Corresponding author: Michaël Manuel (michael.manuel@upmc.fr)
Academic editor: Mariano Michat
Water beetles of the families
Ramahandrison AT, Rakouth B, Manuel M (2022) The aquatic Adephaga of the Makay, central-western Madagascar, with description of two new diving beetle species (Coleoptera, Gyrinidae, Haliplidae, Noteridae, Dytiscidae). ZooKeys 1127: 1–60.
The Makay massif, located in the central-western part of Madagascar (Fig.
Distribution of sampling sites in the study area
The vegetation is adapted to these contrasting life conditions. At places a typical dense rainforest flourishes, highly similar to that usually found in the eastern part of Madagascar, with the presence of remarkable species such as
It was not until 2010 that scientific missions to explore and describe the biodiversity of the Makay started (
Currently four families and 231 species of aquatic
We present here the results of three sampling campaigns targeting aquatic
The study specimens are deposited in the last author’s research collection (
Sampling sites are numbered in chronological order of (first) visit from MAK-1 to MAK-62. They are mapped on Fig.
Three field campaigns were conducted. The first two campaigns (2.–9.VI.2016 and 26.VII.–28.VIII.2017) were conducted in the south-central part of the Makay. In 2016, the area around the Menapanda and the Andranomanintsy rivers was explored (sites MAK-3 to MAK-17, Fig.
All samplings were performed in situ by hand netting using a GB-net professional hand net (NHBS, Totnes, Devon, UK) (25 cm frame; depth of net bag 50 cm; mesh 1 mm), except at site MAK-22 (light trap).
All sites located in the boxes within the map of Fig.
Sites were furthermore categorised according to their vegetation context as determined from field observation completed by inspection of satellite images (Google Earth Pro 7.3.) as “forested”, “semi-forested” or “non-forested”. The context was considered “semi-forested” when a sampling site was located in open or semi-open situation but close to forest edge, or at the bottom of narrow canyons without a proper gallery forest but with a certain density of trees nevertheless present.
In the sampling data given below, the letter between parentheses after the sampling code indicates the vegetation context:
Representative habitats of aquatic
Specimens were morphologically identified to species level by MM (when necessary, with study of dissected genitalia) using the relevant taxonomic literature (reviewed in
For illustration of newly described species, photographs of habitus were made with an Olympus SZX12 trinocular stereomicroscope (Tokyo, Japan) using a Spot FLEX Color Pixel Shift 64 Mp camera (Diagnostic Instruments Inc., Sterling Heights, MI, USA) with SPOT BASIC software (
Label data of type material are given as written in quotation marks, with separate label lines indicated by a slash (/) and separate labels by a double slash (//). Authors’ additional remarks are provided in square brackets.
Relative frequency of occurrence (
Interpolation-extrapolation sampling curves (
In order to compare the groups of observations with each other and to quantify similarity/dissimilarity in species composition, Jaccard (based on occurrence data) and Bray-Curtis (based on abundance data) dissimilarity indices were calculated (with the R software). The data were standardised prior to computation of Bray-Curtis indices. The Jaccard dissimilarity index between two sets of objects A and B is equal to 1 - J(A,B) where J(A,B) = |A∩B| / |A∪B|. For the formula of the Bray-Curtis dissimilarity index see
Madagascar.
1 ♂, 2 ♀♀: MAK-5B; 1 ♂, 1 ♀: MAK-5D; 2 ♂♂: MAK-13; 2 ♂♂, 3 ♀♀: MAK-14B; 2 ♂♂, 1 ♀: MAK _24; 3 ♂♂, 8 ♀ ♀: MAK-27; 1 ♀: MAK-30; 1 ♀: MAK-35C; 2 ♂♂, 2 ♀♀: MAK-37B; 2 ♀♀: MAK-40A; 2 ♂♂, 3 ♀♀: MAK-40B; 3 ♂♂, 3 ♀♀: MAK-48; 1 ♂, 2 ♀♀: MAK-55; 1 ♂, 1 ♀: MAK-59A; ; 1 ♂, 3 ♀♀: MAK-59C.
Madagascar, widespread (
Collected only in inner Makay, in permanent lotic habitats (rivers and streams) with sandy bottom (in a few sites substrate was more rocky) and with clear water, in forested or semi-forested environmental context, with little or no anthropogenic disturbance.
Tibet (erroneous locality?).
1 ♂, 4 ♀♀: MAK-27; 3 ♂♂, 4 ♀♀: MAK-37B; 1 ♂, 1 ♀: MAK-40A; 1 ♂: MAK-40B; 1 ♀: MAK-48; 1 ♂, 5 ♀♀: MAK-52; 5 ♂♂, 9 ♀♀: MAK-55; 1 ♀: MAK-58.
Madagascar, widespread (
Same as
Madagascar, Diego Suarez (Antsiranana), Isokitra.
1 ♂: MAK-15; 2 ♂♂, 3 ♀♀: MAK-36A; 7 ♂♂, 6 ♀♀: MAK-37B; 1 ♀: MAK-40B; 1 ♂, 3 ♀♀: MAK-48.
Madagascar. Previously recorded only from the northern part of the island (
Same as the two preceding species, with a stronger preference for forested and undisturbed habitats.
Madagascar, Antananarivo, Ambodinandohalo Lake.
1 ♀: MAK-19; 1 ♂: MAK-41.
Madagascar, widespread (
This species was only found at two sampling sites, both peripheral. One was a large puddle partially sheltered by trees, with water slowly flowing and with abundant rice straw debris, on a dirty road between two rice fields, and the other was a small puddle in open situation on the sandy bank of a river, with
Madagascar.
1 ♂: MAK-2.
Madagascar, widespread (
Madagascar, Fort-Dauphin, Ankara.
2 ♂♂: MAK-2; 1 ♀: MAK-19.
Zaire (Democratic Republic of the Congo), Madagascar (
La Réunion; Madagascar.
