Research Article |
Corresponding author: Tolotra Ranarilalatiana ( rtolotr@yahoo.fr ) Academic editor: Mariano Michat
© 2019 Tolotra Ranarilalatiana, Lala Harivelo Raveloson Ravaomanarivo, Johannes Bergsten.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ranarilalatiana T, Raveloson Ravaomanarivo LH, Bergsten J (2019) Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species. ZooKeys 869: 19-90. https://doi.org/10.3897/zookeys.869.33997
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The genus Copelatus Erichson, 1832 (Coleoptera, Dytiscidae, Copelatinae) of Madagascar is revised in two parts. This review is restricted to the Copelatus species that have fewer than ten elytral + one submarginal stria, including all species except those of the erichsonii species group. Both morphological and molecular (mitochondrial COI) data are used in an integrative taxonomic approach. Thirteen species are recognised, of which five are described as new: Copelatus ankaratra sp. nov., Copelatus kely sp. nov., Copelatus pseudostriatus sp. nov., Copelatus safiotra sp. nov. and Copelatus vokoka sp. nov. Copelatus unguicularis Régimbart, 1903 and Copelatus apicalis Fairmaire, 1898 are both transferred to the genus Madaglymbus Shaverdo & Balke, 2008 (comb. nov.). Copelatus mimetes
Conservation, distribution, diving beetles, freshwater, gene tree, GMYC, new combination, new species, new synonymy, phylogeny, species delimitation
The subfamily Copelatinae of diving beetles is a diverse group of aquatic beetles represented by eight genera: Agaporomorphus Zimmermann, 1921, Aglymbus Sharp, 1880, Capelatus Turner & Bilton, 2015, Copelatus Erichson, 1832, Exocelina Broun, 1886, Lacconectus Motschulsky, 1855, Liopterus Dejean, 1833, and Madaglymbus Shaverdo & Balke, 2008. This subfamily is relatively homogeneous morphologically, and only the single tribe Copelatini is recognised (
To be able to deal with this diversity, Copelatus species have traditionally been organised in a number of species groups based on the number of elytral and submarginal striae following
The species that occur in Madagascar fall into five of these species groups: the hydroporoides (= formerly haemorrhoidalis, no elytral striae), longicornis (3, 4 or 5 elytral striae), irinus (6 elytral + 1 submarginal stria), consors (10 elytral striae) and erichsonii (10 elytral + 1 submarginal stria) groups. About half of the described species are members of the erichsonii species group, and the other half of the remaining species groups with fewer striae. In this paper, we will treat the Copelatus species of Madagascar with fewer than 10 elytral and one submarginal stria, i.e., all the non-erichsonii group species. Again, this is a practical rather than a phylogenetic division and in fact Copelatus safiotra sp. nov., described below, we believe is most closely related to the species of the erichsonii group due to the shape of male genitalia.
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 irinus species group, together with type material and some additional museum accessions. We use both morphological and molecular data with a species delimitation analysis, in an integrative taxonomic approach to the revision. We provide an identification key and for each species a diagnosis, description, known distribution and habitat and ecology notes. Each species is also illustrated with a dorsal habitus photograph, and ventral and lateral views of male penis and parameres.
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.
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 Museum National d’Histoire Naturelle, Paris (
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 “
Alc. in alcohol tube,
Ex. exemplars (number of individuals),
GP (Genital Preparation) male genitalia have been examined,
HT Holotype,
LT Lectotype,
PLT Paralectotype,
PT Paratype,
ST Syntype,
TL Type locality,
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 cox1) (
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 Copelatus from
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 |
---|---|---|---|---|---|---|---|---|
