Laccophilus bergsteni, Manuel & Ramahandrison, 2020

Manuel, Michaël & Ramahandrison, Andriamirado T., 2020, Four new species of the diving beetle genus Laccophilus Leach, 1815 from Madagascar (Coleoptera, Dytiscidae, Laccophilini), Zootaxa 4822 (4), pp. 482-502 : 486-491

Laccophilus bergsteni sp. nov.

Figures 2 , 6 , 10, 17 , 25, 26

Type locality. Madagascar, former province of Antananarivo, Ankaratra massif, ca. 9.5 km NW Manjakatompo, GPS coordinates S19°20’ E47°15’, altitude 2,400 m .

Type material. Holotype (♂): ” Madagascar. Ex-prov. / Antananarivo. ca. 9.5 km NW / Manjakatompo . 22 XII 2017 / Ramahandrison & Manuel leg. / S19°20’ E47°15’ // Alt. 2,400m. Slowly flowing / stream on flat area in subalpine / grassland. Ankaratra / mountain massif” [pr], ” Holotype, Laccophilus bergsteni sp. nov., Manuel & Ramahandrison 2020” [red, pr] ( MNHN) . Paratypes (in total 168♂♂, 154♀♀) ( BMNH, CMM, DEUA, MNHN, NHRS, NMPC, PBZT / MBC) : 1♂♂, 1♀♀, same data as holotype . 2♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 6 km / WNW Manjakatompo . / 21 XII 2017. Ramahandrison / & Manuel leg. // S19°21’ E47°17’ / Alt. 2,050m. Muddy swamp / in natural forest (Tavolotara) / fed by small stream. / Ankaratra mountain massif” . 3♂♂, 3♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 6 km WNW / Manjakatompo . 21 XII 2017 / Ramahandrison & Manuel leg. / S19°21’ E47°17’ // Alt. 2,000m. Slowly flowing / stream, muddy bottom with / tree leaves and flowering plants. / Natural forest (Tavolotara), / Ankaratra mountain massif” . 41♂♂, 44♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 3 km WNW / Manjakatompo. 14 VII 2018 / Ramahandrison leg. // S19°21’ E47°18’ / Alt. 1,800m. Pond in degraded / forest (pine trees, mimosa) / ( MANS3 ). Ankaratra mountain / massif” . 2♂♂, 3♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 6 km WNW / Manjakatompo . 21 VII 2018 / Ramahandrison leg. // S19°21’ E47°17’ / Alt. 2,000m. Shallow pond / in natural forest (Tavolotara) / ( MANS6 ). Ankaratra / mountain massif” . 1♂♂, 2♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 6 km WNW / Manjakatompo . 19 II 2019 / Ramahandrison leg. / S19°21’ E47°17’ // Alt. 2,000m. Shallow pond / in natural forest (Tavolotara) / ( MANS5 ). Ankaratra / mountain massif” . 1♂♂, 6♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 6 km WNW / Manjakatompo . 20 II 2019 / Ramahandrison leg. // S19°21’ E47°17’ / Alt. 2,000m. Swamp in / natural forest (Tavolotara) / ( MANS8 ). Ankaratra / mountain massif” . 2♂♂, 3♀♀, “ Madagascar. Ex-prov. / Antananarivo. ca. 3 km W / Manjakatompo. 22 II 2019 / Ramahandrison leg. // S19°22’ E47°18’ / Alt. 1,800m. / Pond in forest ( MANS13 ). / Ankaratra mountain massif” . 7 exs., “ Madagascar: Antananarivo: / Vakinankaratra: Manjakatompo Stn. / Forestière : 500m E Lac Froid by the / road: S 19.34485 E 47.33381, 1620 m. / 04:XI:2011: GB Nets and sieves: pond / and inlet stream: Field# MAD11-16 // Leg. J. Bergsten, R. / Bukontaite, T. / Ranarilalatiana & / J.H. Randriamihaja // NHRS-JLKB / 000010394” . 2 ♂♂, 2 ♀♀, “ Madagascar: Antananarivo: Vakinan- / karatra: Manjakatompo Stn. Forestière / river Analamitana by bridge close to / SKOL facility: S 19.36458 E 47.29885, / 1750 m. 05:XI:2011: GB Nets & sieves: / stream & stagn. pool: Field# MAD11-19 // Leg. J. Bergsten, R. / Bukontaite, T. / Ranarilalatiana & / J.H. Randriamihaja // NHRS-JLKB / 000010453” . 4 ♂♂, 4 ♀♀, “ Madagascar: Antananarivo: / Vakinankaratra: Manjakatompo Stn. / Forestière :Analamitana: S 19.363972 E / 47.299083, 1757m. 22:I:2012: swamp / near stream: Field# MJK12-02: Leg. T. / Ranarilalatiana & J. H. Randriamihaja // NHRS-JLKB / 000065530” . 6 ♂♂, 7 ♀♀, “ Madagascar: Antananarivo: / Vakinankaratra: Manjakatompo Stn. / Forestière : Manontongona : S 19.3360 E / 47.295167, 1847m. 23:I:2012: swamp / near stream: Field# MJK12-04: Leg. T. / Ranarilalatiana & J. H. Randriamihaja // NHRS-JLKB / 000065531” . 14 ♂♂, 6 ♀♀, “ Madagascar: Antananarivo: / Vakinankaratra: Manjakatompo Stn. / Forestière : Tavolatara : S 19.349139 E / 47.278361, 2046m. 24:I:2012: swamp / near source: Field# MJK12-08: Leg. T. / Ranarilalatiana & J. H. Randriamihaja // NHRS-JLKB / 000065532” . 