3 ♀: MAK-1A; 1 ♀: MAK-2; 1 ♀: MAK-11A; 52 ♂♂, 39 ♀♀: MAK-19; 1 ♂: MAK-20; 1 ♂, 2 ♀♀: MAK-38B; 1 ♂, 3 ♀♀: MAK-40A; 2 ♂♂, 4 ♀♀: MAK-41; 4♂♂, 8 ♀♀: MAK-42; 17 ♂♂, 11 ♀♀: MAK-60; ; 1 ♂: MAK-61.
Madagascar and Mascarene Islands; widespread and common in Madagascar (
1 ♀: MAK-20; 1 ♂, 3 ♀♀: MAK-60; 3 ♀♀: MAK-61.
This species is very similar to
Madagascar. In addition to the specimens from the Makay, we also have specimens from Namoroka (north-eastern part of the island).
Species collected in permanent lentic environments in open peripheral sites, with clay bottom, clear water and presence of vegetation. One of the collecting sites was a rice field.
Madagascar.
2 ♂♂: MAK-2; 1 ♀: MAK-11A; 8 ♂♂, 16 ♀♀: MAK-19; 1 ♂, 2 ♀♀: MAK-21; 20 exs.: MAK-23; 1 ♂: MAK-38B; 2 ♂♂, 1 ♀: MAK-42; 8 ♂♂, 10 ♀♀: MAK-43; 1 ♂, 4 ♀♀: MAK-44A; 1 ♀: MAK-44C; 9 ♂♂, 17 ♀♀: MAK-60; 2 еxs.: MAK-61.
Africa (Angola, Guinea, Guinea-Bissau, Mali), Madagascar, and Mascarene Islands (
3 ♂♂, 2 ♀♀: MAK-43; 2 ♀♀: MAK-44C; 4 ♂♂, 1 ♀: MAK-56.
This species is smaller than
Madagascar, widespread but not very common.
This species was sampled at three sites in inner Makay, in slowly flowing lotic habitats and a dead river arm, in forested or semi-forested contexts without anthropogenic disturbance. These biotopes had sandy bottoms with moderate abundance of plant debris. Two of the sites were surrounded by a well-developed hygrophilous vegetation and contained cyanobacteria.
Madagascar, Mahajanga, Ambohimanatrika.
3 ♂♂, 2 ♀♀: MAK-19; 1 ♂, 1 ♀: MAK-41.
Madagascar (
This species was collected at two peripheral sites in lentic habitats in areas with intense anthropogenic pressure: large puddle, with water slowly flowing and with abundant rice straw debris, on a dirty road between two rice fields; and puddle on the sandy banks of the Mangoky River.
Madagascar.
1 ♀: MAK-21; 1 ♂: MAK-42.
Madagascar; widespread and common in lowlands (
This species was collected at two peripheral sites located in open areas, in temporary lentic habitats without water renewal, and with vegetation.
Madagascar, Isalo National Parc.
1 ♂, 1 ♀: MAK-6; 3 ♂♂: MAK-7; 1 ♀: MAK-14A; 7 ♂♂, 13 ♀♀: MAK-30; 18 ♂♂, 17 ♀♀: MAK-32; 2 ♂♂, 3 ♀♀: MAK-34A; 2 ♂♂, 1 ♀: MAK-34B; 3 ♀♀: MAK-35A; 1 ♂: MAK-39A; 42 ♂♂, 25 ♀♀: MAK-45; 10 ♂♂, 7 ♀♀: MAK-46; 1 ♀: MAK-47; 26 ♂♂, 44 ♀♀: MAK-49; 10 ♂♂, 3 ♀♀: MAK-50; 1 ♀: MAK-52; 11 ♂♂, 5 ♀♀: MAK-53; 1 ♂, 2 ♀♀: MAK-54A; 4 ♂♂, 5 ♀♀: MAK-54B; 48 ♂♂, 57 ♀♀: MAK-59B; 5 ♂♂, 2 ♀♀: MAK-59C.
Madagascar; previously known only from the sandstone massif of Isalo (
Madagascar, Andobo, Antsingy forest.
4 ♀: MAK-12A; 2 ♀♀: MAK-33; 1 ♂: MAK-57.
Madagascar (
This species was collected in lentic habitats (springs and small puddles), with clay or sandy-clay bottom and a lot of plant debris, located in forested or semi-forested areas and relatively unaffected by human disturbances.
Madagascar, Senegal.
1 ♂: MAK-11A; 1 ♂: MAK-11B; 30 ♂♂, 41 ♀♀: MAK-12A; 1 ♂: MAK-12B; 5 ♂♂, 1 ♀: MAK-12C; 2 ♂♂, 2 ♀♀: MAK-33; 2 ♂♂, 4 ♀♀: MAK-44A; 8 ♂♂, 22 ♀♀: MAK-44B; 23 ♂♂, 22 ♀♀: MAK-44C; 3 ♂♂, 8 ♀♀: MAK-56; 21 ♂♂, 16 ♀♀: MAK-57; 1 ♂: MAK-58.
Continental Africa south from Egypt and Sahara, Madagascar (
Madagascar, Antsiranana, Montagne d’Ambre, Ambohitra National Park.
6 ♂♂, 3 ♀♀: MAK-3; 4 ♂♂, 7 ♀♀: MAK-4; 1 ♂: MAK-7; 1 ♀: MAK-8; 6 ♂♂, 10 ♀♀: MAK-10; 1 ♀: MAK-11B; 2 ♂♂, 6 ♀♀: MAK-12A; 2 ♀♀: MAK-12B; 2 ♀♀: MAK-16; 1 ♀: MAK-25A; 1 ♂: MAK-25B; 3 ♂♂, 5 ♀♀: MAK-28; 1 ♂: MAK-29; 1 ♂, 1 ♀: MAK-33; 1 ♀: MAK-34B; 2 ♂♂, 2 ♀♀: MAK-35A; 4 ♂♂: MAK-38A; 1 ♂, 5 ♀♀: MAK-39A; 6 ♂♂, 5 ♀♀: MAK-39B; 1 ♀: MAK-45; 2 ♂♂: MAK-50.
