Zwicknia ledoarei Reding, Launay, Ruffoni, Vinçon & Boumans, 2016

Reding, Jean-Paul G., Launay, Bertrand, Ruffoni, Alexandre, Vinçon, Gilles & Boumans, Louis, 2016, A new species of Zwicknia Murányi (Plecoptera, Capniidae) from the French and Swiss Jura Mountains, the French Massif Central, and the French Middle Rhône Region, Zootaxa 4121 (2), pp. 133-146 : 135-142

Zwicknia ledoarei Reding, Launay, Ruffoni, Vinçon & Boumans , sp. n.

( Figs. 1–6 )

Capnia bifrons ( Newman, 1838) pro parte

Capnia bifrons ( Newman, 1838) “ Caporb ” race sensu Rupprecht 1997 – Rupprecht 1997: 96 Fig. 4 (drumming signals).

Morphological diagnosis. Male micropterous ( Fig. 1 ); female macropterous. Vesicle small, rounded at apex ( Fig.

5). Process of tergite 9 low, not projecting caudally ( Figs. 2 and 6 ). Main epiproct sclerite strongly bent near mid-

length, banana-shaped in lateral view. Tip of the main epiproct sclerite ogive in lateral view. Main epiproct sclerite

long and slender, with nearly parallel edges and without bulges (dorsal view, Fig. 4 ). Female adults of Z. ledoarei

presently are not separable from other species of Zwicknia .

Type material. Holotype male: FRANCE: Jura Mountains, Ain Department, River Valserine, Mijoux, Golf

D50a bridge, 46° 22.777' N, 6° 01.055' E, 995 m a.s.l., 0 5.05.2014, leg. Bertrand Launay, deposited in the MZL

(catalogue number: GBIFCH00279494).

Paratypes: same locality, 3m, 1f, 0 5.05.2014, leg. B. Launay (paratypes deposited in the MZL under catalogue

number: GBIFCH00279777); Dénériaz coomb, temporary brooklet, 46° 51.206' N, 6° 31.708' E, 1135 m,

18.06.1995, 3m, leg. J.-P. G Reding ( MZL, catalogue number: GBIFCH00279201); Saut de l’Eau coomb,

temporary brooklet, 46° 50.487' N, 6° 30.974' E, 1232 m, 28.04.2015, 2m, leg. J.-P. G. Reding ( MZL, catalogue

number: GBIFCH00278545); Saut de l’Eau coomb, temporary brooklet, 46° 50.487' N, 6° 30.974' E, 1232 m,

3.04.2016, 6L (larvae), leg. J.-P. G. Reding ( MZL, catalogue number: GBIFCH00282526). Additional specimens are held in the collections of Jean-Paul G. and Alexis Reding (RC), Bertrand Launay

(BLC), Gilles Vinçon (GVC), Jacques Le Doaré ( JLDC), Alexandre Ruffoni ( ARC), NHMO and MZL.

1. SWITZERLAND, Jura Mountains, Chasseron region, Areuse River Basin, cantons of Neuchâtel and Vaud, Rhine tributaries:

1.1 Dénériaz coomb, temporary brooklet, 46° 51.206' N, 6° 31.708' E, 1135 m, 25.04.1993, 1m; 27.04.1993, 5m; 0 9.05.1993, 1m; 30.04.1994, 3m; 26.05.1995, 3m; 19.04.1996, 1m; 14.05.1997, 2m; 0 3.05.1998, 3m, 3f; 26.03.2003, 2m; 0 1.05.2003, 1m; 10.05.2008, 2f; 0 4.05.2011, 1L (leg. J.-P. G. Reding; RC)

1.2 Saut de l’Eau coomb, temporary brooklet, 46° 50.487' N, 6° 30.974' E, 1232 m, 31.05.1993, 2m; 12.05.2014, 1m, 1f (tissue collection NHMO, male used for molecular studies with lab number 2020); 28.04.2015, 1L (leg. J.-P. G. Reding; RC); 0 3.04.2016, 1m, 3L (leg. J.-P. G. Reding; RC)

1.3 Poëta Raisse, temporary brooklet, 46° 52.893' N, 6° 36.305' E, 1139 m, 29.03.1997, 1L; 26.04.1998, 1f (leg. J.- P. G. Reding; RC)

