Corresponding author: Tolotra Ranarilalatiana (
Academic editor: Mariano Michat
The genus
Ranarilalatiana T, Raveloson Ravaomanarivo LH, Bergsten J (2019) Taxonomic revision of the genus
The subfamily
To be able to deal with this diversity,
The species that occur in Madagascar fall into five of these species groups: the
This study is motivated by recent collecting efforts of aquatic beetles in Madagascar 2009–2018 in the Water Beetles of Madagascar Project. The project is a collaboration between the Swedish Museum of Natural History and the University of Antananarivo. The current study is based on this rich new material containing five new species of the
New collecting efforts of aquatic beetles were conducted mainly in National Parks, reserves and natural forests but also in degraded forests, open areas and along main roads from all parts of Madagascar, except scant from the very south (Fig.
Maps of Madagascar with records and first level administrative divisions.
Each locality was given a collecting event code and associated metadata included geographic name(s), forest type, waterbody type, habitat description, eventual disturbance, collecting date and collectors. Altitude, latitude and longitude were recorded with a handheld GPS (Garmin). Each locality was also documented with photographs using compact Canon and Olympus digital cameras.
Madagascar is since the 2007 revision of the constitution divided into 22 regions as a first level administrative division, followed by districts as a second level, a recent change from the former six provinces as first level (Fig.
Most of the studied material came from the new fieldwork and is shared between the Swedish Museum of Natural History (
Additional studied material came from earlier expeditions housed at museum collections in
Depositories of studied specimens are referred to by above abbreviations (see Suppl. material
Specimens were examined under dissection microscopes from Leica (M165C and MZ12.5). Genitalia were extracted with a fine forceps or pin from the tip of the abdomen and glued onto cards on the same pin as the specimen. Dry-preserved specimens were first relaxed in warm water for 5–20 minutes before genitalia were carefully extracted. Photos of habitus were taken with a Canon EOS 5D Mark II DSLR camera equipped with a Canon MP-E 65mm 1–5X super macrolens and mounted on a motorised rail (Stackshot) from Cognisys. The system was operated using Canon EOS Utility and Zerene Stacker (Zerene Systems) softwares, the latter also used for stacking the Z-stack of captured images with the PMax or DMap algorithms. Photographs of dry-mounted genitalia were taken with a Canon EOS 7D DSLR camera mounted on a BALPRO 1 Universal bellow from Novoflex with a long working distance 10X Plano apochromatic microscope objective from Mitutoyo. The bellow was mounted on a motorised rail (Stackshot) from Cognisys and operated with the same software given above.
In describing the male penis we use the terminology suggested by
Label data are given as written and separated by “//” if on separate labels and “|” if on different rows on the same label. Text within square brackets “[]” are our comments, explications or interpretations. Most examined specimens (individual mounted specimens, or single alcohol tubes with multiple specimens) have been given unique catalogue numbers and these are listed first, starting with “
DNA was extracted from one mesoleg or from soft abdominal tissue retrieved in association with dissection of male genitalia. The leg or soft tissue was incubated in lysis buffer at 56 °C overnight. Post-incubation protocol followed the cell and tissue DNA kit on a KingFisher Duo Prime. This system provides automated nucleic acid purification at a running time of approx. 25 minutes.
We used ready-to-go beads to prepare 25 μl PCR reactions consisting of 21 μl of water, 1 μl of each primer and 2 μl of DNA template. We used the primers Jerry (F, 5’-CAA CAT TTA TTT TGA TTT TTT GG-3’) and PatDyt (R, 5’-TCA TTG CAC TAA TCT GCC ATA TTA G-3’) to amplify an 825 bp fragment of mitochondrial cytochrome c-oxidase subunit 1 gene (COI or
Successful PCR products were purified using EXOSAP Clean-up mix of two enzymes (Exonuclease and Shrimp Alkaline Phosphatase) and run in a PCR machine with the programme 37 °C for 30 min followed by 80 °C for 15 min and finally 12 °C (∞). PCR products were sent to Macrogen for sequencing.
Gene regions were sequenced in both directions and sequence chromatograms were edited with SEQUENCHER version 4.10.1 (Gene Codes Corporation). The contigs were assembled from forward and reverse reads, and primer regions trimmed. New sequence data were then exported in fasta format and aligned together with GenBank sequences of
Details of material used for DNA analysis and GenBank accession numbers for mitochondrial COI. New sequences submitted to GenBank have accession numbers starting with “MK”. For samples without a separate extract number, the extract is identified by the ID Cat. No.