Madagascar, widespread (
Mayotte.
1 ♀: MAK-44B; 2 ♂♂, 2 ♀♀: MAK-44C; 1 ♀: MAK-56; 2 ♀♀: MAK-60.
Madagascar (widespread), Mayotte (
This species has been captured in a few inner massif sites all situated in northern Makay: a puddle, a blind channel, a small stream (these sites in forest) and an open marsh. These habitats had slightly turbid water and a mineral bottom ranging from clay to sand with moderate quantity of plant debris, and no vegetation.
Madagascar, Toliara province, Malaimbandy municipality, Makay massif (northern part), ca. 20 km WSW Tsimazava, ca.
This species belongs to the
Body elongated and parallel-sided (Fig.
Habitus in dorsal view. Scale bars: 2 mm
Elytra with six discal and one submarginal striae. Stria I very weakly impressed. Striae I, V, and VI abbreviated anteriorly. Submarginal stria very weakly impressed, starting slightly before elytron midlength. Head, pronotum and elytra with fine reticulation and fine punctation. Posterolateral region of pronotum with few short and weakly impressed strioles.
Ventral side rufo-testaceous, slightly darker laterally on metacoxal plate and on abdominal ventrites. Metacoxal plates with sparse and very fine short strioles; visible abdominal ventrites I-III with denser and longer very fine strioles. Prosternal process rather short and broad, with blunt apex. Metacoxal lines rather long, ending anteriorly at quite small distance from posterior margin of metaventrite, diverging anteriorly.
Appendages: Antennae, palps and legs testaceous. Antennae particularly long (Fig.
Median lobe and parameres as in Fig.
Male genitalia. Scale bars: 200 µm
This species is dedicated to the Malavergne family (Dominique, Catherine, Clémence, Jacques, and Laurence, Marie-José) in recognition of their constant help and support to the first author during his PhD thesis. The species epithet is a name in the genitive plural.
So far known only from northern Makay in Madagascar.
The external morphology of this species (very narrow and parallel habitus, broad pronotum, pale colour, long antennae) suggests that it might be a semi-subterranean species. The habitat where the single specimen was found (MAK-44C) was a blind channel connected to River Sakapaly, in northern inner Makay. There was no apparent water flow but the bottom was covered with conspicuous orange masses of iron bacteria, which might be an indication of slow water seepage from underground. The substratum was sandy with moderate amount of decaying vegetal material and the water was red-brown coloured. This water body was fully shaded under trees in forest. There was no vegetation in the water but the surrounding forest floor displayed a typical hygrophilic vegetation of
Madagascar, Toliara province, Malaimbandy municipality, Makay massif (northern part), ca. 21 km W of Tsimazava, ca.
This species belongs to the
Body shape elongate oval, with sides subparallel (Fig.
Elytra with ten well-impressed discal and one submarginal striae. Stria IX abbreviated anteriorly. Striae I and II diverging anteriorly. Submarginal stria starting slightly before elytron midlength, fragmented anteriorly. Head, pronotum and elytra with fine reticulation and fine punctation. Whole surface of pronotum with rather dense, short and fine strioles, in medial disk region strioles even finer.
Ventral side uniformly rufo-testaceous. Metacoxal plates with moderately impressed short strioles; visible abdominal ventrites I-III with denser and longer very fine strioles. Prosternal process short and broad, with rounded apex. Metacoxal lines short, ending anteriorly at large distance from posterior margin of metaventrite, moderately diverging anteriorly.
Appendages: Antennae, palps, forelegs and midlegs testaceous, hindlegs rufo-testaceous. First three pro- and mesotarsomeres widened and ventrally equipped with suction cups; number of suction cups per article (I-III) 7:4:4 on both pro- and mesotarsus. Protibia at base shortly narrow, with shallow protuberance along ventral margin, distally broadened. Pro- and mesotarsal claws unmodified.
Median lobe and parameres as in Fig.
Holotype:
In the male paratype, strioles on pronotum are longer and more deeply impressed than in the holotype, and the metacoxal lines are slightly longer. In the female paratype, the elytral stria V is slightly abbreviated anteriorly.
The species name literally means “son of the rock” in Malagasy. It is an invariable name standing in apposition.
So far known only from northern Makay in Madagascar.
This species was collected at two sites located in two nearby canyons in northern inner Makay. Two specimens were sampled at site MAK-46, an isolated pool (~ 1 m × 3 m) on the bottom of a narrow and dark canyon, and one specimen at site MAK-50, a stagnant temporary pond (~ 3 m × 7 m) situated in a wider canyon and in a more open environment. Both habitats were characterised by sandy bottom with some plant debris, absence of visible inflow / outflow, somewhat turbid water and no vegetation. Other species of aquatic
Madagascar, Maevatanana.
1 ♂, 2 ♀: MAK-3; 2 ♂♂, 8 ♀♀: MAK-10; 1 ♂, 1 ♀: MAK-11B; 2 ♂♂, 2 ♀♀: MAK-12A; 1 ♂: MAK-15; 1 ♂, 1 ♀: MAK-16; 9 ♂♂, 1 ♀: MAK-25A; 3 ♂♂, 1 ♀: MAK-29; 1 ♂: MAK-35A; 1 ♀: MAK-37A; 4 ♂♂, 2 ♀♀: MAK-39A; 1 ♂: MAK-40B.
Madagascar (distribution within the island poorly known) (
Madagascar.
1 ex.: MAK-1A; 1 ♂, 1 ♂: MAK-2; 1 ♂, 3 ♀♀: MAK-19; 4 ♂♂, 1 еx.: MAK-20; 1 ♀: MAK-23; 1 ♀: MAK-41.
Madagascar (widespread and common) (
Madagascar.
1 ♀: MAK-50.