1.4 Breuil Brook, near Fleurier, 46° 54.189' N, 6° 36.982' E, 757 m, 0 5.04.1995, 1m, 2L; 0 9.04.1995, 2m (leg. J.-P. G. Reding; RC)

1.5 Spring of Fleurier Brook, Raisse, 46° 53.445' N, 6° 34.649' E, 800 m, 0 3.03.2012, 1m (leg. J.-P. G. Reding; RC)

1.6 Small tributary of Buttes River, near Buttes, 46° 53.243' N, 6° 33.349' E, 768 m, 21.03.1992, 1m; 13.03.1993, 2m, 2f; 21.03.1993, 3m; 23.03.1993, 1m; 0 6.04.1993, 1m, 1f (in copula); 15.04.2009, 1m (leg. J.-P. G. Reding; RC)

1.7 Noiraigue Brook, Noirvaux, 46° 50.856' N, 6° 30.452' E, 1003 m, 19.04.1996, 1m; 27.05.2008, 2f; 18.5.2013, 1m, 1L; 18.04.2015, 1m (leg. J.-P. G. Reding; RC)

1.8 Buttes River, Longeaigue, 46° 52.281' N, 6° 31.272' E, 812 m, 0 6.02.1996, 3L (leg. J.-P. G. Reding; RC)

1.9 Cambudes brook, waterfall, near Couvet, 46° 56.824' N, 6° 37.918' E, 1003 m, 13.04.1997, 1m; 0 7.06.1997, 1m (leg. J.-P. G. Reding; RC)

1.10 Cambudes brook, Plan du Pré, 46° 57.034' N, 6° 37.757' E, 1047 m, 13.04.1997, 1m; 0 1.05.2007, 1f (leg. J.-P. G. Reding; RC)

1.11 Cambudes brook, Trémalmont, 46° 57.321' N, 6° 38.039' E, 1090 m, 21.03.1998, 1L (leg. J.-P. G. Reding; RC)

1.12 Sucre brook, near Couvet, 46° 56.040' N, 6° 37.123' E, 870 m, 20.04.2005, 3L (leg. J.-P. G. Reding; RC)

2. SWITZERLAND, Jura Mountains, Mont d’Amin mountain, Seyon River Basin, Canton of Neuchâtel, Rhine tributaries:

2.1 Grande Berthière Brook, near La Chaux d’Amin, 47° 05.356' N, 6° 55.103' E, 1205 m, 0 6.11.2012, 5L; (leg. J.- P. G. Reding; RC)

3. SWITZERLAND / FRANCE, Jura Mountains, Jougnène headwaters, Orbe River Basin, Canton of Vaud, Switzerland and Doubs Department, France, Rhine tributaries:

3.1 Gascon Brook, near Grange Neuve, 46° 47.015' N, 6° 27.307' E, 1100 m, 27.04.2014, 14L; (leg. J.-P. G. Reding; RC)

4. SWITZERLAND, Jura Mountains, Vallée de Joux region, Orbe River Basin, Canton of Vaud, Rhine tributaries:

4.1 Spring of Orbe River, near Vallorbe, 46° 42.053' N, 6° 20.770' E, 775 m, 12.05.2015, 3m (leg. J.-P. G. Reding; RC)

4.2 Biblanc Brook, tributary of Orbe River, between Le Brassus and Bois-d’Amont, 46° 56.269' N, 6° 18.152' E, 1050 m, 24.04.1995, 11m, 2f; 30.05.1995, 2m, 3f; 27.06.1995, 2f (leg. G. Vinçon; GVC)

4.3 Biblanc Brook, La Bursine, 46° 33.790' N, 6° 10.898' E, 1067 m, 12.05.1978 – 25.07.1978, 18m, 17f (leg. J. Aubert; Aubert 1989: 262; MZL)

4.4 Epoisats Brook, near Vaulion, 46° 40.930' N, 6° 20.457' E, 1034 m, 22.04.1978, 15m, 28f (leg. J. Aubert; Aubert 1989: 262; MZL)

4.5 Spring of the Nozon River, near Vaulion, 46° 40.643' N, 6° 22.925' E, 980 m, 26.05.1979, 2m, 2f (leg. J. Aubert; Aubert 1989: 262; MZL); 12.05.2015, 1f (leg. J.-P. G. Reding; MZL, catalogue number: GBIFCH00282543)

