goodman and B.D. Patterson, eds), pp. 93–123. Washington: Smithsonian Institution. Maran, J. (1942) De novis generis Prothyma Hope speciebus formisque Insulae Madagascariensis. Coleoptera-Cicindelidae. Sbornik entom. Odd. Zem. Musea v Praze 20, 63–73. Nelson, R. and Horning, N. (1993) Forest/non-forest classification of Madagascar from AVHRR data: AVHRR-LAC estimates of forest area in Madagascar, 1990. Int. J. Remote Sens. 14, 1463–75. Olsoufieff, G. (1934) Les Cicindélides de Madagascar. Deuxième partie: Essai de Révision systématique et Biologie. Mem. Acad. Malgache 20, 31–71. ONE (1997) Environnement: politique, plan d’action, programme à Madagascar. Brochure de l’Office National de l’Environnement. 35 pp. Pearson, D.L. (1993) Tiger beetle species collected at Ranomafana National Park and Masoala Peninsula, Madagascar. Unpublished report. Pearson, D.L. and Cassola, F. (1992) World-wide species richness patterns of tiger beetles (Coleoptera, Cicindelidae): indicator taxon for biodiversity and conservation Studies. Conserv. Biol. 6, 376–91. Pearson, D.L. and Carroll, S.S. (1998) Global patterns of species richness: spatial models for conservation planning using bioindicator and precipitation data. Conserv. Biol. 12, 809–21. Prendergast, J. R. and Eversham, B. C. (1997) Species richness covariance in higher taxa: empirical tests of the biodiversity indicator concept. Ecography 20, 210–16. Rivalier, E. (1950) Démembrement du Genre Cicindela Linné. (Travail préliminaire limité à la faune paléarctique). Revue Fr. Ent. 12, 217–44. Rivalier, E. (1965) Description d’espèces nouvelles et création d’un Genre nouveau de Cicindelidae Malgaches. Ann. Soc. Ent. Fr. (N.S.) 1, 641–57. Rivalier, E. (1967) Le genre Physodeutera Lacordaire (Col. Cicindelidae). Révision et description d’espèces nouvelles. Ann. Soc. Ent. Fr. (N.S.) 3, 261–96. Rivalier, E. (1970) Le genre Pogonostoma (Col. Cicindelidae). Révision avec description d’espèces nouvelles. Ann. Soc. Ent. Fr. (N.S) 6, 269–338. Rosenzweig, M.L. (1995) Species diversity in space and time. Cambridge: Cambridge University Press. Usher, R.R. (1986) Wildlife conservation evaluation. London: Chapman and Hall. Vane-Wright, R.I., Smith, C.R. and Kitching, I.J. (1994) Systematic assessment of taxic diversity by summation. In Systematics and conservation evaluation. (P.L. Forey, C.J. Humphries and R.I. Vane-Wright, eds), pp. 309–26. Oxford: Oxford University Press. Viette, P.D. (1991) Principales localités où des insectes ont été recueillis à Madagascar. Faune de Madagascar. Suppl. 2, 88 pp. Privately published by the author. World Conservation Monitoring Center (WCMC) (1991) Guide de la Diversité Biologique de Madagascar. Map, World Conservation Monitoring Centre, Cambridge. Williams, P.H. (1994) Using WORLDMAP, priority areas for biodiversity. Program and manual. Version 3.1 London (distributed by the author). Williams, P.H. and Gaston, K.J. (1994) Measuring more of biodiversity: can higher-taxon richness predict wholesale species richness? Biol. Conserv. 