Madagascar (widespread but localised) (
This species was captured only once, in northern inner Makay, in a large temporary pond located in a shallow open area on sand. This single capture does not reflect the usual habitat preferences of this species. According to our observations in Isalo and Ankarafantsika, this is a lotic species (an exceptional ecology for the genus) inhabiting the margins of streams and rivers with some vegetation and / or tree roots and / or plant debris, in well-preserved forested or semi-forested environments.
Madagascar.
1 ♂: MAK-2; 2 ♀♀: MAK-19; 1 ♂: MAK-41.
From sub-Saharan Africa to Australia through tropical Asia and the Oriental region (
India.
1 ♂, 1 ♀: MAK-11B.
Southern half of the Palearctic region, Africa, Oriental region (
Madagascar.
1 ♂: MAK-11A; 1 ♀: MAK-11B; 1 ♂, 1 ♀: MAK-12A; 1 ♂: MAK-14A; 1 ♂: MAK-18; 8 ♂♂, 6 ♀♀: MAK-19; 1 ♂: MAK-23.
Whole tropical Africa to Arabia, Madagascar (where it is widespread and common) (
Madagascar.
1 ♂: MAK-11B.
Sub-Saharan Africa, Mauritius, Madagascar (
Madagascar.
1 ♂: MAK-52.
Madagascar, widespread (
This species was collected once, in northern inner Makay, in a calm pool (~ 3 m × 7 m) on the bottom of a canyon, with inflow and outflow from the nearby Ampasimaiky River. This pool was rather deep (> 1 m), with bottom of sand covered with a thin layer of clay, almost without vegetal detritus, with moderately turbid water and no vegetation. The environment was semi-forested and the pond was surrounded by
South Africa, Western Cape, Cape of Good Hope.
1 ♀: MAK-1A; 1 ♂: MAK-2; 13 ♂♂, 7 ♀♀: MAK-19; 1 ♂, 3 ♀♀: MAK-23; 2 ♂♂: MAK-40A; 1 ♂, 6 ♀♀: MAK-61; 1 ♂: MAK-62.
Sub-Saharan Africa, Madagascar (
Madagascar, Mascarene Islands (
1 ♀: MAK- 8; 2 ♀: MAK-50; 2 ♀♀: MAK-53; 1 ♂: MAK-54A; 2 ♀♀: MAK-59C.
Mauritius, La Réunion, Madagascar. In Madagascar, widespread.
This species was collected only in inner Makay, in well-preserved forested or semi-forested areas. The habitats were isolated pools and very slowly flowing streams, with clear or slightly turbid water, sandy bottom (sometimes with stones), with moderate amount of plant debris and no vegetation.
Madagascar, Antsiranana,
1 ♀: MAK-1A; 1 ♂: MAK-2; 2 ♀♀: MAK-19; 17 еxs.: MAK-42; 1 еx.: MAK-60; 1 еx.: MAK-61.
Madagascar, widespread (
Madagascar, South inner part.
1 ex.: MAK-1A; 4 ♂♂, 1 ex.: MAK-2; 1 ♂: MAK-17; 82 exs.: MAK-18; 4 ♂♂, 25 exs.: MAK-19; 3 exs.: MAK-60; 1 ex.: MAK-61.
Madagascar, widespread (
Madagascar, Iharanandriana mountain.
3 ♂♂, 3 ♀♀: MAK-2; 1 ♂: MAK-61.
Madagascar, widespread (
Madagascar, Bas Mangoky agricultural station.
1 ♂: MAK-20.
Madagascar, widespread but not common (
A single specimen of this species was taken in a rice field on the banks of the Mangoky River (a peripheral site), at shallow depth. The bottom was composed of sand and clay with some plant debris; there was no visible inlet or outlet, the water was clear and there was a moderate presence of green algae. The environment was open with no trees in the surroundings.
1 ♂: MAK-2.
Madagascar (known to us only from site MAK-2).
This is an undescribed species, close to
Madagascar, Antsiranana.
13 ♂♂, 20 ♀: MAK-1A; 1 ♂, 1 ♀: MAK-4; 5 ♂♂: MAK-61.
Seychelles, Madagascar (
Madagascar, Antsiranana.
6 ♂♂, 3 ♀: MAK-1A; 1 ♂, 2 ♀: MAK-2; 6 ♂♂, 3 ♀♀: MAK-4; 2 ♂♂, 10 ♀♀: MAK-17; 1 ♂: MAK-18; 4 ♂♂, 3 ♀♀: MAK-19; 4 ♂♂, 1 ♀: MAK-61.
Sub-Saharan Africa, Mauritius, La Réunion, Madagascar (
Eastern part of Madagascar.
1 ♂: MAK-17.
Madagascar, common in the Central Highlands (
This species seems very rare in the Makay since only one specimen was found, in an inner massif site located in a semi-forested environment. The habitat was a small, isolated, sun-exposed puddle on rock mass, on the bank of a river. The bottom consisted of sandstone, sand and clay without organic debris, the water was clear and there was no vegetation.
Mascarene Islands, Mauritius, Curepipe.
1 ♂: MAK-16.
Mauritius, La Réunion, Comoros, Madagascar (
This species was sampled at a single inner massif site, a small pond, partly shaded, at the edge of a gallery forest close to River Andranomanintsy. The water was slowly renewed from the nearby river, the bottom was sandy with moderate plant debris and the water was clear. This small water body was filled in with subaquatic
Madagascar, Imanombo.
4 ♂♂, 2 ♀♀: MAK-1A; 3 ♂♂, 11 ♀♀: MAK-1B; 12 ♂♂, 26 ♀♀: MAK-1C; 2 ♂♂: MAK-20; 2 ♂♂: MAK-22.