Species | ID Cat. No. | Extract | Field ID | Place | Lat/Long | Alt | Date | Accession numbers |
---|---|---|---|---|---|---|---|---|
|
BMNH-797876 | 294:A4 | P57BI31 | Marojejy NP |
|
162 | 10/12/07 |
|
BMNH-797894 | 294:B10 | P61BI15 | Andasibe NP |
|
940 | 06/01/07 |
|
|
BMNH-797906 | 294:C10 | P58BI14 | Masoala NP, E. of Marofototra |
|
10 | 17/12/06 |
|
|
BMNH-797907 | 294:C11 | P58BI14 | Masoala NP, E. of Marofototra |
|
10 | 17/12/06 |
|
|
BMNH-797908 | 294:C12 | P58BI14 | Masoala NP, E. of Marofototra |
|
10 | 17/12/06 |
|
|
BMNH-797909 | 294:D01 | P58BI14 | Masoala NP, E. of Marofototra |
|
10 | 17/12/06 |
|
|
BMNH-797910 | 294:D02 | P58BI14 | Masoala NP, E. of Marofototra |
|
10 | 17/12/06 |
|
|
JB196 | MAD09-07 | Ankarafantsika NP, Ampijoroa |
|
74 | 29/11/09 |
|
||
JB197 | MAD09-46 | Kirindy Res. |
|
52 | 12/12/09 |
|
||
JB198 | MAD09-13 | Ankarafantsika NP, Ampijoroa |
|
75 | 30/11/09 |
|
||
JB199 | MAD09-07 | Ankarafantsika NP, Ampijoroa |
|
74 | 29/11/09 |
|
||
JB200 | MAD09-59 | Tsingy de Bemaraha NP, Bekopaka |
|
41 | 15/12/09 |
|
||
JB201 | MAD09-03 | Ankarafantsika NP, Ampijoroa |
|
87 | 29/11/09 |
|
||
JB202 | MAD09-29 | Mahavavy Kinkony Res., Mitsinjo |
|
24 | 05/12/09 |
|
||
JB203 | MAD09-65 | Tsingy de Bemaraha NP, Antsalova |
|
119 | 17/12/09 |
|
||
JB191 | MAD09-14 | Ankarafantsika NP, Ampijoroa |
|
77 | 30/11/09 |
|
||
JB192 | MAD09-24 | Mahavavy Kinkony Res., Makary village |
|
9 | 04/12/09 |
|
||
JB193 | MAD09-25 | Mahavavy Kinkony Res., Makary village |
|
19 | 04/12/09 |
|
||
JB194 | MAD09-28 | Mahavavy Kinkony Res., Mitsinjo |
|
22 | 05/12/09 |
|
||
JB189 | MAD09-30 | Mahavavy Kinkony RS, Mitsinjo |
|
55 | 05/12/09 |
|
||
JB809 | MAD09-24 | Mahavavy Kinkony RS, Makary village |
|
9 | 04/12/09 |
|
||
TR18L14 | Ambohidray Res., Andriambe |
|
1044 | 23/05/18 |
|
|||
TR18L04 | Ambohidray Res., Andriambe |
|
1044 | 07/04/18 |
|
|||
TR18L04 | Ambohidray Res., Andriambe |
|
1044 | 07/04/18 |
|
|||
TR18L07 | Ambohidray Res., Andriambe |
|
1046 | 07/04/18 |
|
|||
|
JB204 | MAD09-74 | btw Morafenobe-Ambohijanahary Res. |
|
290 | 19/12/09 |
|
|
|
BLF4432 | Tsingy de Bemaraha NP |
|
150 | 16/11/01 |
|
||
|
BMNH-670601 | 007:E07 | P27MD31 | Ranomafana NP |
|
1123 | 06/12/04 |
|
BMNH-729896 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
BMNH-792954 | P39EM08 | Andringitra NP |
|
1420 | 09/05/06 |
|
||
BMNH-792955 | P39EM08 | Andringitra NP |
|
1420 | 09/05/06 |
|
||
BMNH-792956 | P39EM08 | Andringitra NP |
|
1420 | 09/05/06 |
|
||
BMNH-792962 | P36C | RN7, Col de Tapias |
|
1717 | 06/05/06 |
|
||
BMNH-792963 | P36C | RN7, Col de Tapias |
|
1717 | 06/05/06 |
|
||
BMNH-792964 | P36C | RN7, Col de Tapias |
|
1717 | 06/05/06 |
|
||
BMNH-792976 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
BMNH-792977 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
BMNH-792978 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
BMNH-792979 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
BMNH-792980 | P30MD33 | Sahatsiho Ambohimanjaka |
|
1442 | 08/12/04 |
|
||
MAD14-81 | RN2, Betsabora river |
|
900 | 24/11/14 |
|
|||
MAD16-47 | Manjakatompo Ankaratra Res., Ankafotra Mtn. |
|
2466 | 18/09/16 |
|
|||
|
MAD14-81 | RN2, Betsabora river |
|
900 | 24/11/14 |
|
||
MAD18-91 | Zahamena NP, Sect. Antanandava |
|
1040 | 08/03/18 |
|
|||
MAD11-26 | Analamazaotra NP |
|
930 | 08/11/11 |
|
|||
MAD14-18 | Analamazaotra NP |
|
930 | 27/11/14 |
|
|||
BMNH-797895 | P60BI15 | Zahamena NP, Sect. Antanandava |
|
1075 | 31/11/06 |
|
||
MAD11-37 | Mantadia NP |
|
1000 | 11/11/11 |
|
|||
MAD14-70 | Anjanaharibe Sud res. |
|
910 | 16/11/14 |
|
|||
MAD14-04 | Ranomafana NP |
|
1130 | 02/11/14 |
|
|||
MAD12-03 | Isalo NP, Canyon des Makis |
|
700 | 13/11/12 |
|
|||
MAD12-03 | Isalo NP, Canyon des Makis |
|
700 | 13/11/12 |
|
|||
MAD13-55 | Ivohibe RS |
|
874 | 09/12/13 |
|
|||
MAD13-55 | Ivohibe RS |
|
874 | 09/12/13 |
|
|||
|
MAD14-81 | RN2, Betsabora river |
|
900 | 24/11/14 |
|
||
MAD14-81 | RN2, Betsabora river |
|
900 | 24/11/14 |
|
|||
JB190 | MAD09-58 | Tsingy de Bemaraha NP, Bekopaka |
|
66 | 15/12/09 |
|
||
JB195 | MAD09-33 | Mahavavy Kinkony Res., Anjohibe |
|
24 | 06/12/09 |
|
||
JB802 | MAD09-25 | Mahavavy Kinkony Res., Makary village |
|
19 | 04/12/09 |
|
||
MJK12-13 | Manjakatompo Ankaratra Res., Anosiarivo |
|
2073 | 24/01/12 |
|
|||
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. |
|
2597 | 07/02/16 |
|
|||
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. |
|
2597 | 07/02/16 |
|
|||
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. |
|
2597 | 07/02/16 |
|
|||
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. |
|
2597 | 07/02/16 |
|
|||
|
JB808 | MAD09-25 | Mahavavy Kinkony RS, Makary village |
|
19 | 04/12/09 |
|
|
MAD14-14 | Analamazaotra NP |
|
930 | 27/11/14 |
|
|||
MAD14-14 | Analamazaotra NP |
|
930 | 27/11/14 |
|
|||
Analamazaotra NP, Andasibe |
|
938 | 17/01/15 |
|
||||
JB206 | MAD09-07 | Ankarafantsika NP, Ampijoroa |
|
74 | 29/11/09 |
|
||
JB205 | MAD09-03 | Ankarafantsika NP, Ampijoroa |
|
87 | 29/11/09 |
|
||
MAD13-61 | Ivohibe RS, Andranovory |
|
1106 | 10/12/13 |
|
|||
MAD13-61 | Ivohibe RS, Andranovory |
|
1106 | 10/12/13 |
|
|||
MAD11-52 | RN5, Ivoloina |
|
0 | 15/11/15 |
|
|||
MAD17-12 | Analalava Res., Analalava forest |
|
39 | 09/03/17 |
|
|||
BMNH-672727 | 027:A05 | P32 | Tsaratanana massif, Mangindrano |
|
1700 | 20/12/04 |
|
|
BMNH-672728 | 027:A06 | P32 | Tsaratanana massif, Mangindrano |
|
1700 | 20/12/04 |
|
|
BMNH-672729 | 027:A07 | P32 | Tsaratanana massif, Mangindrano |
|
1700 | 20/12/04 |
|