5 ♂♂, 4 ♀♀, “ Madagascar: Antananarivo: / Vakinankaratra: Manjakatompo Stn. / Forestière : Antandraolatra : S 19.35425 E / 47.294861, 1808m. 24:I:2012: w. hole / near river: Field# MJK12- 11: Leg. T. / Ranarilalatiana & J. H. Randriamihaja // NHRS-JLKB / 000065533” . 7 ♂♂, 5 ♀♀, “MAD: ANTA: Analamanga: Ambo / hitantely special reserve: / Mad 14-76: forest stream / with waterfalls and over bedrock: / 18.1808S 47.2901E: / 1340m: 22.XI.2014 // Leg. J. Bergsten, / R. Bukontaite, S. / Holmgren, / J. H. Randriamihaja / & T. Ranarilalatiana // NHRS-JLKB / 000026505” . 17 ♂♂, 12 ♀♀, “MAD: ANTA: Analamanga: Anjoz / orobe forest reserve: stream / next to Saha forest camp, 10km / E of Anjozorobe: Mad 14- / 77: small forest stream: 1240m: / 18.4133S 47.9443E: 23.XI.2014 // Leg. J. Bergsten, / R. Bukontaite, S. / Holmgren, / J. H. Randriamihaja / & T. Ranarilalatiana // NHRS-JLKB / 000026501” . 13 ♂♂, 9 ♀♀, “MAD: ANTA: Analamanga: Anjoz / orobe forest reserve: marsh next / to the stream by Saha forest , / 10km E of Anjozorobe: Mad 14- / 78: forestmarsh: 18.4128S / 47.9439E: 1320m: 23.XI.2014 // Leg. J. Bergsten, / R. Bukontaite, S. / Holmgren, / J. H. Randriamihaja / & T. Ranarilalatiana // NHRS-JLKB / 000026499” . 3 ♂♂, 1 ♀, “ Madagascar – Vakinankaratra / Ambatolampy: MAD16-01 / Andalantsoavaly ruisseau / S19° 37588: E047° 35180; / Alt. 1599m; 03/02/2016 / Leg. T. Ranarilalatiana // NHRS-JLKB / 000065574” . 2 ♂♂, 2 ♀♀, “ Madagascar – Vakinankaratra / Ambatolampy: MAD16-02 / Manjakatompo Reserve / Ankaratra ruisseau / S19° 34409: E047° 32322; / Alt. 1796m; 03/02/2016 / Leg. T. Ranarilalatiana // NHRS-JLKB / 000065575” . 20 ♂♂, 9 ♀♀, “ Madagascar – Vakinankaratra / Ambatolampy: MAD16-05 / Manjakatompo Reserve / Source ruisseau Tavolotara / S19° 34917: E047° 27736; / Alt. 2044m; 04/02/2016 / Leg. T. Ranarilalatiana // NHRS-JLKB / 000065578” . 9 ♂♂, 10 ♀♀, “ Madagascar – Vakinankaratra / Ambatolampy: MAD16-06 / Manjakatompo Reserve / Analamitana ruisseau / with side ponds / S19° 36397: E047° 29907; / Alt. 1759m; 05/02/2016 / Leg. T. Ranarilalatiana // NHRS-JLKB / 000065579” . 1 ♂, “ Madagascar – Analamanga / Ankazobe: MAD16-19 / Ambohitantely Special Reserve / forest stream / S18° 18411: E047° 30251 / Alt. 1533m: 09/03/16 / T. Ranarilalatiana // NHRS-JLKB / 000065595” . 3 ♂♂, 1 ♀, “ Madagascar – Analamanga / Ankazobe: MAD16-22 / Ambohitantely Special Reserve / Zone 2, ruisseau 8 Ambohitraingy / S18° 16321: E047° 28335 / Alt. 1553m: 11/03/16 / T. Ranarilalatiana // NHRS-JLKB / 000065580” . 5 ♂♂, 3 ♀♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-32 / Forêt d’Anjozorobe Angavo / Ruisseau Antavinamboa Ambosarinala / S18° 45036: E047° 94942 /Alt. 1306m: 02/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000065581” . 4 ♀♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-33 / Forêt d’Anjozorobe Angavo / Ruisseau Ambatonisana Ambosarinala / S18° 45113: E047° 94677 / Alt. 1298m: 02/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000065582” . 1 ♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-35 / Forêt d’Anjozorobe Angavo / Ruisseau Sahalemaka Ambosarinala / S18° 44919: E047° 95928 / Alt. 1367m: 03/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000070601” . 1 ♂, 2 ♀♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-36 / Ruisseau Ambatovikinina / Ambosarinala , Hors réserve / S18° 45792: E047° 93438 / Alt. 1367m: 04/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000070602” . 1 ♂, 1 ♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-37 / Forêt d’Anjozorobe Angavo / Ruisseau Ankorona Ambosarinala / S18° 42555: E047° 95121 / Alt. 1331m: 04/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000070605” . 2 ♀♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-38 / Forêt d’Anjozorobe Angavo / Ruisseau Andrianarivo Ambosarinala / S18° 42407: E047° 95590 / Alt. 1356m: 04/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000070607” . 1 ♀, “ Madagascar – Analamanga / Anjozorobe: MAD16-39 / Forêt d’Anjozorobe Angavo / South of Angavo / S18° 42192: E047° 93801 / Alt. 1308m: 05/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000065601” . 1 ♂, “ Madagascar – Analamanga / Anjozorobe: MAD16-40 / Forêt d’Anjozorobe Angavo / Ruisseau Antanketsandramanitra / circuit bambou Antsahabe / S18° 41266: E047° 94380 / Alt. 1341m: 05/04/16 / T. Ranarilalatiana // NHRS-JLKB / 000065602”. All paratypes with the respective printed red label .