67, 211–17. Appendix List of species of Cicindelidae included in the database. Species that were re-observed during recent fieldworks are marked with (∗), species for which there is no distributional data available are marked with (?). Subfamily 1: Cicindelinae Sloane, 1906 Genus 1: Hipparidium Jeannel, 1946 H. equestre Dejean, 1826∗ H. perroti Fairmaire, 1897 H. clavator Jeannel, 1946∗ H. sahy Alluaud, 1902 H. osa Alluaud, 1902 H. albo-sinuatum Olsoufieff, 1934 Genus 2: Myriochile Motschulsky, 1862 M. melancholica Fabricius, 1798∗ Genus 3: Lophyra Motschoulsky, 1862 L. abbreviata Klug, 1832∗ L. quadraticollis Chaudoir, 1835 L. tetradia Fairmaire, 1899∗ L. vittula Rivalier, 1951 Genus 4: Habrodera Motschulsky, 1862 H. ovas Bates, 1878∗ H. truncatilabris Fairmaire, 1897∗ Genus 5: Chaetodera Jeannel, 1946 C. maheva Kunckel, 1887∗ C. andriana Alluaud, 1900∗ C. perrieri Fairmaire, 1897 ∗ C. antatsima Alluaud, 1902∗ Genus 6: Lophyridia Jeannel, 1946 L. cristipennis Walther Horn, 1905∗ Genus 7: Cylindera Westwood, 1831 C. fallax Coquerel, 1851∗ C. umbratilis Fairmaire, 1903 C. constricticollis Walther Horn, 1913 C. sakalava Cassola & Andriamampianina, 1998∗ C. zaza Alluaud, 1902∗ Biogeography and conservation of Madagascan tiger beetles Genus 8: Cicindelina Jeannel, 1946 C. oculata Chaudoir, 1843∗ Genus 9: Chaetotaxis Jeannel, 1946 C. rugicollis Fairmaire, 1871∗ C. cicindeloides Walter Horn, 1905 C. descarpentriesi Deuve, 1987∗ C. soalalae Fairmaire, 1903 ∗ C. ankarahitrae Jeannel, 1946 C. serieguttata Walter Horn, 1934 C. macropus Chaudoir, 1865 C. grandidieri Kunckel, 1887∗ C. semi-confluens Rivalier, 1965 C. leptographa Rivalier, 1965 Genus 10: Ambalia Jeannel, 1946 A. aberrans Fairmaire, 1871∗ A. satura Rivalier, 1965 Genus 11: Calyptoglossa Jeannel, 1946 C. frontalis Audouin et Brullé, 1839∗ Genus 12: Peridexia Chaudoir, 1860 P. fulvipes Dejean, 1831∗ P. hilaris Fairmaire, 1883∗ Genus 13: Prothyma Hope, 1838 P. radama Kunckel, 1887∗ Genus 14: Physodeutera Lacordaire, 1843 P. adonis Castelnau, 1835 P. intermedia Rivalier, 1967 P. parcepunctata Jeannel, 1946 P. uncifera Jeannel, 1946 P. lateralis Olsoufieff, 1934 P. bellula Fleutiaux, 1886 P. trimaculata Fleutiaux, 1889∗ P. sobrina Rivalier, 1967 P. debilis Rivalier, 1967 P. uniguttata Fairmaire, 1871 P. komorousi Maran, 1942 (?) P. fairmairei Walter Horn, 1899∗ P. hajni Maran, 1942 (?) P. rufosignata Audouin et Brullé, 1839∗ P. mocquerysi Fleutiaux, 1899∗ P. punctum Rivalier, 1951∗ P. umbrosa Rivalier, 1967 P. cyanea Audouin et Brullé, 1839 P. minima Walter Horn, 1893 P. madari Maran, 1942 (?) P. vadoni Rivalier, 1951 P. rubescens Jeannel, 1946 P. pseudorubescens Deuve, 1987 P. perroti Rivalier, 1967 P. gigantea Walter Horn, 1913 P. pokornyi Maran, 1942 (?) P. megalommoı̈des Walter Horn, 1896 P. belalonensis Deuve, 1987 P. flagellicorne Walter Horn, 1897∗ P. pseudo-trimaculata Walter Horn, 1934 P. andriai Rivalier, 1965 P. dorri Fleutiaux, 1899 P. rectipenis Walter Horn, 1934 P. centropunctata Walter Horn, 1934 P. biguttula Fairmaire, 1903 P. dubia Maran, 1942 (?) P. marginemaculata Walter Horn, 1934 P. peyrierasi Rivalier, 1967 P. rectolabialis Walter Horn, 1913 P. punctipenne