We first performed a Bayesian phylogenetic analysis to produce a CO1-genetree. As the taxon selection here is not aimed at producing a phylogeny, but to interprete genetic variation in light of morphological delimitations of a diagnosable set of species in a certain geographic region (Madagascar), the genetree was artificially rooted using
To explicitly compare our morphological delimitations with a single-locus species delimitation method we implemented the GMYC-method (
Our amplification of CO1 was successful for 53 samples which, together with sequences downloaded from GenBank, gave 77 terminals (Table
Specimens of morphologically identified species all clustered as monophyletic in the Bayesian analysis except
Majority-rule consensus tree from Bayesian analysis of CO1. Values next to nodes indicate posterior probabilities.
Uncorrected P genetic distances between closely related species or populations in the group near
|
|
|
|
|||||
---|---|---|---|---|---|---|---|---|
|
0–0.007 | |||||||
(0–0.007) | ||||||||
0.005–0.009 | 0 | |||||||
(Ankaraf.) | (0.005–0.009) | (0) | ||||||
|
0.001–0.017 | 0.006–0.011 | 0–0.017 | |||||
(0.001–0.017) | (0.006–0.011) | (0–0.017) | ||||||
|
0.032–0.036 | 0.028–0.031 | 0.028–0.038 | 0–0.004 | ||||
(peak) | (0.033–0.038) | (0.029–0.032) | (0.029–0.039) | (0–0.004) | ||||
|
0.030–0.039 | 0.029–0.031 | 0.028–0.038 | 0.023–0.025 | NA | |||
(0.031–0.040) | (0.030–0.032) | (0.029–0.040) | (0.023–0.025) | NA | ||||
|
0.026–0.029 | 0.028 | 0.024–0.029 | 0.033–0.035 | 0.033 | 0 | ||
(0.026–0.030) | (0.028–0.029) | (0.024–0.030) | (0.034–0.036) | (0.034) | (0) | |||
|
0.046–0.048 | 0.044–0.045 | 0.048–0.050 | 0.059–0.060 | 0,061 | 0.051–0.052 | 0 | |
(Ivohibe) | (0.048–0.049) | (0.045–0.047) | (0.050–0.053) | (0.062–0.064) | (0.065) | (0.054) | (0) | |
|
0.051–0.054 | 0.047–0.049 | 0.052–0.053 | 0.063–0.066 | 0.066 | 0.058–0.059 | 0.023–0.024 | 0 |
(N. Toam.) | (0.053–0.057) | (0.049–0.051) | (0.054–0.056) | (0.067–0.071) | (0.070–0.071) | (0.061–0.062) | (0.023–0.024) | (0) |
The GMYC species delimitation analysis of the strict clock tree resulted in 11 separate evolutionary units that were largely but not entirely consistent with our morphological delimitation (Fig.
Result of the single-locus GMYC species delimitation using an ultrametric CO1 genetree from BEAST. Black branches indicate interspecific divergences, red branches represent intraspecific coalescence events. Values above interspecific nodes indicate posterior probability from the Beast analysis under a strict clock model. The * indicates two nodes of further splitting from the GMYC analysis that is within the confidence interval of 2Log likelihood units from the optimal solution.
Key to
1 | Impressed elytral striae absent (rows of points may be present) (Fig. |
|
– | Impressed elytral striae present (Figs |
|
2 | Body length less than 4 mm (Fig. |
|
– | Body length between 5.7 and 6.6 mm (Fig. |
|
3 | Elytra with five discal but no submarginal stria; dorsal surface reddish brown with a lighter elytral base; first elytra stria abbreviated and only present in posterior third; body length 4.1–4.2 mm (Fig. |
|
– | Elytra with six discal and one submarginal stria (Figs |
|
4 | First, third, and fifth elytral striae distinctly abbreviated anteriorly; dorsal colouration largely black (Fig. |
|
– | Third and fifth elytral striae not abbreviated anteriorly; first elytral stria abbreviated or not; dorsal colouration rarely mostly black; penis shape variable |
|
5 | Body shape more broadly oval (Fig. |
|
– | Body shape narrow, elongate, subparalell (Figs |
|
6 | Elytra without a transverse testaceous band at base, black except laterally and posteriorly (Fig. |
|
– | Elytra with basal testaceous transverse band; penis apex in ventral view right-curved |
|
7 | Body broadly oblong; first elytral stria often longer and basal testaceous elytral band narrower but both characters overlapping; penis in lateral view with dorsal knob and ventral invagination; penis in ventral view not broadly expanded apically, but with a small tooth (Fig. |
|
– | Body more narrowly oblong and attenuating posteriorly; first elytral stria often shorter and elytra with a broader testaceous transverse band at base; penis in ventral view bisinuate and widened at apex (Fig. |
|
8 | Elytra with irregular traces of intermediate striae or “pseudostriae” between first and second stria and between second and third stria; body length larger, 5.3 to 5.6 mm (Fig. |
|
– | Elytra without pseudostriae in elytral intervals; body length smaller, less than 5.3 mm; Penis curvature in lateral view stronger |
|
9 | Body subparallell and head with a broad interocular distance; pronotum without strioles (Fig. |
|
– | Body more attenuating anteriorly and posteriorly and interocular distance narrower; pronotum with strioles posterolaterally; penis slender, without subapical expansion in lateral view |
|
10 | Dark colouration dorsally, with narrow testaceous band at base of elytra and testaceous, sometimes strongly contrasting, anterolateral pronotal corners; head distinctly infuscated (Fig. |
|
– | Colouration usually lighter brown, testaceous band at base of elytra variable; base of penis in posteroventral view not angled; apex in lateral view with or without a dorsal ridge crossing posterior inner outline |
|
11 | Body length smaller, 3.8 to 4.3 mm (Fig. |
|
– | Body length larger, 3.9 to 5.0 mm; penis in lateral view with a dorsal ridge crossing posterior inner outline near apex |
|
12 | Elytra usually with a more narrow basal testaceous band; penis in lateral view with distinct shoulder definition interrupting an evenly curved outer outline (Fig. |
|
– | Elytra often with broader basal testaceous band; penis in lateral view evenly curved without a distinct shoulder definition (Fig. |
|
Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere.
Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere.
Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere.
The following three taxa were described as
New Caledonia.
Madagascar, Suberbieville [= Maevatanana].
Madagascar, Suberbieville [= Maevatanana].
This group is defined by the lack of elytral striae (
Madagascar, Massif Ankaratra, Manjakatompo, alt. 1700–1800 m.
based on a single female specimen (holotype), collected December 1951 by R. Benoist.
Small size (4 mm). Elytra uniformly dark brown ferrugineous, without a basal testaceous area (Fig.
(based on holotype ♀):
Body length 4 mm. Body shape elongate oval and dark brown to blackish ferrugineous. Head uniformly dark brown ferrugineous to slightly darker posteriorly inside eyes, with thin sparse punctation. Pronotum dark brown ferrugineous, same colour medially and laterally but darker along anterior and posterior third. Disc of pronotum less densely punctuated, posterolateral corners with dense superficial strioles. The entire dorsal surface covered with a microsculpture. Elytra uniformly coloured in same dark brown to blackish ferrugineous colour as anterior and posterior parts of pronotum (Fig.
Habitus, dorsal view.
Ventral side testaceous to weakly infuscated. Prosternal process carinate also onto apical process. Lateral parts of metaventrite (“metasternal wings”) rather broad. Metacoxal lines anteriorly diverging and abbreviated before metaventral margin. Metacoxa with fine and long longitudinal strioles, continuing onto abdominal ventrites, and with 6–7 transverse “wrinkles” anterolaterally.
Male: unknown.
Madagascar, central highlands, only known from type locality Manjakatompo, Ankaratra Massif (Fig.
Unknown, but according to original description collected at at an altitude of 1700–1800 m. See
No other specimen than the female holotype is known of this species and it is a bit of a “mystery species”. We have conducted fieldwork at the type locality Manjakatompo multiple times (2011, 2012, 2014, and 2016) but never found any specimens resembling this species. The species belongs in the
Elisabethville, Zaire [DR Congo, Haut-Katanga, Lubumbashi].
-
-1♂(
Similar to
Body length 5.7–6.6 mm. Body shape elongate oval, from midpoint uniformly tapering towards a rather pointed apex. Head and pronotum both in the same dark brown to blackish ferrugineous colouration; elytra anteriorly in the same colour as pronotum, but posterior part brown ferrugineous with largely testaceous lateral margins (Fig.
Ventral side dark ferrugineous. Prosternal process strongly carinate anteriorly and with a rather short process. Lateral parts of the metaventrite medium-broad. Metacoxal lines short, anteriorly diverging and abbreviated well before metaventral margin. Metacoxa with short fine strioles continuing onto abdominal ventrites.
Male: first three pro- and mesotarsomeres widened. Protibia modified, narrow at base and with an early abrupt bend, extended and broadened towards middle with a straight ventral side but angled dorsal side. Pro- and mesotarsal claws unmodified.
Penis in ventral view with apical part more or less straight and even, gently pointed at apex but with the very apex minutely twisted to the right (Fig.
Female: dorsal sculpture similar to male.
Likely a more widespread distribution in at least central and eastern continental Africa than the current records (Lubumbashi and Kivu in DR Congo) indicate. In Madagascar, only know from the eastern central parts: Betongolo (Antananarivo), the Analamazaotra NP, and P.K.57 Route d’Anosibe [RN23] (Fig.
A series of six specimens collected with light trap (“piège lumineux”) in the capital Antananarivo 1946, indicates flight capacity and anthropogenically disturbed habitats. All records from DR Congo are also from light trap catches (
This species was described from Lubumbashi, DR Congo, and has not been recorded from Madagascar before. Earliest record found is from November 1946. It seems to be a dispersive good flier and often collected at light so its presence in Madagascar is therefore not surprising. However, it is not widespread in Madagascar as far as we know. In fact, the known distribution is restricted to the surroundings of the capital and east of the capital along the main national route towards Toamasina, which could suggest a recent incidental human-mediated introduction from mainland Africa.
Note that it may be that this is a species with intraspecific variation with regards to elytral striation, ranging from five puncture lines out of which two are more distinct, to five weakly impressed striae (see further discussion under
Madagascar, Morondava, forest south of Befasy.
based on an unknown number of female type specimens but holotype and paratypes are distinguished in introduction. Collected in January 1956 by R. Paulian.
Similar to
Body length 4.1–4.2 mm. Body shape elongate and subparallel, dorsal surface reddish brown with a lighter elytral base. Head and pronotum uniformly rufus brown. Head, pronotum and elytra with dense punctation. Elytra and pronotum covered with dense punctures and the whole dorsal surface with a microsculpture. Lateral sides of pronotum striolate with the widest striolate area in the posterior corners. Elytra light brown with a distinct testaceous band basally (Fig.