Description. See under Laccophilus leguyaderi sp. nov. Laccophilus bergsteni sp. nov. is morphologically closely similar to L. leguyaderi sp. nov., therefore we provide only a differential diagnosis of the species.

Differential diagnosis. This species differs from L. leguyaderi sp. nov. by the following characters:

- body length on average slightly smaller;

- pale area between anterior elytral margin and anterior extremity of dark longitudinal markings on average a bit narrower and slightly paler (yellow-whitish, vs. more frankly yellow in L. leguyaderi ; compare Figs. 1 and 2 );

- metacoxal plates much darker, laterally dark brown to black, medially gradually transitioning to ferrugineous (compare Fig. 5 and 6 );

- ventral surface of prothorax also on average slightly darker;

- mesotibia much narrower, unmodified (compare Fig. 5 and 6 ) and identical in both sexes (in shape and microsculpture); along dorsal edge without row of long setae; anterior surface as in L. leguyaderi females;

- protibia also distinctly narrower;

- pro- and mesotarsomeres I–III much less broadened, with ventral adhesive setae smaller;

- protarsal and mesotarsal claws a bit shorter;

- median lobe of aedeagus in lateral view ( Fig. 10 ) as in L. leguyaderi ; in ventral view a bit narrower ( Fig. 17 ). Parameres ( Figs. 25, 26 ) as in L. leguyaderi .

