Taxic Richness Patterns and Conservation Evaluation of Madagascan Tiger Beetles (Coleoptera: Cicindelidae)
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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,† 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. 5Facult´e des Sciences, Universit´e 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, occur- ring 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 local- ities. When a given locality was too large to fit into one Perinet Analamazaotra Kirindy/ cell or when locality