Fairmaire, 1903 P. virgulata Fairmaire, 1904 P. bucephala Walter Horn, 1900 P. sikorae Walter Horn, 1896∗ P. subtilivelutina Walter Horn, 1934b P. viridi-cyanea Audouin et brullé, 1839∗ P. maximum Fleutiaux, 1899 P. janthina Fairmaire, 1903 P. natalia Walter Horn, 1934 P. catalai Jeannel, 1946 P. alluaudi Fleutiaux, 1903∗ P. tricolorata Walter Horn, 1934 P. perrieri Rivalier, 1967 Genus 15: Walterhornia Olsoufieff, 1934 W. speculifera Walter Horn, 1934∗ Genus 16: Stenocosmia Rivalier, 1965 S. angusta Rivalier, 1965∗ S. tenuicollis Fairmaire, 1904 Subfamily 2: Collyrinae Csiki, 1906 Genus 17: Pogonostoma Klug, 1835 P. cylindricum Fleutiaux, 1899∗ P. brevicorne Walter Horn, 1898 P. vestitum Fairmaire, 1900∗ P. violaceum Fleutiaux, 1902 P. septentrionale Fleutiaux, 1903 P. cyanescens Klug, 1835∗ P. caerulea Castelnau & Gory, 1837∗ P. violaceolevigata Walter Horn, 1927 P. sambiranense Rivalier, 1965 P. affine Horn, 1893 P. chalybaeum Klug, 1835∗ P. spinipennis Castelnau et Gory, 1837 P. rugosoglabra Walter Horn, 1923 P. propinquum Rivalier, 1970∗ 127 128 L. Andriamampianina et al. P. globicolle Rivalier, 1965 P. perroti Rivalier, 1970 P. malleatum Rivalier, 1970 P. peyrierasi Rivalier, 1970 P. densepunctatum Rivalier, 1970 P. subtilis Walter Horn, 1904 P. hamulipenis Walter Horn, 1934 P. tortipenis Walter Horn, 1934 P. impressum Rivalier, 1970 P. externespinosa Walter Horn, 1934 P. elegans Brullé, 1834∗ P. abadiei Rivalier, 1965 P. brullei Castelnau & Gory, 1837 P. alluaudi Walter Horn, 1898 P. rufidens Rivalier, 1970 P. atrorotundata Walter Horn, 1934 P. sudiferum Rivalier, 1965 P. subtiligrossa Walter Horn, 1934∗ P. litigiosum Rivalier, 1970 P. ankaranense Deuve, 1986 P. srnkai Walter Horn, 1893 P. gibbosum Rivalier, 1970 P. meridionale Fleutiaux, 1899 P. sikorae Walter Horn, 1894∗ P. flavomaxillaris Cassola & Andriamampianina, 1998∗ P. pseudo-minimum Walter Horn, 1934 P. flavopalpale Jeannel, 1946 P. infimum Rivalier, 1970 P. gladiator Walter Horn, 1934 P. phalangioı̈de Rivalier, 1970 P. levigatum Walter Horn, 1908 P. comptum Rivalier, 1970 P. surdum Rivalier, 1970 P. sericeum Klug, 1835 P. perrieri Fairmaire, 1900 P. kraatzi Walter Horn, 1894 P. ovicolle Walter Horn, 1893 P. microtuberculatum Walter Horn, 1934 P. longicolle Jeannel, 1946 P. anthracina Castelnau et Gory, 1837 P. maculicorne Walter Horn, 1934 P. humbloti Rivalier, 1970 P. excisoclavipenis Walter Horn, 1934 P. basidilatatum Walter Horn, 1909 P. pusilla Castelnau et Gory, 1937 P. laportei Walter Horn, 1900∗ P. inerme Jeannel, 1946 P. flavomaculatum Walter Horn, 1892 P. beananae Rivalier, 1963 P. moestum Rivalier, 1970 P. schaumi Walter Horn, 1893∗ P. mathiauxi Jeannel, 1946 P. fleutiauxi Walter Horn, 1905∗ P. basale Fleutiaux, 1899 P. globulithorax Jeannel, 1946 P. parallelum Walter Horn, 1909 P. geniculatum Jeannel, 1946 P. horni Fleutiaux, 1899 P. rugosiceps Rivalier, 1970 P. nigricans Klug, 1835 P. vadoni Jeannel, 1946 P. pallipes Rivalier, 1970 P. mocquerysi Fleutiaux, 1899 P. delphinense Jeannel, 1946 P. minimum Fleutiaux, 1899 P. sicardi Walter Horn, 1927 P. parvulum Rivalier, 1970