4.6 Nozon River, Vaulion, 46° 41.423' N, 6° 23.724' E, 920 m, 13.04.2010, 1L (leg. SESA, MZL, catalogue number: GBIFCH00280271)

5. FRANCE, Jura Mountains, Doubs Department, Doubs drainage basin, Rhône tributaries:

5.1 Spring of Doubs River, Mouthe (25), 46° 42.295' N, 6° 12.545' E, 945 m, 29.04.2011, 4f (leg. J. Le Doaré; JLDC)

5.2 Small tributary of Lhaut River, bridge over Lhaut River, near Labergement-Sainte-Marie (25), 46° 45.503' N, 6° 15.768' E, 860 m, 12.05.1998, 1m; (leg. J.-P. G. Reding; RC)

5.3 Spring at Les Vurpillières, National Nature Reserve of the Lake of Remoray (25), 46° 45.485' N, 6° 15.283' E, 852 m, 15.05.1993, 1L; 0 5.06.1993, 1f; (leg. J.-P. G. Reding; RC)

5.4 Spring of Les Capucins, tributary of Lhaut River, near Brey-et-Maison du Bois, 46° 45.418' N, 6° 15.73' E, 870m, 11.04.2009, 2m (leg. J. Le Doaré; JLDC)

5.5 Small tributary of Drugeon River, Levier coomb, near Bonnevaux, 46° 49.338' N, 6° 14.073' E, 867 m, 10.06.2008, 1f; (leg. J.-P. G. Reding; RC); 11.04.2009, 7L (leg. J. Le Doaré; JLDC)

6. FRANCE, Jura Mountains, Jura dpt, Ain drainage basin, Rhône tributaries:

6.1 Spring of Ain River, near Conte, 46° 44.991' N, 6° 01.386' E, 708 m, 13.04.1991, 27m, 3f; 19.07.1991, 2f (leg. J. Aubert; GVC); 10.04.2009, 1m, 8f (leg. J. Le Doaré; JLDC); 0 2.04.2010, 5m, 2f, 12L; 24.05.2010, 1m, 15f (leg. A. Reding; females RC, male NHMO and used for molecular studies with lab number 2019); 0 2.05.2015, 15m, 5f (leg. B. Launay; BLC)

6.2 Spring of Saine River, near Foncine-le-Haut, 46° 40.108' N, 6° 04.678' E, 900 m, 25.04.2008, 1m, 2f (leg. M. Genoud, JLDC); 0 2.04.2010, 4L; 24.05.2010, 1f, 1L; 20.07.2010, 1f; 0 1.03.2012, 1f, 5L; 30.04.2014, 3m, 2f (leg. J.-P. G. Reding; RC)

6.3 Saine River, Chez Vallet near Foncine-le-haut, 46° 39.527' N, 6° 04.267' E, 877 m, 12.06.2008, 1m, 1f (leg. J. Le Doaré, JLDC)

7. FRANCE, Jura Mountains, Ain dpt, Valserine and Albarine drainage basins, Rhône tributaries:

7.1 Tributary of Taillis brook, forest of Côte Aubert, Cormaranche en Bugey, 45° 58.025' N, 5° 37.477' E, 1000m, 31.03.2013, 7m, 16L; 14.04.2013, 1m; 26.05.2013, 1m, 1f; 25.01.2014, 1m, 4L (leg. B. Launay; BLC)

7.2 Mélogne brook, upstream D8 bridge, Hauteville-Lompnes (01), 45° 58.072' N, 5° 36.523' E, 868m, 0 7.04.2013, 6m, 1L (leg. B. Launay; BLC)

7.3 Valserine, spring, Lajoux (01), 46° 25.027' N, 6° 03.507' E, 1158m, 0 6.04.2014, 1L (leg. B. Launay; BLC)

7.4 Valserine, La Villette bridge, Mijoux (01), 46° 23.813' N, 6° 02.37' E, 1050 m, 26.04.2015, 1m (leg. B. Launay; BLC)

7.5 Valserine, D50a bridge (golf), Mijoux (01), 46° 22.777' N, 6° 01.055' E, 995 m, 0 7.04.2014, 4m, 1f; 0 5.05.2014, 9m, 1f; 31.05.2014, 3m, 2f; 11.04.2015, 8m, 9L; 26.04.2015, 5m, 1f (leg. B. Launay; BLC & RC)