C. marginipennis | BMNH-797876 | 294:A4 | P57BI31 | Marojejy NP | 14.4573S, 49.7908E | 162 | 10/12/07 | HQ382912 |
BMNH-797894 | 294:B10 | P61BI15 | Andasibe NP | 18.9375S, 48.4167E | 940 | 06/01/07 | HQ382926 | |
BMNH-797906 | 294:C10 | P58BI14 | Masoala NP, E. of Marofototra | 15.7587S, 49.9932E | 10 | 17/12/06 | HQ382937 | |
BMNH-797907 | 294:C11 | P58BI14 | Masoala NP, E. of Marofototra | 15.7587S, 49.9932E | 10 | 17/12/06 | HQ382938 | |
BMNH-797908 | 294:C12 | P58BI14 | Masoala NP, E. of Marofototra | 15.7587S, 49.9932E | 10 | 17/12/06 | HQ382939 | |
BMNH-797909 | 294:D01 | P58BI14 | Masoala NP, E. of Marofototra | 15.7587S, 49.9932E | 10 | 17/12/06 | HQ382940 | |
BMNH-797910 | 294:D02 | P58BI14 | Masoala NP, E. of Marofototra | 15.7587S, 49.9932E | 10 | 17/12/06 | HQ382941 | |
|
JB196 | MAD09-07 | Ankarafantsika NP, Ampijoroa | 16.3034S, 46.8107E | 74 | 29/11/09 | MK878864 | |
|
JB197 | MAD09-46 | Kirindy Res. | 20.0743S, 4.6631E | 52 | 12/12/09 | MK878871 | |
|
JB198 | MAD09-13 | Ankarafantsika NP, Ampijoroa | 16.3027S, 46.8100E | 75 | 30/11/09 | MK878865 | |
|
JB199 | MAD09-07 | Ankarafantsika NP, Ampijoroa | 16.3034S, 46.8107E | 74 | 29/11/09 | MK878869 | |
|
JB200 | MAD09-59 | Tsingy de Bemaraha NP, Bekopaka | 19.0342S, 44.7750E | 41 | 15/12/09 | MK878872 | |
|
JB201 | MAD09-03 | Ankarafantsika NP, Ampijoroa | 16.3035S, 46.8107E | 87 | 29/11/09 | MK878870 | |
|
JB202 | MAD09-29 | Mahavavy Kinkony Res., Mitsinjo | 16.0665S, 45.7767E | 24 | 05/12/09 | MK878866 | |
|
JB203 | MAD09-65 | Tsingy de Bemaraha NP, Antsalova | 18.7564S, 44.7140E | 119 | 17/12/09 | MK878873 | |
|
JB191 | MAD09-14 | Ankarafantsika NP, Ampijoroa | 16.3142S, 6.8173E | 77 | 30/11/09 | MK878862 | |
|
JB192 | MAD09-24 | Mahavavy Kinkony Res., Makary village | 16.1465S, 45.9493E | 9 | 04/12/09 | MK878863 | |
|
JB193 | MAD09-25 | Mahavavy Kinkony Res., Makary village | 16.1334S, 45.9578E | 19 | 04/12/09 | MK878874 | |
|
JB194 | MAD09-28 | Mahavavy Kinkony Res., Mitsinjo | 16.0578S, 5.8059E | 22 | 05/12/09 | MK878868 | |
|
JB189 | MAD09-30 | Mahavavy Kinkony RS, Mitsinjo | 16.0565S, 45.7637E | 55 | 05/12/09 | MK878867 | |
|
JB809 | MAD09-24 | Mahavavy Kinkony RS, Makary village | 16.1465S, 45.9493E | 9 | 04/12/09 | MK878861 | |
C. kely sp. nov. |
|
TR18L14 | Ambohidray Res., Andriambe | 18.6132S, 8.3259E | 1044 | 23/05/18 | MK878839 | |
|
TR18L04 | Ambohidray Res., Andriambe | 18.6132S, 48.3262E | 1044 | 07/04/18 | MK878842 | ||
|
TR18L04 | Ambohidray Res., Andriambe | 18.6132S, 48.3262E | 1044 | 07/04/18 | MK878843 | ||
|
TR18L07 | Ambohidray Res., Andriambe | 18.6131S, 48.3257E | 1046 | 07/04/18 | MK878844 | ||
C. befasicus |
|
JB204 | MAD09-74 | btw Morafenobe-Ambohijanahary Res. | 18.1909S, 45.1999E | 290 | 19/12/09 | MK878825 |
Copelatus sp. ♀ (Bemaraha) | CASENT-8135000 | BLF4432 | Tsingy de Bemaraha NP | 19.1323S, 4.8147E | 150 | 16/11/01 | MK878860 | |
C. distinguendus | BMNH-670601 | 007:E07 | P27MD31 | Ranomafana NP | 21.2359S, 47.3963E | 1123 | 06/12/04 | HQ381662 |
BMNH-729896 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 47.1002E | 1442 | 08/12/04 | HQ381870 | ||
BMNH-792954 | P39EM08 | Andringitra NP | 22.1043S, 6.9207E | 1420 | 09/05/06 | HQ382583 | ||
BMNH-792955 | P39EM08 | Andringitra NP | 22.1043S, 46.9207E | 1420 | 09/05/06 | HQ382584 | ||
BMNH-792956 | P39EM08 | Andringitra NP | 22.1043S, 46.9207E | 1420 | 09/05/06 | HQ382585 | ||
BMNH-792962 | P36C | RN7, Col de Tapias | 20.7729S, 47.1792E | 1717 | 06/05/06 | HQ382591 | ||
BMNH-792963 | P36C | RN7, Col de Tapias | 20.7729S, 47.1792E | 1717 | 06/05/06 | HQ382592 | ||
BMNH-792964 | P36C | RN7, Col de Tapias | 20.7729S, 7.1792E | 1717 | 06/05/06 | HQ382593 | ||
BMNH-792976 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 47.1002E | 1442 | 08/12/04 | HQ382604 | ||
BMNH-792977 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 47.1002E | 1442 | 08/12/04 | HQ382605 | ||
BMNH-792978 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 47.1002E | 1442 | 08/12/04 | HQ382606 | ||
BMNH-792979 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 47.1002E | 1442 | 08/12/04 | HQ382607 | ||