Madagascar, western and southern parts of the island (
5 ♂♂, 2 ♀♀: MAK-2; 25 ♂♂, 22 ♀♀: MAK-3; 8 ♂♂, 4 ♀♀: MAK-4; 3 ♂♂, 1 ♀: MAK-5A; ; 8 exs.: MAK-5D; 2 ♀♀: MAK-8; 29 ♂♂, 14 ♀♀: MAK-14A; 5 ♂♂, 5 ♀♀: MAK-16; 24 exs.: MAK-28; 24 exs.: MAK-29; 3 exs.: MAK-36B; 91 exs.: MAK-37A; 3 exs: MAK-38B; 53 exs.: MAK-40A; 1 ex.: MAK-43; 1 ex.: MAK-44C; 2 exs.: MAK-45; 28 exs.: MAK-47; 5 exs.: MAK-49; 31 exs.: MAK-50; 15 exs.: MAK-51; 37 exs.: MAK-52; 10 exs.: MAK-53; 2 exs.: MAK-54B; 6 exs.: MAK-58; 39 exs.: MAK-59A; 25 exs.: MAK-59C.
This is a probably undescribed species close to
Madagascar. The exact distribution of this species within the island remains to be established in the context of the upcoming revision, but it is not endemic to the Makay (sampled by us notably in the Isalo Massif).
1 ♂: MAK-5A; 2 ♂♂: MAK-5D; 1 ♂, 1 ♂: MAK-14A; 1 ♂, 1 ♀: MAK-29; 1 ♀: MAK-45; 8 ♂♂, 4 ♀♀: MAK-50; 1 ♂, 4 ♀♀: MAK-51; 57 ♂♂, 37 ♀♀: MAK-52; 1 ♀: MAK-59C.
Madagascar. So far endemic to the Makay massif.
This is an undescribed species, rather large for the genus, and very close to another undescribed species which lives in the Isalo massif.
1 ♀: MAK-2; 1 ♂: MAK-61.
Madagascar (widespread).
This species corresponds to
Same as
Madagascar, Maroantsetra.
1 ♂, 1 ♀: MAK-19.
Madagascar (
This species was collected only once, in a peripheral site located in a semi-open area impacted by human activities. The habitat was a large puddle partially sheltered by trees, with water slowly flowing and with abundant rice straw debris, on a dirty road between two rice fields.
Madagascar, Antsiranana.
1 ♀: MAK-1A; 2 ♀♀: MAK-2; 1 ♀: MAK-18; 4 ♂♂, 2 ♀♀: MAK-19; 4 exs.: MAK-60.
Madagascar, widespread in lowlands (
Madagascar, Sainte Marie Island.
5 exs.: MAK-1A; 2 exs.: MAK-1C; 4 exs.: MAK-19; 6 exs.: MAK-20; 4 exs.: MAK-41; 1 ex.: MAK-42; 1 ex.: MAK-60; 3 exs.: MAK-62.
Madagascar, widespread and common (
South-East Asia (Indian continent).
1 ♂, 1 ♀: MAK-1A; 6 ♂♂, 6 ♀♀: MAK-2; 1 ♀: MAK-3; 2 ♂♂, 1 ♀: MAK-19; 1 ♂, 2 ♀♀: MAK-21; 1 ♂: MAK-23; 1 ♀: MAK-44C; 1 ♂, 2 ♀♀: MAK-60; 1 ♂, 1 ♀: MAK-61; 1 ♂: MAK-62.
Sub-Saharan Africa, Madagascar, Seychelles, Turkey, Egypt, Arabian Peninsula, south-eastern Palearctic region from India to south Japan, Oriental region (
Zambia (Congo Belge), Musosa.
1 ♂: MAK-2; 1 ♂, 1 ♀: MAK-42.
Sudan, Democratic Republic of the Congo, Madagascar (
Eastern part of Madagascar
1 ♂, 1 ♀: MAK-42; 1 ♂: MAK-60; 1 ♂, 1 ♀: MAK-61.
Madagascar, widespread (
This species was collected at two peripheral and one inner massif sites. The habitats were located in open areas and were lentic water bodies (with or without slow water circulation) with sand-clay bottom and with plant debris, with water clear to moderately turbid, with discontinuous marginal belts of helophytes and at one site with filamentous green algae.
Eastern part of Madagascar.
1 ♂: MAK-2; 1 ♀: MAK-42.
Madagascar, widespread (
Same as
Zambia, Luangwa valley, Chibembe.
1 ♂: MAK-2.
Zambia, South Africa (
Madagascar, Antananarivo, Ikopa River.
3 ♂♂, 2 ♀♀: MAK-2; 3 ♂♂, 1 ♀: MAK-62.
Madagascar, widespread (
Madagascar, Antongil Bay.
2 ♂♂: MAK-2; 2 exs.: MAK-43; 3 exs.: MAK-61.
Sub-Saharan Africa, Madagascar (
Madagascar.
5 ♂♂, 1 ♀: MAK-2; 1 ♂: MAK-62.
Sub-Saharan Africa, Madagascar (
Same as
Madagascar, Antananarivo, Ambodinandohalo Lake.
7 ♂♂, 3 ♀♀: MAK-2.
Madagascar, widespread but not common (
Same as
1 ♂, 1 ♀: MAK-2.
This large species may be
Unknown.
Same as
Madagascar, Montagne d’Ambre, Ambohitra National Park.
2 ♂♂, 8 ♀♀: MAK-3; 2 ♀♀: MAK-4; 30 exs.: MAK-5A; 8 exs.: MAK-5B; 15 exs.: MAK-5C; 85 exs.: MAK-5D; 5 ♂♂, 6 ♀♀: MAK-8; 1 ♀: MAK-10; 1 ♀: MAK-11B; 3 ♂♂, 5 ♀♀: MAK-14A; 1 ♀: MAK-16; 1 ♂, 1 ♀: MAK-25B; 4 exs.: MAK-28; 14 exs.: MAK-29; 1 ♂: MAK-35B; 1 ♂, 1 ♀: MAK-37A; 5 ♂♂: MAK-40A; 1 ex.: MAK-40B; 1 ♂: MAK-50; 2 ♂♂, 1 ♀: MAK-52; 1 ♂: MAK-59C.
Comoro Islands, Madagascar (
Madagascar.
1 ♂: MAK-29.