7.6 Valserine, under Mijoux (01), 46° 21.753' N, 5° 59.008' E, 970 m, 13.04.1991, 1m (leg J. Aubert, GVC) 7.7 Valserine, la Rosselle, under Mijoux (01), 46° 20.998' N, 5° 58.498' E, 940 m, 26.04.2015, 1m, 1f (leg. B. Launay; BLC)

7.8 Séran River, D9 bridge, Ruffieu (01), 45° 59.863' N, 5° 40.662' E, 664m, 14.04.2013, 12m, 3f, 5L (leg. B. Launay; BLC & RC)

7.9 Bief du Ravinet, upstream Sous Panafay, Torcieu (01), 45° 54.672' N, 5° 24.962' E, 412m, 16.03.2013, 1m; 25.01.2014, 1m; 22.03.2015, 4m (leg. B. Launay; BLC)

8. FRANCE: Middle Rhône Region, Rhône tributaries:

8.1 Roubion River, near Pont de Barret (26), 44° 36.5' N, 5° 00.655' E, 250m, 27.01.2014, 4m; 0 5.03.2015, 5m, 1f (leg. B. Launay; BLC & RC)

8.2 Lez River, bridge on road D130, Teyssières (26), 44° 28.222' N, 05° 08.183' E, 545m, 0 4.03.2015, 2m, 1f (leg. B. Launay; BLC)

8.3 Lez River, ford opposite to La Borie, Roche-Saint-Secret-Béconne Region (26), 44° 29.9' N, 05° 03.648' E, 390m, 0 4.03.2015, 6m, 3f (leg. B. Launay; BLC)

8.4 Méouge River, near Les Iscles, côte 817m, Séderon (26), 44° 11.788' N, 5° 32.35' E, 817m, 0 3.03.2015, 1m, 1f (leg. B. Launay; BLC)

9. FRANCE: Massif Central, Rhône tributaries:

9.1 Tributary to Ay River, Sarras Town Center (07), 45° 10.918' N, 4° 47.172' E, 200 m, 23.02.2007, 1m (leg. J. Le Doaré; JLDC)

9.2 Ribeyrette River, near Chamborigaud (30), 44° 18.138' N, 3° 58.623' E, 295 m, 12.02.2015, 2m (leg. B. Launay, BLC)

9.3 Valencize River, bridge on road D34, côte 339m, Pélussin (42), 45° 25.477' N, 4° 41.29' E, 330m, 14.03.2015, 26m, 2f, 2L (leg. B. Launay; BLC)

10. FRANCE: Massif Central, Loire tributaries:

10.1 Boisserand temporary brook, Arroux tributary, near Auxy (71), 46° 57.473' N, 4° 28.002' E, 415 m, 18.02.2011, 3m (leg. A. Ruffoni; ARC)

10.2 Gazeille River, between Les Estables and Le Monastier-sur-Gazeille (43), 44° 55.408' N, 4° 02.833' E, 940 m, 15.04.2004, 1m, 2f (leg. G. Vinçon; GVC)

[11. SWITZERLAND: northeastern Prealps, canton of Thurgau, Thur drainage basin, Rhine tributaries: 11.1 Tobelbach, Gschmelltobel, 47° 37.341' N, 9° 1.762' E, 490 m, 20.04.2006, 1m (leg. U. Mürle, MZL, catalogue number: GBIFCH 00279956)]

Description. Head, thorax, appendages and basal segments of the abdomen are typical for the genus. Males are micropterous ( Fig. 1 ), females macropterous. Body length of males 4.5 to 7.6 mm, females 5.4 to 10.1 mm. Forewing length of males 0.7 to 1.2 mm.