BMNH-792980 | P30MD33 | Sahatsiho Ambohimanjaka | 20.2388S, 7.1002E | 1442 | 08/12/04 | HQ382608 | ||
|
MAD14-81 | RN2, Betsabora river | 18.9247S, 48.1828E | 900 | 24/11/14 | MK878834 | ||
|
MAD16-47 | Manjakatompo Ankaratra Res., Ankafotra Mtn. | 19.3375S, 47.2453E | 2466 | 18/09/16 | MK878835 | ||
C. insuetus |
|
MAD14-81 | RN2, Betsabora river | 18.9247S, 8.1828E | 900 | 24/11/14 | MK878836 | |
|
MAD18-91 | Zahamena NP, Sect. Antanandava | 17.5225S, 48.7227E | 1040 | 08/03/18 | MK878841 | ||
|
MAD11-26 | Analamazaotra NP | 18.9357S, 48.4174E | 930 | 08/11/11 | MK878840 | ||
|
MAD14-18 | Analamazaotra NP | 18.9357S, 48.4174E | 930 | 27/11/14 | MK878837 | ||
BMNH-797895 | P60BI15 | Zahamena NP, Sect. Antanandava | 17.52S, 48.721E | 1075 | 31/11/06 | HQ382927 | ||
C. safiotra sp. nov. |
|
MAD11-37 | Mantadia NP | 18.8340S, 8.4378E | 1000 | 11/11/11 | MK878829 | |
|
MAD14-70 | Anjanaharibe Sud res. | 14.7414S, 49.4975E | 910 | 16/11/14 | MK878833 | ||
|
MAD14-04 | Ranomafana NP | 21.2395S, 47.3947E | 1130 | 02/11/14 | MK878830 | ||
|
MAD12-03 | Isalo NP, Canyon des Makis | 22.4866S, 45.3797E | 700 | 13/11/12 | MK878831 | ||
|
MAD12-03 | Isalo NP, Canyon des Makis | 22.4866S, 45.3797E | 700 | 13/11/12 | MK878832 | ||
|
MAD13-55 | Ivohibe RS | 22.4567S, 46.9563E | 874 | 09/12/13 | MK878845 | ||
|
MAD13-55 | Ivohibe RS | 22.4567S, 46.9563E | 874 | 09/12/13 | MK878846 | ||
C. mahajanga |
|
MAD14-81 | RN2, Betsabora river | 18.9247S, 48.1828E | 900 | 24/11/14 | MK878826 | |
|
MAD14-81 | RN2, Betsabora river | 18.9247S, 48.1828E | 900 | 24/11/14 | MK878827 | ||
|
JB190 | MAD09-58 | Tsingy de Bemaraha NP, Bekopaka | 19.0357S, 44.7751E | 66 | 15/12/09 | MK878875 | |
|
JB195 | MAD09-33 | Mahavavy Kinkony Res., Anjohibe | 16.0133S, 46.0038E | 24 | 06/12/09 | MK878876 | |
|
JB802 | MAD09-25 | Mahavavy Kinkony Res., Makary village | 16.1334S, 45.9578E | 19 | 04/12/09 | MK878828 | |
C. ankaratra sp. nov. |
|
MJK12-13 | Manjakatompo Ankaratra Res., Anosiarivo | 19.3449S, 47.3041E | 2073 | 24/01/12 | MK878851 | |
|
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. | 19.3516S, 47.2428E | 2597 | 07/02/16 | MK878847 | ||
|
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. | 19.3516S, 47.2428E | 2597 | 07/02/16 | MK878848 | ||
|
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. | 19.3516S, 47.2428E | 2597 | 07/02/16 | MK878849 | ||
|
MAD16-11 | Manjakatompo Ankaratra Res., Tsiafajavona Mtn. | 19.3516S, 47.2428E | 2597 | 07/02/16 | MK878850 | ||
C. pulchellus |
|
JB808 | MAD09-25 | Mahavavy Kinkony RS, Makary village | 16.1334S, 45.9578E | 19 | 04/12/09 | MK878859 |
|
MAD14-14 | Analamazaotra NP | 18.9355S, 48.4166E | 930 | 27/11/14 | MK878856 | ||
|
MAD14-14 | Analamazaotra NP | 18.9355S, 48.4166E | 930 | 27/11/14 | MK878857 | ||
Copelatus sp. ♀ (Andasibe) |
|
Analamazaotra NP, Andasibe | 18.94S, 48.43E | 938 | 17/01/15 | MK878858 | ||
Copelatus ? insuetus ♀ (Ankaraf.) |
|
JB206 | MAD09-07 | Ankarafantsika NP, Ampijoroa | 16.3034S, 46.8107E | 74 | 29/11/09 | MK878877 |
|
JB205 | MAD09-03 | Ankarafantsika NP, Ampijoroa | 16.3035S, 46.8107E | 87 | 29/11/09 | MK878838 | |
Copelatus sp. ♀ (Ivohibe) |
|
MAD13-61 | Ivohibe RS, Andranovory | 22.4751S, 46.9559E | 1106 | 10/12/13 | MK878854 | |
|
MAD13-61 | Ivohibe RS, Andranovory | 22.4751S, 46.9559E | 1106 | 10/12/13 | MK878855 | ||
Copelatus sp. ♀ (N Toam.) |
|
MAD11-52 | RN5, Ivoloina | 18.0649S, 49.3786E | 0 | 15/11/15 | MK878852 | |
|
MAD17-12 | Analalava Res., Analalava forest | 17.7106S, 49.4500E | 39 | 09/03/17 | MK878853 | ||
C. pseudostriatus sp. nov. | BMNH-672727 | 027:A05 | P32 | Tsaratanana massif, Mangindrano | 14.1824N, 48.9448E | 1700 | 20/12/04 | HQ381767 |
BMNH-672728 | 027:A06 | P32 | Tsaratanana massif, Mangindrano | 14.1824N, 48.9448E | 1700 | 20/12/04 | HQ381768 | |
BMNH-672729 | 027:A07 | P32 | Tsaratanana massif, Mangindrano | 14.1824N, 48.9448E | 1700 | 20/12/04 | HQ381769 |
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 Copelatus befasicus. We used PARTITIONFINDER Ver. 2.1.1 (
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 C. insuetus and C. kely sp. nov. (Fig.