Ventral side light brown. Metacoxa and abdominal ventrites punctate and striolate. Prosternal process rather short and medially raised, triangular in cross-section. Lateral parts of metaventrite medium broad. Metacoxal lines anteriorly diverging but rather weakly so, abbreviated well before metaventral margin. Antennae, palps and legs testaceous.
Male: unknown.
Known only from two localities in the western part of Madagascar, the deciduous forest south of Befasy, Morondava, and at one locality between Morafenobe and Beravina village (Fig.
Paulian collected the species in 1956 in the western dry deciduous forest south of Befasy, Morondava. We rediscovered the species in 2009, when we found one female specimen of
This is the only species in the
The
Environs de Tananarive [surroundings of Antananarivo] and Fianarantsoa, Madagascar.
Based on an unknown number of specimens (syntypes) collected by Sikora (Antananarivo) and Perrot (Fianarantsoa).
Type material in
Body shape elongate oval, convex, and attenuate posteriorly, with uniform black colouration (Fig.
Body length: 5.3–6.3 mm. Body shape elongate oval, convex, and attenuate posteriorly. Head, pronotum, and elytra all the same colour, ferrugineous black and finely punctate.
Lateral margin of pronotum rusty ferrugineous, with short sparse strioles. Pronotum with puncture rows and microsculpture. Elytra narrowly testaceous to ferrugineous posterolaterally. Six elytral striae present and one submarginal stria (Fig.
Ventral side ferrugineous dark brown. Metacoxa and abdominal sternite II, III, IV, striolate. Prosternal process raised medially but rounded, not carinate. Lateral parts of metaventrite broad. Metacoxal lines short and strongly diverging anteriorly. Antenna, palps, pro- and mesothoracic legs brown to yellowish, but metathoracic legs dark brown.
Male: protibia slightly widened at apex, somewhat curved and angulate basally. Penis in lateral view curved with two points where curvature is more abrupt, constricted before apex at a narrow “neck” and expanding to apical part (Fig.
Female: dorsal sculpture similar to male.
Occurs on Madagascar and Mauritius (
This species has been collected in various localities, mostly from open, partly deforested areas at mid- to high altitudes or open forest marshes. It occurs at altitudes above 900 m and is often associated with grass vegetation along lake shores and in marshes, found by stamping, and at vegetation-rich margins of rivers.
Egypt, Sinai.
-
Similar to
(based on Malagasy specimens):
Body length 5.5–6.1 mm. Body shape oblong oval, rather convex and attenuate posteriorly, dark brown to blackish ferrugineous. Head infuscated brown ferrugineous, somewhat lighter posteriorly, covered with dense microreticulation and sparser punctation.
Pronotum dark brown to ferrugineous black with testaceous anterolateral corners. Disc covered with fine microsculpture forming regular cells and regularily spread small punctures of about same size as cells. Punctuation becomes coarser in posterolateral corners with a weak tendency to corrugate.
Elytra predominantely dark brown to ferrugineous black on disc and along striae with or without a rather narrow and vague basal testaceous band (Fig.
Ventral side ferrugineous dark brown, with testaceous spots laterally on abdominal ventrites. Metacoxa and ventrites with strioles. Prosternal process more elongate lanceolate and with blunter apex compared with
Female: elytral striolation limited to the medial parts of the outer three elytral intervals in the single female studied from Madagascar. From other parts of the distribution a female form is known that has the entire elytra striolated (
Male: protibia bisinuate and angled at base, distally expanded. Penis thin, strongly angled at middle in lateral view, and apex somewhat twisted to the left in ventral view (Fig.
Habitus, dorsal view.
As the species
On Madagascar we have collected the species associated with a small forest stream with sidepools in a karstic limestone area (“tsingy”) and in a muddy stagnant pool in a dried-out river bed. Both localities are in dry deciduous forests of lowland western Madagascar.
Senegal [possibly mislabelled].
housed in Buquet collection and originating from Senegal; of
-
Body length 5.2–6.6 mm. Body shape oval, rather convex and attenuate posteriorly, dark brown to brown ferrugineous. Head, pronotum and elytra in the same dark brown ferrugineous, covered with fine dense punctation. Lateral sides of pronotum more brownish, with short strioles and the widest striolate area in the posterior corners. Elytra dark brown to brown ferrugineous, with a testaceous transverse band at base (Fig.
Ventral side brownish to ferrugineous, metacoxa with microsculpture, densely and finely punctate. Metacoxa and abdominal ventrites striolate. Prosternal process rather short and spear-shaped, medially only weakly raised and rounded. Lateral parts of metaventrites rather broad. Metacoxal lines short and rather strongly diverging anteriorly. Antennae and palps both in the same brown colour. Pro- and mesothoracic legs brown ferrugineous. Metathoracic legs dark brown ferrugineous.
Male: Protibia strongly angled basally and expanded in apical two thirds. Penis in ventral view with a small preapical tooth on right side; in lateral view very characteristic with a subbasal dorsal knob, and post-middle with a deep ventral invagination (Fig.
Females from Madagascar usually with elytral striolation rather weak and restricted to outer intervals, but rarely the elytra are entirely and distincly striolate. Females on average smaller than males.
Endemic to the western Indian Ocean islands as far as is modernly known, but the 1835 type locality of the original description and labels read “Senegal”, which indicates either mislabeling or that the species in fact also occurs on continental Africa. Known from Madagascar, Reunion, Comores, Seychelles, and Aldabra Island (
This is a lowland species that seems to be most common in the deciduous western parts of Madagascar. It was common in the newly designated protected areas of Mahavavy Kinkony when we visited it in 2009. In the deciduous forest biome it has been recorded from Kirindy in the south to Ankarana NP in the north. The species seems to be a generalist and can as well show up on lowland humid east coast and at midaltitudes up to at least 1000 m. It is an apt flier collected with light traps and often in very temporary and small shallow pools including water-filled wheel tracks. Found in all kinds of temporary pools, as well as in streams and in residual pools in dried-up riverbeds. It occurred sympatrically with
Madagascar, Mahajanga Distr., Mahajanga env. [ca. 15°43'S, 46°18'E]
Based on male (holotype), 3 male and 4 female (paratypes). I. Janis 1–10 December 1996.
