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.
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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