Uncorrected P genetic distances between closely related species or populations in the group near Copelatus insuetus. Numbers in parenthesis are genetic distances calculated under a Kimura-2-parameter model. NA = Not Applicable because of single sample in category.
C. insuetus | C. insuetus? (Ankaraf.) | C. kely | C. ankaratra (peak) | C. ankaratra | C. vokoka | C. sp. female (Ivohibe) | C. sp. female (N. Toam.) | |
---|---|---|---|---|---|---|---|---|
C. insuetus | 0–0.007 | |||||||
(0–0.007) | ||||||||
C. insuetus ? | 0.005–0.009 | 0 | ||||||
(Ankaraf.) | (0.005–0.009) | (0) | ||||||
C. kely | 0.001–0.017 | 0.006–0.011 | 0–0.017 | |||||
(0.001–0.017) | (0.006–0.011) | (0–0.017) | ||||||
C. ankaratra | 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) | ||||
C. ankaratra | 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 | ||||
C. vokoka | 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) | |||
C. sp. female | 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) | |
C. sp. female | 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 Copelatus species of Madagascar with fewer than ten discal and one submarginal elytral striae. Note that for some species males are necessary.
1 | Impressed elytral striae absent (rows of points may be present) (Fig. |
2 (hydroporoides group) |
– | Impressed elytral striae present (Figs |
3 |
2 | Body length less than 4 mm (Fig. |
C. baculiformis |
– | Body length between 5.7 and 6.6 mm (Fig. |
C. peridinus |
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. |
C. befasicus (longicornis group) |
– | Elytra with six discal and one submarginal stria (Figs |
4 (irinus group) |
4 | First, third, and fifth elytral striae distinctly abbreviated anteriorly; dorsal colouration largely black (Fig. |
C. distinguendus |
– | Third and fifth elytral striae not abbreviated anteriorly; first elytral stria abbreviated or not; dorsal colouration rarely mostly black; penis shape variable | 5 |
5 | Body shape more broadly oval (Fig. |
6 |
– | Body shape narrow, elongate, subparalell (Figs |
8 |
6 | Elytra without a transverse testaceous band at base, black except laterally and posteriorly (Fig. |
C. pulchellus |
– | Elytra with basal testaceous transverse band; penis apex in ventral view right-curved | 7 |
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. |
C. marginipennis |
– | 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. |
C. mahajanga |
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. |
C. pseudostriatus sp. nov. |
– | Elytra without pseudostriae in elytral intervals; body length smaller, less than 5.3 mm; Penis curvature in lateral view stronger | 9 |
9 | Body subparallell and head with a broad interocular distance; pronotum without strioles (Fig. |
C. safiotra sp. nov. |
– | Body more attenuating anteriorly and posteriorly and interocular distance narrower; pronotum with strioles posterolaterally; penis slender, without subapical expansion in lateral view | 10 |
10 | Dark colouration dorsally, with narrow testaceous band at base of elytra and testaceous, sometimes strongly contrasting, anterolateral pronotal corners; head distinctly infuscated (Fig. |
C. ankaratra sp. nov. |
– | 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 |
11 | Body length smaller, 3.8 to 4.3 mm (Fig. |
C. kely sp. nov. |
– | Body length larger, 3.9 to 5.0 mm; penis in lateral view with a dorsal ridge crossing posterior inner outline near apex | 12 |
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. |
C. insuetus |
– | Elytra often with broader basal testaceous band; penis in lateral view evenly curved without a distinct shoulder definition (Fig. |
C. vokoka sp. nov. |
The following three taxa were described as Copelatus species from Madagascar and were listed as such by
Copelatus subjectus Sharp, 1882: 568.
Copelatus bilunatus
Guignot, 1955: 73; TL: Madagascar [mislabelled, likely New Caledonia; see
New Caledonia.
Copelatus bilunatus Guignot, 1955 is a synonym of Exocelina subjecta (Sharp, 1882) following
Copelatus apicalis Fairmaire, 1898: 465.
Madagascar, Suberbieville [= Maevatanana].
Mahajanga. Betsiboka: Maevatanana: -ST ♂ (GP) (
Copelatus apicalis was described by
Copelatus unguicularis Régimbart, 1903:19.
Madagascar, Suberbieville [= Maevatanana].
Mahajanga. Betsiboka: Maevatanana: -HT ♂ (GP) (
Copelatus unguicularis was described by
This group is defined by the lack of elytral striae (
Copelatus baculiformis Guignot, 1955b: 193.
Madagascar, Massif Ankaratra, Manjakatompo, alt. 1700–1800 m.
based on a single female specimen (holotype), collected December 1951 by R. Benoist.
Antananarivo. Vakinankaratra: Ambatolampy: -HT ♀ (
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.
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 hydroporoides species group of Copelatus, but two other Malagasy species placed in this group have turned out to be misidentified Madaglymbus or Exocelina (see above). Copelatus baculiformis was described by Guignot the same year (1955) that he described C. bilunatus, considered mislabeled (
Copelatus peridinus Guignot, 1955c: 188.