Madagascar, widespread (
This species was collected at a single site located in inner Makay, in a semi-forested small valley. The habitat was a sun-exposed isolated pool on sandstone rock, rather deep (70 cm), with bottom made up of sand and stones with a moderate quantity of plant debris, with clear water and no vegetation. This site was well preserved from anthropogenic disturbance.
4 ♂♂, 3 ♀♀: MAK-44C; 1 ♂, 2 ♀♀: MAK-60.
The Malagasy species of the genus
Unknown (but not confined to the Makay area).
This species was collected at two inner massif sites in northern Makay, with very different habitat characteristics. One was a blind channel in forest connected to River Sakapaly; this site is the locus typicus of
1 ♀: MAK-2; 3 ♂♂, 3 ♀♀: MAK-42; 1 ♀: MAK-43; 1 ♂: MAK-44C; 1 ♂, 3 ♀♀: MAK-60; 19 ♂♂, 24 ♀♀: MAK-61; 3 ♂♂, 1 ♀: MAK-62.
The Malagasy species of the genus
Unknown (but not confined to the Makay area).
Madagascar, Isalo National Park, Canyon des Singes.
1 ♀: MAK-8; 8 ♂♂, 8 ♀♀: MAK-9; 1 ♀: MAK-36B; 1 ♀: MAK-39B.
Madagascar; previously known only from the sandstone massif of Isalo (
Madagascar, Ambalavao region.
5 ♂♂, 8 ♀♀: MAK-5A; 1 ♀: MAK-5B; 1 ♀: MAK-8; 2 ♀♀: MAK-13; 3 ♂♂, 4 ♀♀: MAK-14A; 1 ♀: MAK-26; 1 ♀: MAK-39A; 2 ♂♂: MAK-40A
Sub-Saharan Africa, Madagascar (
Madagascar.
1 ♀: MAK-2; 1 ♀: MAK-5A; 28 ♂♂, 19 ♀♀: MAK-11A; 1 ♀: MAK-11B; 2 ♂♂, 1 ♀: MAK-19; 2 ♂♂, 1 ♀: MAK-25A; 1 ♂: MAK-26; 1 ♂: MAK-38A; 1 ♂, 1 ♀: MAK-62.
Madagascar, widespread (
Kenya, Winam, Kavirondo Bay.
1 ♂: MAK-2.
Eastern Sub-Saharan Africa, Madagascar (
Madagascar, Ankarafantsika National Park, Mahajanga, Boeny.
2 ♂♂, 2 ♀♀: MAK-5A.
Madagascar, widespread (
Madagascar.
1 ♂, 1 ♀: MAK-21.
Madagascar (widespread in lowlands) (
This species was encountered only once in a peripheral site located in an open area. The habitat was a small isolated temporary puddle with sandy bottom and clear water under
Madagascar, Toliara, Makay massif, 10.7 km NW of Tsivoky.
8 ♂♂, 7 ♀♀: MAK-3; 2 ♂♂, 1 ♀: MAK-4; 4 ♂♂, 1 ♀: MAK-5A; 3 ♂♂: MAK-5B; 2 ♂♂: MAK-5C; 7 ♂♂, 5 ♀♀: MAK-5D; 1 ♂, 2 ♀♀: MAK-6; 1 ♂, 1 ♀: MAK-7; 11 ♂♂, 9 ♀♀: MAK-8; 2 ♂♂, 4 ♀♀: MAK-10; 13 ♂♂, 6 ♀♀: MAK-14A; 3 ♂♂, 2 ♀♀: MAK-15; 1 ♂, 1 ♀: MAK-16; 1 ♂, 1 ♀: MAK-25A; 9 ♂♂, 22 ♀♀: MAK-25B; 1 ♂, 1 ♀: MAK-26; 4 ♂♂, 4 ♀♀: MAK-28; 1 ♀: MAK-29; 5 ♂♂, 2 ♀♀: MAK-30; 3 ♂♂, 5 ♀♀: MAK-31A; 5 ♂♂, 2 ♀♀: MAK-31B; 4 ♂♂, 1 ♀: MAK-31C; 12 ♂♂, 20 ♀♀: MAK-32; 1 ♂, 4 ♀♀: MAK-33; 1 ♂: MAK-34A; 2 ♂♂, 3 ♀♀: MAK-34B; 11 ♂♂, 3 ♀♀: MAK-35A; 4 ♂♂: MAK-35B; 5 ♂♂: MAK-35C; 6 ♂♂, 5 ♀♀: MAK-36B; 8 ♂♂, 11 ♀♀: MAK-38A; 1 ♂, 1 ♀: MAK-39A; 3 ♂♂, 1 ♀: MAK-39B; 1 ♂: MAK-44C; 68 exs.: MAK-45; 2 ♂♂, 5 ♀♀: MAK-46; 4 ♂♂, 6 ♀♀: MAK-47; 21 exs.: MAK-49; 130 exs.: MAK-50; 26 exs.: MAK-51; 111 exs.: MAK-52; 108 exs.: MAK-53; 16 ♂♂, 19 ♀♀: MAK-54A; 64 exs: MAK-54B; 8 ♂♂, 1 ♀: MAK-58; 37 exs.: MAK-59A; 5 ♂♂, 2 ♀♀: MAK-59B; 62 exs: MAK-59C.
So far endemic to the Makay massif, Madagascar (
Southern Madagascar, Pays Androy.
1 ♀: MAK-1A; 8 ♂♂, 5 ♀♀: MAK-2; 1 ♂: MAK-18; 1 ♂: MAK-19.
Sub Saharan-Africa, Arabian Peninsula, Madagascar (
Mascarene Islands, Mauritius.
5 ♂♂: MAK-1A; 1 ♂: MAK-2; 1 ♀: MAK-3; 3 ♀♀: MAK-4; 4 ♂♂, 3 ♀♀: MAK-11A; 1 ♂: MAK-11B; 1 ♂: MAK-17; 1 ♂: MAK-18; 243 exs.: MAK-19; 1 ♀: MAK-21; 3 ♂♂, 9 ♀♀: MAK-23; 1 ♀: MAK-28; 1 ♂, 2 ♀♀: MAK-40A; 11 ♂♂, 14 ♀♀: MAK-41; 37 exs.: MAK-42; 1 ♂: MAK-44A; 1 ♂: MAK-59C; 4 ♂♂, 6 ♀♀: MAK-60; 6 ♂♂, 1 ♀: MAK-62.