Male terminalia: process of tergite 9 low, rising sharply near the distal region of the corresponding tergite ( Figs. 1 , 2 , 3 and 6 ), not projecting caudally ( Figs. 2 and 6 ). In caudal view, the apical part of the process of tergite 9 has neatly marked angles at the upper corners ( Fig. 3 ). The process itself is narrow, only about twice as wide as the main epiproct sclerite, in dorsal view ( Fig. 4 ). Vesicle small, rounded at apex ( Fig. 5 ), width about ¼ subgenital plate, sometimes 1/5 width in specimens from higher locations (> 1100 m). The heart-shaped subgenital plate is densely covered with long setae ( Figs. 2 , 3 and 5 ). In lateral view, the main epiproct sclerite (Ep-scl, sensu Murányi et al. 2014) is strongly bent near mid-length, banana-shaped; its tip is ogive, neither up-curved nor down-curved ( Figs. 2 , 3 and 6 ). In dorsal view, the main epiproct sclerite is long and slender, with nearly parallel edges and without bulges ( Fig. 4 ). The tips of the pairwise arranged upper sclerites on the main epiproct sclerite are long and pointed.

Female adults of Z. ledoarei presently not separable from other species of Zwicknia .

Morphological affinities. Zwicknia ledoarei is morphologically most similar to Z. rupprechti but differing by the following: In lateral view, the main epiproct sclerite of Z. rupprechti is pointed with the tip slightly up-curved (Figs. 92 and 107, Murányi et al. 2014), whereas Z. ledoarei has a tip that is blunt and ogive in shape ( Figs. 2 , 3 and 6 ). In dorsal view, the main epiproct sclerite of Z. rupprechti bulges out slightly medially (Fig. 90, Murányi et al. 2014), whereas the shape of the main epiproct sclerite is uniformly parallel and is also slightly longer in Z. ledoarei ( Fig. 4 ). The tips of the pairwise arranged upper sclerites of the main epiproct sclerite of Z. rupprechti (Fig. 90, Murányi et al. 2014) are much shorter than those of Z. ledoarei ( Fig. 4 ). Moreover, the process of tergite 9 is projecting caudally in Z. rupprechti (Fig. 92, Murányi et al. 2014), instead of upwards in Z. ledoarei ( Figs. 2 , 3 and 6 ). In caudal view, the process of tergite 9 of Z. rupprechti (Fig. 93, Murányi et al. 2014) is much broader than that of Z. ledoarei ( Fig. 4 ).

Zwicknia ledoarei differs more markedly from Z. bifrons . This latter species has a much larger ventral vesicle (Fig. 83, Murányi et al. 2014) than Z. ledoarei ( Fig. 5 ). Additionally, this vesicle is mussel shaped in Z. bifrons , whereas it is rounded in Z. ledoarei ( Fig. 5 ). The main epiproct sclerite of Z. ledoarei is strongly bent, bananashaped, in lateral view ( Figs. 2 , 3 and 6 ) and only slightly bent in Z. bifrons . In lateral view, the main epiproct sclerite of Z. bifrons is pointed (Fig. 84, Murányi et al. 2014), whereas Z. ledoarei has a tip that is blunt and ogive in shape ( Figs. 2 , 3 and 6 ). Process of tergite 9 in Z. bifrons is elevated and perpendicular (Fig. 84, Murányi et al. 2014), whereas it is low in Z. ledoarei ( Figs. 1 , 2 , 3 and 6 ). The process of tergite 9 rises gradually from the proximal to the distal part of the corresponding tergite in Z. bifrons (Fig. 84, Murányi et al. 2014), whereas it rises sharply only in the distal region of the corresponding tergite in Z. ledoarei ( Figs. 2 and 6 ).

Zwicknia westermanni , the fourth species of Zwicknia known from the Jura Mountains and the Massif Central, has no close morphological affinities with Z. ledoarei . The main epiproct sclerite is much longer and slender in lateral view ( Fig. 3 , Boumans & Murányi 2014). The male is brachypterous with a large ventral vesicle ( Fig. 4 , Boumans & Murányi 2014). In contrast, the male of Z. ledoarei has a smaller and thicker main epiproct sclerite, banana-shaped in lateral view ( Figs. 2 , 3 and 6 ), and is micropterous ( Fig. 1 ) with a very small and rounded vesicle ( Fig. 5 ).

Molecular identification. Both COI and 28S support the new species as a distinct lineage within Zwicknia . While many relationships are unresolved in the 656 bp COI-based mitochondrial phylogeny, the three specimens morphologically identified as Z. ledoarei clearly stand out as a well-supported haploclade ( Fig. 7 ). The two COI sequences from the Swiss and French Jura Mountains are identical and 0.6% different from the SwissBOL sequence from Thurgau. The haplotypes most similar to Z. ledoarei belong to French and German Z. rupprechti (4.3%–5.5% uncorrected difference) and Andalusian Z. bifrons (5.0%–5.2% difference).