Copelatus seydeli Guignot, 1958: 107; TL: Elisabethville, Zaire [DR Congo, Haut-Katanga, Lubumbashi].
Elisabethville, Zaire [DR Congo, Haut-Katanga, Lubumbashi].
-PT ♀ (
-1♂(GP) (
Similar to C. baculiformis based on shape and colouration, but C. peridinus is bigger with body length between 5.7 and 6.6 mm (Fig.
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 Copelatus sp. 2 below).
Copelatus befasicus Guignot, 1956: 79.
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.
Toliara. Menabe: Morondava: -HT♀ (
Mahajanga. Melaky: Morafenobe: -1♀ (
Similar to C. insuetus and related species in habitus by being small, elongate, and subparalell, but C. befasicus is distinguished from C. insuetus and from all other Malagasy Copelatus species by the presence of only five elytral striae, and without submarginal striae. In addition, the first stria is shortened, present only in the posterior third (Fig.
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 C. befasicus also in the western part but a bit further north than the type locality, along the road between Morafenobe and Beravina village at an altitude of 290 m. This locality consisted of dry savannah with mixed wood and grassland ecosystem after deforestation. The habitat consisted of muddy/sandy residual pools with some dead leaves in a temporary stream after the rainy season. The dry deciduous forest ecosystem in western Madagascar has suffered immensely from deforestation and very little of this habitat remains (
This is the only species in the Copelatus longicornis group from Madagascar. The longicornis species group currently contains 38 species distributed mainly in the Neotropical and Afrotropical regions but also with few species present in Japan, New Guinea, and Fiji Islands (
The irinus group is characterised by the presence of six discal and one submarginal elytral striae (
Copelatus distinguendus
Régimbart, 1903: 19 [nom. nov., referring to his description of Malagasy material under the name Copelatus duodecimstriatus Aubé in
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
Fianarantsoa. Matsiatra Ambony: Ambalavao, Ambohimahasoa, Lalangina: -2♀, 1♂(GP) (
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.
Agabus pulchellus
Copelatus africanus Sharp, 1882: 583; TL: Botswana, Lake Ngami;
? Copelatus basalis Boheman, 1848: 244; TL: South Africa (Caffraria interiore);
Copelatus discoideus Sharp, 1882: 582; TL: Mesopotamia;
Copelatus obtusus Boheman, 1848: 242; TL: South Africa (Caffraria orientali);
Copelatus strigulosus Sharp, 1882: 582; TL: Mesopotamia;
Copelatus mimetes Guignot, 1957: 73; TL: Madagascar, Sakaraha, Lambomakandro; syn. nov.
Egypt, Sinai.
-HT♂ (GP) (Copelatus mimetes) (
Antsiranana. Diana: Antsiranana: -1♂(GP) (
Similar to C. marginipennis (Laporte, 1835) and C. mahajanga in overall habitus, but body shape more like the elongated shape of C. mahajanga. Copelatus pulchellus on Madagascar is ferrugineous black in overall colouration with or without a rather narrow or vague testaceous band basally on elytra (Fig.
(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 C. marginipennis and C. mahajanga. Lateral parts of metaventrite medium broad. Metacoxal lines short and rather strongly diverging anteriorly. Antennae, palps, pro- and mesolegs testaceous, metalegs somewhat darker testaceous.
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.
As the species C. pulchellus is currently interpreted, this is a very widely distributed Afrotropical and Middle Eastern 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.
Copelatus pulchellus forms part of a diverse species group with many externally very similar species.
Copelatus pulchellus is now interpreted as a widespread Afrotropical and Middle Eastern species with the male penis similar to that illustrated in Figure
Copelatus basalis Boheman, 1848 was synonymised with C. pulchellus by Omer-Cooper (1965). We have studied the same type material as Omer-Cooper and agree with this conclusion. However, we are not convinced that the material housed at
Colymbetes marginipennis Laporte, 1835: 102.
Copelatus aldabricus J. Balfour-Browne, 1950: 368 syn. nov.; TL: Seychelles, Aldabra Islands.
Copelatus aldabricus var. simplex
Senegal [possibly mislabelled].
housed in Buquet collection and originating from Senegal; of aldabricus: based on male (holotype), J.C. Fryer collection, collected 1908-9 from Aldabra; of simplex: based on male and female syntypes from Madagascar without further locality data.
-LT♂ (lectotype here designated) (Colymbetes marginipennis) (
Toamasina. Alaotra Mangoro: Andilamena, Moramanga: -1♀ (
Copelatus marginipennis is distinguished from all other Malagasy Copelatus except C. pulchellus and C. mahajanga by the presence of six discal elytral striae and a broadly oval body shape. Copelatus marginipennis is most easily separated by the distinct shape of the male penis in lateral view (Fig.
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 C. mahajanga in Betsabora river near Moramanga and at a 22W black light trap by a forest pool in Ankarafantsika NP.
Copelatus aldabricus was described by J.
Copelatus marginipennis (Laporte, 1835) has been treated as a junior synonym of Copelatus pulchellus Klug, 1834 since
Copelatus mahajanga Pederzani & Hájek, 2005: 104.