Mauritius, Aldabra, Madagascar (
Madagascar.
1 ♀: MAK-2; 2 ♂♂, 7 ♀♀: MAK-19.
Madagascar, widespread but not very common (
Southern Madagascar, near Bekily.
1 ♀: MAK-2.
South and south-western Madagascar (very rare); the present record is the first since the original description of the species (
Same as
Madagascar, Isalo, Menamaty River.
1 ♂, 1 ♀: MAK-12C; 1 ♀: MAK-26; 1 ♀: MAK-52.
Madagascar, widespread outside from the Central Highlands (
This species was found at three inner massif sites, two in south-central and one in northern Makay. The surrounding was forested or semi-forested and without visible anthropogenic disturbance. The habitats were: a small pond with water slowly renewed, just downstream from a spring, partly sheltered by trees, with clay bottom, with plant debris and clear water, with sparse helophytes (
1 ♂: MAK-21.
This is an undescribed species close to
Madagascar (widespread in lowlands).
Same as
Madagascar, Annanarivo (= Antananarivo?).
35 ♂♂, 50 ♀♀: MAK-5A; 3 ♂♂: MAK-5B.
Central and southern Madagascar (
Same as
Madagascar, Sainte Marie Island.
1 ♂: MAK-2.
There is a complex of very similar species around
Madagascar (
Same as
Relative frequencies of occurrence of species across samplings for different sets of sampling sites (all, inner Makay, peripheral Makay, forested sites, semi-forested sites, non-forested sites) are given in Table
List of the species sampled, with indication of status and values of relative frequencies of occurrence (
Species | Status | NbOc ( |
|||||
---|---|---|---|---|---|---|---|
|
|||||||
|
E | 15 (17.2) | 20.6 | 0 | 28.6 | 14.7 | 0 |
|
E | 8 (9.2) | 11.0 | 0 | 14.3 | 8.8 | 0 |
|
E | 5 (5.7) | 6.9 | 0 | 11.4 | 2.9 | 0 |
|
|||||||
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
|||||||
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
W | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E* | 11 (12.6) | 5.5 | 50.0 | 5.7 | 2.9 | 44.4 |
? | 3 (3.4) | 1.4 | 14.3 | 0 | 2.9 | 11.1 | |
|
W | 12 (13.8) | 8.2 | 42.9 | 8.6 | 5.9 | 38.9 |
? | 3 (3.4) | 4.1 | 0 | 5.7 | 2.9 | 0 | |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
|||||||
|
|||||||
|
E | 20 (23.0) | 27.4 | 0 | 25.7 | 29.4 | 5.6 |
|
E | 3 (3.4) | 4.1 | 0 | 5.7 | 2.9 | 0 |
|
W | 12 (13.8) | 16.4 | 0 | 20.0 | 8.8 | 11.1 |
|
E | 21 (24.1) | 28.8 | 0 | 25.7 | 32.3 | 5.6 |
|
E* | 4 (4.6) | 5.5 | 0 | 8.6 | 2.9 | 0 |
E | 1 (1.1) | 1.4 | 0 | 2.9 | 0 | 0 | |
E | 2 (2.3) | 2.7 | 0 | 0 | 5.9 | 0 | |
|
E | 12 (13.8) | 16.4 | 0 | 20.0 | 11.8 | 5.6 |
|
|||||||
|
E | 6 (6.9) | 0 | 42.9 | 0 | 0 | 33.3 |
|
E | 1 (1.1) | 1.4 | 0 | 0 | 2.9 | 0 |
|
|||||||
|
W | 3 (3.4) | 0 | 21.4 | 0 | 0 | 16.7 |
|
|||||||
|
W | 1 (1.1) | 1.4 | 0 | 0 | 0 | 5.6 |
|
|||||||
|
W | 7 (8.0) | 5.5 | 21.4 | 2.9 | 2.9 | 27.8 |
|
W | 1 (1.1) | 1.4 | 0 | 0 | 0 | 5.6 |
|
E | 1 (1.1) | 1.4 | 0 | 0 | 2.9 | 0 |
|
W | 7 (8.0) | 1.4 | 42.9 | 2.9 | 0 | 33.3 |
|
E* | 5 (5.7) | 6.8 | 0 | 5.7 | 8.8 | 0 |
|
|||||||
|
E | 6 (6.9) | 1.4 | 35.7 | 0 | 2.9 | 27.8 |
|
E | 7 (8.0) | 2.7 | 35.7 | 0 | 5.9 | 27.8 |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
? | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 | |
|
E* | 3 (3.4) | 1.4 | 7.1 | 0 | 2.9 | 5.6 |
|
W | 7 (8.0) | 2.7 | 35.7 | 0 | 5.9 | 27.8 |
|
E | 1 (1.1) | 1.4 | 0 | 0 | 2.9 | 0 |
|
E* | 1 (1.1) | 1.4 | 0 | 2.9 | 0 | 0 |
|
E | 5 (5.7) | 0 | 35.7 | 0 | 0 | 27.8 |
E | 27 (31.0) | 35.6 | 7.1 | 31.4 | 44.1 | 5.6 | |
E | 9 (10.3) | 12.3 | 0 | 5.7 | 20.6 | 0 | |
? | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 | |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
E | 5 (5.7) | 1.4 | 28.6 | 0 | 2.9 | 22.2 |
|
E | 8 (9.2) | 1.4 | 50.0 | 0 | 2.9 | 38.9 |
|
|||||||
|
W | 10 (11.5) | 4.1 | 50.0 | 2.9 | 5.9 | 38.9 |
|
W | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E | 3 (3.4) | 1.4 | 7.1 | 0 | 2.9 | 5.6 |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
W | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
W | 3 (3.4) | 1.4 | 7.1 | 0 | 2.9 | 5.6 |
|
W | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
? | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 | |
|
|||||||
|
E* | 21 (24.1) | 28.8 | 0 | 25.7 | 32.4 | 5.6 |
|
E | 1 (1.1) | 1.4 | 0 | 0 | 2.9 | 0 |
|
|||||||
? | 2 (2.3) | 2.7 | 0 | 2.9 | 2.9 | 0 | |
? | 7 (8.0) | 4.1 | 28.6 | 2.9 | 5.9 | 22.2 | |
|
|||||||
|
E | 4 (4.6) | 5.5 | 0 | 8.6 | 0 | 5.6 |
|
W | 8 (9.2) | 11.0 | 0 | 17.1 | 5.9 | 0 |
|
E | 9 (10.3) | 8.2 | 21.4 | 5.7 | 5.9 | 27.8 |
|
W | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
E | 1 (1.1) | 1.4 | 0 | 0 | 2.9 | 0 |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
E | 48 (55.2) | 65.6 | 0 | 62.9 | 73.5 | 5.6 |
|
W | 4 (4.6) | 0 | 28.6 | 0 | 0 | 22.2 |
|
E* | 19 (21.8) | 13.7 | 64.3 | 8.6 | 14.7 | 61.1 |
|
E | 2 (2.3) | 0 | 14.3 | 0 | 0 | 11.1 |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
|
E | 3 (3.4) | 4.1 | 0 | 2.9 | 5.9 | 0 |
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 | |
|
E | 2 (2.3) | 2.7 | 0 | 0 | 5.9 | 0 |
|
E | 1 (1.1) | 0 | 7.1 | 0 | 0 | 5.6 |
According to vegetation contexts, dominant species (