Close to
Body length 5.2–6.2 mm. Body shape oblong oval, rather convex, brown to dark brown. Head ferrugineous brown, paler in front, sometimes darker around eyes, finely microreticulate and punctate, two shallow depressions between eyes. Pronotum dark brown, paler at sides. Dorsal surface with fine microsculpture and scattered punctures, lateral sides of pronotum striolate with the widest striolate area in the posterior corners. Elytra brown, paler at sides and at apex, with a broad testaceous transverse band at base (Fig.
Ventral side ferrugineous brown, metacoxa and abdominal ventrites striolate and punctate. Prosternal process similar to
Male: protibia modified, widened in front, strongly angled after base, with several long spines on the outer side of distal half. Tarsomeres I–III of pro- and mesolegs enlarged, with pads of numerous setae. Penis unique: in ventral view with a bisinuate shape and a widening asymmetrical apex; in lateral view abruptly curved near middle, with a right angle of more or less 90 degrees, the basal part robust (Fig.
Female: similar but smaller than the male, legs not modified.
Endemic to Madagascar. Known from several places in Mahajanga province and near Moramanga (Fig.
Type series probably collected by light trap (
Madagascar, Perinet [= Analamazaotra NP].
Based on single male specimen (holotype). Madagascar, Perinet.
Body shape elongate oval, elytra brown with basal testaceous band, with six discal and one submarginal stria (Fig.
Body length 4.3–5.0 mm. Body shape elongate, weakly oval, some specimens appearing subparalell. Head rufotestaceous anteriorly and posteriorly but infuscated by two blotches inbetween the eyes. Blotches usually meet in middle and form an M-mark, but sometimes infuscation reduced to nearly absent. Pronotum usually with extensive medial infuscation leaving only lateral, anterolateral and posterolateral areas testaceous. Some specimens with less extensive medial infuscation (Fig.
Elytra with six clearly impressed discal and one submarginal stria. Fifth and sixth striae abbreviated anteriorly, and also first stria may be somewhat abbreviated. Submarginal stria starting between one third from the base and midway between base and apex. Head, pronotum, and elytra microreticulate and finely micropunctate. Posterolateral corners of pronotum with strioles that extend along posterior surface, reduced or absent medially.
Ventral side ferrugineous brown, except prothorax, epipleura, appendages, and gula of head which are testaceous. Area around metacoxal lines lighter (not all individuals) and also spots laterally on abdominal sternites II–VII and along some sternite margins are lighter. Metacoxa and abdominal sternites II–IV marked with strioles. Prosternal process short, broad, with blunt apex. Lateral parts of metaventrite broad. Metacoxal lines long, abbreviated only slightly before metaventral margin, diverging anteriorly.
Male: first three pro- and mesotarsomeres widened and ventrally equipped with suction cups; number of suction cups per segment for I–III: 7:4:4 for both pro- and mesotarsus. Protibia modified, narrow with a bisinuate angulate ventral margin at base, broadened distally. Pro- and mesotarsal claws unmodified. Penis in ventral view thin and simple, apical part slightly left-turned. Penis in lateral view angled after basal third forming a “shoulder”. Apex in left lateral view with a characteristic dorsal ridge crossing posterior dorsal margin (Fig.
Female: anterior half of elytra finely striolated from second or third elytral interval to external margin. Degree of elytral striolation in females quite variable between specimens. These strioles are finer than the strioles on pronotum found in both sexes.
Habitus, dorsal view.
Endemic to Madagascar. Known with certainty from the eastern escarpment, Ranomafana NP in the south, Analamazaotra NP, and Mantadia NP in the central region, and Zahamena NP and Andilamena further north (Fig.
We have collected this species in Analamazaotra NP in stagnant forest pools with vegetation or with plentiful of dead leaves. Also found in a sidepool next to a river in semi-open degraded area near Moramanga. The eastern escarpment localities range in altitude between 900–1300 m, but if the localities of Nosy Be and Maroantsetra are correct this species can also occur at lowland sea level altitudes. It seems to be most abundant in tropical eastern forests, but if the Ankarafantsika females belong to this species, then it can also occur in western deciduous forests.
Eventual older records of
Ambohidray reserve, Andriambe [18.61317S, 048.32593E] [Madagascar, Alaotra Mangoro region, Moramanga district].
Similar body shape to
Body length 3.8–4.3 mm. Body shape elongate oval to subparallel. Head rufo-testaceous and only vaguely infuscated medially and around eyes. Pronotum largely rufotestaceous with only faint infuscation medially. Lateral margins lighter testaceous but pronotum also medially lighter than elytra. This gives the habitus appearance of a lighter more rufous anterior part of body contrasting with brown elytra. Elytra brown with a testaceous band basally. Testaceous band narrower and with less of a tendancy to be extended posteriorly in second interval (Fig.
Elytra with six impressed discal and one submarginal striae. First to fourth striae full length or first stria slightly abbreviated at base; fifth and sixth striae abbreviated anteriorly; submarginal stria starting approx. one third to one half from the base. Head, pronotum, and elytra microreticulate and finely micropunctate. Striolation of pronotum rather restricted and present only in posterolateral corners and somewhat inwards along posterior margin but not posteromedially.
Ventral side largely testaceous to faintly infuscated, similar to
Male: first three pro- and mesotarsomeres widened, but less so than in
Female: with very weak, faint and dispersed strioles on anterior half of elytra from third or fourth interval to the lateral margin.
The new species is named after the Malagasy word for small, “kely”, referring to the small body size. It is the smallest species so far known from the
Known from the eastern central part of Madagascar, at Ambohidray Reserve and in Analamazaotra NP (Fig.