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. HT & 5PT in
Mahajanga. Boeny: Mahajanga I/Mahajanga II [district cannot be verified based on original description or labels]: -HT♂(GP) (
Mahajanga. Boeny: Ambato-Boeny, Mitsinjo, Soalala: -3♂ (GP), 1♀ (
Close to C. marginipennis in habitus appearance but body shape more elongate. On average C. mahajanga have a broader transverse testaceous band at base of elytra (Fig.
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 C. marginipennis but lateral parts of metaventrite slightly narrower and metacoxal lines less divergent anteriorly. Antennae, palps, pro- and mesothoracic legs brown, but metathoracic legs ferrugineous dark brown.
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 (
Copelatus mahajanga was the most recently described Copelatus species from Madagascar (Peredzani and Hájek 2005) and was previously only known from the type series and lowland west type locality “Mahajanga env.” without further details. The discovery of the species in Betsabora River at mid-altitude in the east indicates that the species may have a much wider distribution and ecological niche. As the species is very similar to C. marginipennis habitually, it might be misidentified as such in collections if male genitalia were not examined.
Copelatus insuetus Guignot, 1941: 39.
Madagascar, Perinet [= Analamazaotra NP].
Based on single male specimen (holotype). Madagascar, Perinet.
Toamasina. Alaotra Mangoro: Moramanga: -HT♂ (GP) (
Toamasina. Alaotra Mangoro: Moramanga, Ambatondrazaka, Andilamena: -2♀ (
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.
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 C. insuetus should be regarded with caution in light of the habitually very similar new species described below.
Ambohidray reserve, Andriambe [18.61317S, 048.32593E] [Madagascar, Alaotra Mangoro region, Moramanga district].
Toamasina. Alaotra Mangoro: Moramanga: -HT♂(GP) (
Similar body shape to C. insuetus but C. kely is smaller than all other species in the Copelatus insuetus complex on Madagascar; body length 3.8–4.4 mm (Fig.
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 C. insuetus but lighter. Prosternal process and lateral parts of metaventrite similar to C. insuetus but metacoxal lines less strongly diverging anteriorly.
Male: first three pro- and mesotarsomeres widened, but less so than in C. insuetus, and ventrally equipped with suction cups (same constellation as in C. insuetus). Protibia modified, bisinuate and angled basally and broadened distally. Pro- and mesotarsal claws unmodified. Penis thin and simple, in ventral view with apical part slightly leftturned; in lateral view slightly angled after basal third giving a suggested “shoulder” (Fig.
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 C. insuetus species complex on Madagascar. It is a non-latinised adjective.
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 Mantella frogs, but we’ve also found it in muddy, stagnant, forest pools with dead leaves. The altitude of known localities ranges from 930 to 1050 m. At Analamazaotra NP, we found one specimen occurring sympatrically with Copelatus insuetus at the same locality.
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 Mantella frog species. Copelatus kely adds to the importance of this reserve for conservation of rare and microendemic eastern rainforest species. Ambohidray reserve was established in 2013 and managed through collaboration between the local people association (VOI MMA) and Antananarivo University. During fieldwork at the reserve in April 2018 however, we observed worrying signs of disturbances; “tavy” or slash and burn of the forest for agriculture, the cutting of woods for charcoal, signs of zebu-cattle along forest paths. These factors could cause a serious threat to the aquatic insect fauna of the reserve. The reserve of Ambohidray harbours some species not known from anywere else on Madagascar. If the reserve has any ambition to serve as a refugium for these species, activities destroying or degrading the forests or aquatic habitats should be avoided. Copelatus kely is very close to C. insuetus, and the two species were not reciprocally monophyletic in the CO1 gene tree (Fig.
Ivohibe Special Reserve [22.456683S, 46.956283E] [Madagascar, Ihorombe region, Ivohibe district]
Fianarantsoa. Ihorombe: Ivohibe: -HT♂ (GP) (
Body shape slightly shorter, less elongate and slightly more oval than C. insuetus, and eyes smaller. Elytral striae more deeply impressed and intervals therefore slightly more convex (Fig.
Body length 3.9–4.5 mm. Body shape elongate oval, but slightly less elongate compared with C. insuetus. Head rufotestaceous with only a faint suggesstion of an M-shaped infuscation between eyes. Pronotum rufotestaceous as the head, with weak medial infuscation Elytra dark brown, with testaceous band at base. Testaceous band generally broader and more diffusely transitioning into the darker elytral colour posteriorly (Fig.
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 C. insuetus. Head, pronotum, and elytra microreticulate and finely micropunctate. Pronotum extensively and coarsely striated on posterior surfaces, although reduced posteromedially. On some specimens striolation covers most of pronotum, also on anterior surfaces, but is reduced medially.
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 C. insuetus and C. kely, and lateral parts of metaventrite not as broad. Metacoxal lines long and not strongly diverging, as in C. kely.
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups (same constellation as in C. insuetus). Protibia modified, bisinuate and angled basally and broadened distally. Pro- and mesotarsal claws unmodified. Penis long, thin and simple; apex in ventral view straight and not leftturned; in lateral view rather evenly arched, lacking the distinct “shoulder” characteristic of C. insuetus but with a different type of postmedial and preapical suggested humps in the curvature. Apex in lateral view also broder closer to apex. Apex in left lateral view with a dorsal ridge crossing posterior dorsal margin but at a more acute angle (Fig.
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 C. insuetus. It is a non-latinised adjective.
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 Poaceae (M. Manuel pers. comm.).