We also computed dissimilarity indices based on all species to see how the whole community varies across categories of sites (Jaccard dissimilarity index, calculated from occurrence data; Bray-Curtis dissimilarity index, taking into account numbers of individuals captured for each species; Fig.
Comparison across site categories of percentages of endemics for dominant or rare species, and calculated dissimilarity indices for species composition
Observed percentages of species endemic to Madagascar are 58.1% for all samplings, 55.3% for inner Makay, 53.3% for peripheral Makay, 48.3% for forested sites, 58.5% for semi-forested sites and 53.7% for non-forested sites. Thus, when all sampled species are considered, rather surprisingly, endemicity is very similar in inner Makay vs. in the (deforested) peripheral plain and is also similar across categories of environment with the lowest value for forested areas, the highest for semi-forested areas, and non-forested areas standing in-between. When looking at percentages of endemic species among dominant species (
Observed species richness (number of species counted in the samplings) was 74 for all samplings, 45 for peripheral Makay, 47 for inner Makay, 29 for forested sites, 41 for semi-forested sites and 54 for deforested sites. Because observed species richness is a very poor proxy for species diversity, we ran interpolation-extrapolation analyses to obtain estimates of the H0, H1 and H2 metrics (see Material and methods) for the various categories of sites (Fig.
Interpolation-extrapolation graphs for the whole Makay dataset (All), for samplings in inner and in peripheral Makay, and for samplings in different vegetation contexts. Coloured lines represent the interpolated (solid line) or extrapolated (dashed line) estimate of the metric against number of individuals; the surface of lighter colour surrounding each curve materialises the 95% confidence interval
This faunistic study represents the first survey of predaceous water beetles (aquatic
In the current state of knowledge, five species of aquatic
Notwithstanding this singularity and an undeniable patrimonial value, the aquatic
There is a critical lack of published studies with comparable datasets on which to confront quantitatively species diversity and endemism level of inner Makay with those of other Malagasy massifs, in order to substantiate the conclusion that species diversity and endemism level of aquatic
The Makay massif bears strong similarities and a geographical proximity with the massif of Isalo, making comparison of the aquatic
We are able to provide comparative estimates of species diversity, based on our own unpublished sampling data in southern Isalo (obtained in May 2016, using similar collecting techniques to those deployed in the Makay; with satisfying sample coverage as shown in Fig.
Interpolation-extrapolation graphs for inner Makay and inner Isalo. Coloured lines represent the interpolated (solid line) or extrapolated (dashed line) estimate of the metric against number of individuals; the surface of lighter colour surrounding each curve materialises the 95% confidence interval
Why is species diversity of aquatic
Finally, we would like to point out a few remarkable findings from our samplings in the peripheral plain surrounding the Makay massif. At ~ 15 km south-west of Makaikely, a small and shallow stream (MAK-2) with sandy bottom and very slowly flowing water, strongly impacted by cattle trampling and highly eutrophicated, provided an impressive sampling with 32 species of aquatic
We warmly acknowledge for their decisive contribution to our field work our colleague Catherine Reeb and our guide and driver Parsson. We are very grateful to Bernard Forgeau for sharing with us his deep knowledge of the Makay and its people, for guidance in the field and assistance in field work, and for ensuring good interface with the local population of the Makay. Local guides are also thanked for their help. Jean-François Cart is warmly acknowledged for help in field work during one of the campaigns and for constant material support to ATR during his PhD thesis. ATR addresses a special acknowledgement to Clémence, his parents and other members of the Malavergne family, for their constant help and support, and notably financial support to help his sister when she needed health care as well as their decisive material help toward the completion of his PhD thesis, notably through financing his last stay in France, and much more. The association Nature Evolution partly organised the field trips of 2017 and 2018 in which ATR participated. We thank Muriel Jager for technical help and Antoine Mantilleri for access to material of the
Table S1
Cumulated abundances of species per categories of sampling sites in Makay
This table gives for each category of sites and for each species the total number of specimens sampled. The second column indicates the status of the species with respect to endemism. These data were used to build the graphs of Figs
Table S2
Cumulated abundances of species sampled in inner Makay vs. in inner Isalo.
This table gives for each species the total number of specimens sampled in inner Makay and in inner Isalo. These data were used to build the graphs of Fig.