This species occurs in the eastern central rainforest. Most specimens were collected from small waterfilled pitfalltrap holes for
This species may be endemic to a very limited area and apart from one specimen found in Analamazaotra NP, we collected the remaining series from the Ambohidray reserve. Most specimens were in fact found in water-filled pitfall trap holes set for a microendemic
Ivohibe Special Reserve [22.456683S, 46.956283E] [Madagascar, Ihorombe region, Ivohibe district]
Body shape slightly shorter, less elongate and slightly more oval than
Body length 3.9–4.5 mm. Body shape elongate oval, but slightly less elongate compared with
Elytra with six discal and one submarginal striae. First to fourth more or less full length, fifth and sixth slightly abbreviated anteriorly; submarginal stria starting approximately one half to one third from base. Striae more deeply impressed and intervals therefore slightly more convex compared to
Ventral side entirely testaceous except metacoxal plate which is variably infuscated, especially laterally in some individuals; metacoxa and abdominal sternites II–IV striolate. Prosternal process with a slightly more pointed apex than in
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups (same constellation as in
Female: very faint to no striolation on outer elytral intervals in the series from Ivohibe. All females from Ranomafana NP were densely striolated over the entire elytral surface except at the apical part, and entire pronotum. Density of striation approx. 5–8 striae in breadth across each elytral interval.
Vokoka is a Malagasy adjective for curvature, also used for an old person with a hunched back. Here it refers to the even curvature of the male aedeagus in lateral view where the even curvature sets it apart from
Known from the mountainous escarpment of southeastern Madagascar at Ivohibe Special Reserve and Ranomafana NP (Fig.
We collected this species in 2013 from forest pools with dead leaves next to streams at the Ivohibe reserve in pristine humid forest at an altitude of 870 m. The reserve of Pic d’Ivohibe was established in 1964 and is managed through collaboration between Madagascar National Parks, local people associations, and other partners. During our visit in 2013, there were little signs of degradation except at the entrance and at the west edge of the reserve. Local people sometimes take zebu cattle with them on a path through the forest. At a larger scale zebu excrement could influence freshwater quality and species assemblages through eutrophication, but there were no signs of this inside the intact pristine forest. In Ranomafana NP it was collected in a shallow clear-water shaded forest pond with large quantity of dead tree leaves and some
Manjakatompo Ankaratra Reserve, Tsiafajavona mountain [19.35163S, 47.24278E] [Madagascar, Vakinankaratra region, Ambatolampy district]
The best diagnostic character for the species is the angled base of penis in posteroventral view (Fig.
Body length 4.4–5.2 mm. Body shape very elongate and subparallel. Head infuscated around eyes and medially, testaceous on clypeus and as a posterior band. Pronotum entirely infuscated except lateral margins and especially the anterolateral corners more broadly testaceous. Elytra in same dark brown colour as infuscated parts of head and pronotum, except an irregular basal testaceous band (Fig.
Habitus, dorsal view.
Elytra with six clearly impressed discal and one submarginal stria. First to fourth elytral striae more or less full length, fifth and sixth striae slightly abbreviated anteriorly; submarginal stria starting 1/3rd to 1/2 posterior of base and does not reach apex. Head, pronotum, and elytra microreticulate and finely micropunctate. Posterior third to posterior half of pronotum striolate. Strioles on average coarser than in
Ventral side similar to
Male: first three pro- and mesotarsomeres widened and ventrally equipped with suction cups (same pattern as in
Female: all but one specimen examined lack finer elytral striolation and is in elytral structure similar to males. However, the entire pronotum and elytra except apical quarter are coarsely longitudinally striolate in one female specimen examined. Striolation coarser and made up of longer irregular but connected strioles and very different to the short separate fine strioles seen in external intervals of
The new species is named after the mountain massif Ankaratra where it was found and in honour of the newly created Ankaratra Massif Reserve in 2015. The epithet is a noun in apposition (ICZN 11.9.1.2).
Known only from a few localities in the Ankaratra Massif Reserve on the central highland plateau of Madagascar (Fig.
This new species was collected in the mountains of Ankaratra at altitudes above 2000 m. The first locality, Anosiarivo forest, consisted of water from a source flowing over grass vegetation at an altitude of 2070 m. The second locality, Tsiafajavona Mountain, was a small stream with grass vegetation downstream but very near to the source at an altitude of 2600 m near Ankaratra peak.
The Ankaratra Massif is an area known for several microendemic species not known from anywhere else on Madagascar. This includes two critically endangered micro-endemic frogs,
Manjakatompo forestry station was established in 1923 in the forested part of the mountains to preserve an area of 8320 ha, out of which only 650 ha is natural forest and 2300 ha has been replanted with exotic trees (
Tsaratanana reserve, Antetikalambazaha Camp [14.1824S, 48.9448E] [Madagascar, Sofia region, Bealanana district].
The best diagnostic character for the species is the pseudostriae between first and second, and second and third striae (Fig.
Body length 5.3–5.6 mm. Body shape elongate oval. Head and pronotum of all three specimens exposed to DNA extraction lysis buffer which has discoloured them slightly. Colour descriptions of head and pronotum below should therefore be taken with caution and can differ somewhat from other specimens. Head and pronotum rather uniformly brown but head likely infuscated between and around eyes (more visible in one paratype) and pronotum may have been infuscated medially prior to exposure to lysis buffer. Elytra uniformly testaceous brown with a faint suggestion of a darker transverse field preapically (Fig.
Elytra with six clearly impressed striae and one submarginal stria. Stria five distincly shorter basally and the submarginal stria starts 1/4th to 1/3rd posterior from the base. Between first and second, and between second and third striae, there are irregular traces of intermediate striae, or “pseudostriae”, extending from just after base until 1/4th from the apex (Fig.
Ventral side largely ferrugineous, a little lighter testaceous-ferrugineous around metacoxal processes, medially on the metaventrite and on sternite II. Prosternal process short, medially raised and rounded and with a fairly pointed apex. Lateral parts of metaventrite medium broad. Metacoxal lines anteriorly diverging and abbreviated well before metaventral margin. Metacoxal plate distinctly striolate with short strioles.