Copelatus vokoka sp. nov. falls in the irinus group, based on the number of elytral striae. The new species is most closely related to C. ankaratra according to the mitochondrial gene CO1 and belongs to the C. insuetus species complex on Madagascar.
Manjakatompo Ankaratra Reserve, Tsiafajavona mountain [19.35163S, 47.24278E] [Madagascar, Vakinankaratra region, Ambatolampy district]
Antananarivo. Vakinankaratra: Ambatolampy: -HT♂ (GP) (
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.
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 C. insuetus and in some specimens more extensive onto disc and posteromedially.
Ventral side similar to C. insuetus, slightly darker on average so that medial light area around metacoxal lines may be more contrasting. Strioles on metacoxa rather short. Prosternal process more rhomboid and apex more strongly raised medially than in C. insuetus. Lateral parts of metaventrite rather broad. Metacoxal lines long and anteriorly diverging, similar to C. insuetus.
Male: first three pro- and mesotarsomeres widened and ventrally equipped with suction cups (same pattern as in C. insuetus). Protibia modified, bisinuate, angled basally, and broadened distally. Pro- and mesotarsal claws unmodified. Penis in posteroventral view distinctly angled at base (Fig.
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 C. insuetus females. Density approx. 5–7 strioles in breadth across an elytral interval.
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. Copelatus ankaratra seems to be a high-altitude alpine species associated with spring water.
Copelatus ankaratra sp. nov. belongs to the irinus group, based on the number of elytral striae. It belongs to the Copelatus insuetus species complex radiation on Madagascar and based on its CO1 it is most closely related to C. vokoka from Ivohibe. Notably, both these species have a densely striolated female form. There was a surprisingly large genetic distance (2.3–2.5% uncorrected p-distance) between the locality near the peak (2600 m) and the locality in the forested region ca. 5 km away at a lower altitude (2070 m). We find the male genitalia and other characters very similar and treat them here as conspecific.
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, Boophis williamsi (Guibé, 1974) and Mantidactylus pauliani Guibé, 1974 only found in montane streams at elevations above 2000 m.
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].
Mahajanga. Sofia: Bealanana: -HT♂ (GP) (
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 pseudostriatus is a compound word formed from pseudo- (false) and striatus (furrowed or striated) and refers to the intermediate non-complete striae in-between the complete continuous striae on the elytra in this species. It is the only species of Copelatus on Madagascar with this characteristic. The word striatus (masculine) is a participle (verb as adjective) in the nominative singular (ICZN 11.9.1.1).
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 Copelatus pseudostriatus sp. nov. is hardly possible: based on the complete striae it would fall in the irinus group, but the incomplete pseudostriae are likely reduced striae of an ancestor with a higher number of complete striae. This is a very distinct species with no recognisable close relatives, either based on genitalia or the CO1 gene. Tsaratanana Massif contains the highest peak in Madagascar at 2876 m and possibly this species is endemic to the Tsaratanana Massif. However,
Anjanaharibe Sud reserve, [14.7414S, 049.4975E] [Madagascar, Sava region, Andapa district]
Antsiranana. Sava: Andapa: -HT♂ (GP) (
Somewhat similar to C. insuetus on habitus appearance, but sturdier, broader pronotum and head with a much greater interocular distance compared to width of eyes, and subparalell along a longer distance of body with more rapidly attenuating anterior and posterior ends; easily distinguished from all other irinus group species of Madagascar by the penis shape, which is of a type otherwise found in species closely related to Copelatus owas; penis has a large medial expansion in lateral view followed by an apical blade (Fig.
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 C. insuetus. Compared with C. insuetus, the metacoxal lines are shorter, and the anterior traces suggest an inward curve towards the posterior metaventral margin. The lateral part of metaventrite is narrower than in C. insuetus at level of mesocoxa, equal to the width of mesofemur at middle. Posterior metaventral margin not straight but slightly angular at level of apex of mesotrochanter. Prosternal process is slightly more elongate.
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups. Pattern of suction cups same as for C. insuetus but tarsomeres not as wide and less developed as an integrated protarsal palett. Protibia modified, bisinuate, angled basally, and broadened distally. Pro- and mesotarsal claws unmodified. Penis very characteristic, rather broad and short and in lateral view with a medial expansion followed by a sharp constriction before the narrow blade-like apex (Fig.