Variability. Specimens vary slightly in habitus (length / width ratio) and dorsal colour pattern (intensity of the elytral brown longitudinal markings; degree of fragmentation of the pale interspaces between these dark bands; extension of the pale area between anterior margin of elytra and anterior extremity of dark longitudinal markings). Some specimens have the metacoxal plates largely dark brown to black as in the holotype ( Fig. 6 ) whereas in other specimens the ferrugineous medial area is more extended.

Measurements. Holotype: TL = 4.0 mm, TL without head = 3.7 mm, MW = 2.2 mm, ratio TL/MW = 1.81. Paratypes: TL = 3.65–4.3 mm (4.03±0.15), TL without head = 3.5–4.1 mm (3.81±0.13), MW = 2.0– 2.4 mm (2.24±0.08), TL/MW = 1.75–1.86 (1.80±0.02).

Habitat. This species was collected at an altitude range of 1,240 –2,430 m. Its ecology is similar to that described above for L. leguyaderi . Notably, in the Ankaratra massif, it is present in natural or degraded mountain forests ( Fig. 45 ), as well as above tree line in subalpine grasslands ( Fig. 46 ).

Distribution. So far known from the Ankaratra, Ambohitantely and Anjozorobe-Angavo mountain ranges, in central Madagascar ( Fig. 53 ).

Derivatio nominis. This species is dedicated to Dr. Johannes Bergsten (Department of Zoology, Swedish Museum of Natural History, Stockholm). The species name is a noun in the genitive singular.

Figures

FIGURE 53. Map showing the distribution of known localities of the newly described species. Red dot: L. leguyaderi; Blue dots: L. bergsteni; Magenta dot: L. bergsteni and L. rakouthae; yellow dot: L. makay. Names of corresponding protected areas or massifs indicated in italics above dots.

FIGURES 1–4. Habitus in dorsal view of: (1) Laccophilus leguyaderi sp. nov. (holotype); (2) Laccophilus bergsteni sp. nov. (holotype); (3) Laccophilus makay sp. nov. (holotype); (4) Laccophilus alluaudi (male specimen from Isalo massif). Scale bar: 1 mm.

FIGURES 5–8. Ventral surface of: (5) Laccophilus leguyaderi sp. nov. (holotype); (6) Laccophilus bergsteni sp. nov. (holotype); (7) Laccophilus makay sp. nov. (holotype); (8) Laccophilus alluaudi (male specimen from Isalo massif). Scale bar: 1 mm.

FIGURES 9–22. Median lobe of aedeagus in lateral (9–15) and ventral (16–22) view of: (9, 16) Laccophilus leguyaderi sp. nov. (holotype); (10, 17) Laccophilus bergsteni sp. nov. (holotype); (11, 18) Laccophilus makay sp. nov. (holotype); Laccophilus alluaudi, male specimens from (12, 19) Montagne d’Ambre, (13, 14, 20, 21) Isalo massif, and (15, 22) Ibity massif. Scale bar: 0.1 mm.

FIGURES 23–30. Left (23, 25, 27, 29) and right (24, 26, 28, 30) parameres of: (23, 24) Laccophilus leguyaderi sp. nov. (holotype); (25, 26) Laccophilus bergsteni sp. nov. (holotype); (27, 28) Laccophilus makay sp. nov. (holotype); (29, 30) Laccophilus alluaudi (male specimen from Isalo massif). Scale bar: 0.1 mm.

FIGURES 43–46. Habitats of Laccophilus leguyaderi sp. nov. (43–44) and Laccophilus bergsteni sp. nov. (45–46). (43) Shallow brooklet with peaty bottom in subalpine grassland (Andringitra massif, 2,000 m); (44) Rather deep brook with peaty bottom, slowly flowing among subalpine shrubs (Andringitra massif, 1,970 m; type locality of L. leguyaderi); (45) Slowly flowing stream in natural forest with muddy bottom and lots of tree leaves (Ankaratra massif, Tavolotara, 2,000 m); (46) Slowly flowing stream on flat area in subalpine grassland (Ankaratra massif, 2,400 m; type locality of L. bergsteni).