Male: first three pro- and mesotarsomeres widened. Protibia modified, narrow but not bisinuate with an angulate ventral margin at base, broadened, almost club-like, distally. Protarsal and mesotarsal claws unmodified. Penis in ventral view narrowed one third from apex and the very last apical tip angled to the left; in lateral view evenly and weakly curved from base to apex (Fig.
Female: elytral structure similar to male.
The name
Endemic to Madagascar, only known from the type series from Tsaratanana Massif (Fig.
Not known, but the type series of specimens were collected in 2004 likely from a stream, near Antetikalambazaha Camp at an altitude of 1700 m.
Species group assignment of
Anjanaharibe Sud reserve, [14.7414S, 049.4975E] [Madagascar, Sava region, Andapa district]
Somewhat similar to
Body length 4.3–5.2 mm. Body shape elongate and subparallel along a very long part of the body. Pronotum and head broad and eyes small creating a very wide interocular distance. Maximum width of pronotum clearly in front of hind corners. Head rufotestaceous with weak or absent infuscation in between and posterior of eyes. Pronotum infuscated medially with broadly testaceous lateral sides. Elytra brown, with a broad testaceous transverse band basally (Fig.
Elytra with six discal and one submarginal stria. Fifth stria abbreviated anteriorly, variably also first and third striae. In some individuals especially fifth but also first and sixth striae are rudimentary or with very shallow impressions. Interval between fifth and sixth striae narrow, approx. half interval between first and second striae. Submarginal stria short, starting around middle. Head, pronotum, and elytra microreticulate and finely micropunctate. Pronotum not striolate.
Ventral side entirely testaceous except last three abdominal ventrites may be vaguely infuscated and have lighter lateral spots. Metacoxa and abdominal sternite II–IV striolate, but strioles shallower and finer than in
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups. Pattern of suction cups same as for
Female: on average smaller than males (Table
Measurements of body length summarised as Min, Max, Mean, and Standard Deviation (SD) for each species, separated by sex. N = number of measured individuals, F = females, M = males.
Species | Sex | N | Min | Max | Mean | SD |
---|---|---|---|---|---|---|
|
F | 4 | 4.13 | 4.19 | 4.15 | 0.03 |
M | ||||||
|
F | 26 | 5.16 | 6.32 | 5.75 | 0.30 |
M | 45 | 5.48 | 6.58 | 6.02 | 0.26 | |
|
F | 12 | 5.16 | 6.13 | 5.63 | 0.24 |
M | 16 | 5.42 | 6.19 | 5.88 | 0.20 | |
|
F | 1 | 5.74 | 5.74 | 5.74 | |
M | 3 | 5.48 | 6.06 | 5.83 | 0.30 | |
|
F | 44 | 5.29 | 6.32 | 5.75 | 0.21 |
M | 26 | 5.48 | 6.26 | 5.92 | 0.18 | |
|
F | 7 | 5.74 | 6.58 | 6.05 | 0.27 |
M | 7 | 5.81 | 6.26 | 6.10 | 0.16 | |
|
F | 1 | 4.00 | 4.00 | 4.00 | |
M | ||||||
F | 7 | 3.81 | 4.26 | 4.01 | 0.16 | |
M | 5 | 3.87 | 4.32 | 4.03 | 0.17 | |
|
F | 27 | 4.26 | 5.03 | 4.60 | 0.20 |
M | 24 | 4.26 | 4.90 | 4.61 | 0.17 | |
F | 3 | 4.32 | 4.71 | 4.58 | 0.22 | |
M | 13 | 4.52 | 5.16 | 4.86 | 0.19 | |
F | 6 | 3.94 | 4.32 | 4.18 | 0.13 | |
M | 8 | 4.06 | 4.45 | 4.29 | 0.12 | |
F | 19 | 4.39 | 5.03 | 4.70 | 0.17 | |
M | 30 | 4.65 | 5.16 | 4.87 | 0.16 | |
F | 2 | 5.42 | 5.61 | 5.52 | 0.14 | |
M | 1 | 5.29 | 5.29 | 5.29 |
The species name
Endemic to Madagascar. This species has a rather large distribution in the eastern humid forest from Anjanaharibe Sud reserve in the NE, all along the eastern escarpment including Zahamena NP, Mantadia NP, and Ranomafana NP, and even extending to the rather isolated western patch of subhumid forest at Isalo NP (Fig.
This species seems to be strongly associated with clean streams having sandy substrate in humid forests. At these localities, individuals can be found in sidepools, at margins or sites protected from waterflow (e.g., by fallen logs) where dead leaves and debris accumulate. The species has been found at altitudes between 700 and 1300 m but most numerous at elevations above 900 m in primary humid forest. The discovery in Isalo NP, in a sandy river running through a Canyon, indicates that subhumid forests may also be part of the species’ niche.
We are aware of several
[
This species is broad and oval in body shape and belongs to the
It is plausible that it was one or several female specimens of this species that
Distribution maps of
This species has a configuration of elytral striae not found in any of the other species treated here. It has five discal striae and is lacking a submarginal stria and would fall in the
The DNA data revealed that these females represent one or possibly two additional new species in the
Distribution maps of
Madagascar is known for an extremely rich endemic flora and fauna which, together with the unfortunate level of deforestation, has rewarded the island with a top spot among the world’s biodiversity hotspots (
The
We are grateful to the “Ministère de l’Environnement, de l’Ecologie et des Forêts” (MEEF) and the Madagascar National Parks (MNP) for permits, access and continuous long-term support over the years. We thank collection curators Antoine Mantilleri (
Note that on the continent this species may have a basal testaceous band although we have yet not seen this colour form from Madagascar. It is not impossible that this colour form may show up eventually also in Madagascar, which is why the shape of male genitalia should always be verified for identification
Examined specimens, species occurrence records
species data