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 |
---|---|---|---|---|---|---|
C. befasicus | F | 4 | 4.13 | 4.19 | 4.15 | 0.03 |
M | ||||||
C. marginipennis | F | 26 | 5.16 | 6.32 | 5.75 | 0.30 |
M | 45 | 5.48 | 6.58 | 6.02 | 0.26 | |
C. mahajanga | F | 12 | 5.16 | 6.13 | 5.63 | 0.24 |
M | 16 | 5.42 | 6.19 | 5.88 | 0.20 | |
C. pulchellus | F | 1 | 5.74 | 5.74 | 5.74 | |
M | 3 | 5.48 | 6.06 | 5.83 | 0.30 | |
C. distinguendus | F | 44 | 5.29 | 6.32 | 5.75 | 0.21 |
M | 26 | 5.48 | 6.26 | 5.92 | 0.18 | |
C. peridinus | F | 7 | 5.74 | 6.58 | 6.05 | 0.27 |
M | 7 | 5.81 | 6.26 | 6.10 | 0.16 | |
C. baculiformis | F | 1 | 4.00 | 4.00 | 4.00 | |
M | ||||||
C. kely sp. nov. | F | 7 | 3.81 | 4.26 | 4.01 | 0.16 |
M | 5 | 3.87 | 4.32 | 4.03 | 0.17 | |
C. insuetus | F | 27 | 4.26 | 5.03 | 4.60 | 0.20 |
M | 24 | 4.26 | 4.90 | 4.61 | 0.17 | |
C. safiotra sp. nov. | F | 3 | 4.32 | 4.71 | 4.58 | 0.22 |
M | 13 | 4.52 | 5.16 | 4.86 | 0.19 | |
C. vokoka sp. nov. | F | 6 | 3.94 | 4.32 | 4.18 | 0.13 |
M | 8 | 4.06 | 4.45 | 4.29 | 0.12 | |
C. ankaratra sp. nov. | F | 19 | 4.39 | 5.03 | 4.70 | 0.17 |
M | 30 | 4.65 | 5.16 | 4.87 | 0.16 | |
C. pseudostriatus sp. nov. | F | 2 | 5.42 | 5.61 | 5.52 | 0.14 |
M | 1 | 5.29 | 5.29 | 5.29 |
The species name safiotra is a Malagasy noun for hybrid, here referring to the unusual combination of a male genitalia type, typical of the Copelatus owas species complex, in a body with a 6+1 striated elytra very much resembling the C. insuetus complex of species. It is a non-latinised noun in apposition.
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.
Copelatus safiotra sp. nov. falls in the irinus group, based on the number of elytral striae (six discal and one submarginal striae). However, the genitalia is of a very different type and characteristic of the complex of species close to C. owas in the erichsonii group with ten discal and one submarginal striae. We hypothesize that this species, despite the body shape and number of elytral striae, is not related to the other species treated here, but belongs to the radiation of species around C. owas. This would reinforce the idea that the number of elytral striae is a very homoplastic character and not reliable to create phylogenetically sound groups (
We are aware of several Copelatus species not of the erichsonii group on Madagascar but of which we have only seen females. Some of these are undoubtedly new, confirmed with DNA data (Figs
[of C. nodieri & C. nodieri var. somalicus for comparison with sp1]: -LT♀ (
Mahajanga. Melaky: Antsalova: -1♀ (
This species is broad and oval in body shape and belongs to the pulchellus complex. Three species of the pulchellus complex are known from Madagascar, C. pulchellus, C. marginipennis, and C. mahajanga. It is most similar to C. marginipennis based on its very short and broad body shape. However, it differs in being almost entirely black dorsally, only slightly rufous laterally on the elytra, pronotum, and part of head. As such it is most similar in colouration to the dark form of C. pulchellus on Madagascar, but the body is shorter and broader. A second distinguishing characteristic is the very abbreviated first stria, present only in the posterior third, hence abbreviated even more than in C. mahajanga. Finally, this female specimen has fine but dense striolations mediolaterally on the elytra, and is densely punctured in the posterolateral corners of the pronotum (Fig.
It is plausible that it was one or several female specimens of this species that
Toamasina. Alaotra Mangoro: Moramanga: -1♀ (coll. Wewalka): // Data in
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 longicornis species group together with C. befasicus. However, the five striae are likely not homologous as C. befasicus has an abbreviated first stria, whereas the first stria is entirely lacking in this specimen, therby creating an interstriae space double in width compared to the outer intervals. But despite the lack of the first stria, it has five striae on the central and lateral parts of elytra where C. befasicus only has four. In addition, the striae are very faintly impressed, intermediate between real striae and the puncture lines found in C. peridinus. The colouration is uniformly brownish black like in C. peridinus and the body size is also similar. This specimen is possibly a different species compared to all others presented here; however, we cannot rule out that intraspecific variation of C. peridinus ranges from two defined puncture lines to five weakly impressed striae. They are very similar in all other aspects. We managed to sequence a partial fragment of CO1 (447 bp) from the Andasibe specimen (
Fianarantsoa. Ihorombe: Ivohibe: -3♀ (
The DNA data revealed that these females represent one or possibly two additional new species in the C. insuetus complex (Figs
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 Copelatus diversity on Madagascar represents four of the traditional species groups in the genus based on the number of elytral striae (
Copelatus as a genus is widespread all over Madagascar. Species can be found from the humid forest in the east to the dry forest in the west and from lowlands to the highest peaks. But the different species complexes have particular niches. Species in the C. insuetus complex are predominantely inhabitants of the eastern humid forests. Copelatus distinguendus and related species (C. peridinus, C. sp. 2, and likely C. baculiformis) mostly occupy open, often anthropogenically disturbed, habitats of the Central Highlands. The C. pulchellus complex (C. marginipennis, C. mahajanga, C. pulchellus, and C. sp. 1), and C. befasicus seem to be most abundant in the dry deciduous western forests, lowlands with open to semiopen landscapes. Finally, some species are specialists like the high altitude crenophile C. ankaratra sp. nov. and the sandy stream specialist C. safiotra sp. nov. Not until we have this knowledge – what are the species, how can we recognise them, where do they occur and how do they live – can we attempt to protect them and their habitats in the face of constantly increasing human pressure on the environment.
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 (
Examined specimens, species occurrence records
Data type: species data