The 28S alignment of 758 bp contains only a small number of nucleotide substitutions and is therefore represented as a network ( Fig. 8 ). Within the available data set, the 28S allele of Z. ledoarei is species-specific. The least similar alleles belong to Z. rupprechti from France and Germany. The latter are further distinguished from all other Zwicknia by an indel of 2 base pairs.

Identifications based on drumming signals. Zwicknia ledoarei was distinguished as early as 1993 by Rupprecht (1997). He recognised the temporal organisation of the duetting behaviour as unique and reported it as “ Caporb ” of the C. bifrons species complex. Rupprecht analysed the drumming behaviour of populations from four locations in the Swiss Jura Mountains, the karstic spring of the Orbe River at Vallorbe (Canton of Vaud, Switzerland) as well as three sampling sites in the headwaters of the Areuse River Basin (cantons of Neuchâtel and Vaud, Switzerland, stations 1.1, 1.4 and 1.7, see above). He found that while the signals of the male-female drumming duets of Z. ledoarei are similar to those of Z. rupprechti , the interval between the call of the male and the answer of the female of Z. ledoarei is much longer (about one second, Rupprecht 1997, Fig. 4 ) than the one of Z. rupprechti (only about half a second, see also Murányi et al. 2014, Figs. 188 and 189). Moreover, the behaviour of the male of Z. ledoarei , who runs after producing a signal and stops shortly before the female response is to be expected, is also different from that of Z. rupprechti ( Rupprecht 1997) .

Distribution ( Fig. 9 ). In France, outside of the Jura Mountains, Z. ledoarei occurs in the eastern foothills of the Massif Central, but does not seem to extend to the Langres Plateau and the headwaters of the Seine River. Its distribution extends from the foothills of the Massif Central into the French Middle Rhône Region on both sides of the river, to the French Prealps. The species is then found further in the Bugey Region, mainly in tributaries of the Rhône (Séran, Valserine and Ain drainage basins). The occurrence of Z. ledoarei in the Jura Mountains is patchy, despite being the major center of distribution area for the species. Zwicknia ledoarei occurs in many karstic springs of the French portion of the Jura Mountains, within an area that includes the springs of the Doubs, the Drugeon, the Saine, the Ain, the Valserine, and the Séran. In Switzerland, Z. ledoarei occupies the western parts of the Jura Mountains, where it is very abundant in the Areuse drainage basin and the Vallée de Joux ( Aubert 1989; Reding 1998), hydrologically linked via the Orbe River, whose headwaters are in dispersal range from Valserine Spring. The occurrence of Z. ledoarei is less frequent in the eastern portions of the Swiss Jura Mountains and it seems to be absent from the Birs drainage basin (Tabular Jura, Küry 1994). In France, Z. ledoarei is not known from the Loue and Lison drainage basins.

Within the above regions, three additional species of Zwicknia occur. Zwicknia bifrons and Z. westermanni are to be found only at lower altitudes at the foothills of the Jura Mountains; the distribution area of Z. rupprechti does not extend beyond the Massif Central.

Ecology. Zwicknia ledoarei can be collected from small perennial as well as temporary brooks and springs up to 1250 m in the High Jura Chain of France and Switzerland, and karstic springs at lower altitudes in the Bugey Region. The species seems absent, however from springs that are isolated and not connected to larger watercourses. In the Massif Central and the French Prealps, the species rarely occurs in headwaters and springs, but more often in larger streams and rivers with gravel substrates (for example the Roubion River).

No other species of the genera Zwicknia or Capnia Pictet are to be found in sympatry with Z. ledoarei in the High Jura Chain. In the Massif Central, however, Z. ledoarei occurs in at least one sampling-station (intermittent Boisserand Brook, station 10.1) in sympatry with Z. westermanni and Z. rupprechti .

The flight period of Z. ledoarei extends from early spring to early summer. Adults of both sexes emerge from the end of February until mid-May; females may be encountered up to the end of July, especially in karstic springs. In the High Jura Chain, oviposition often takes place in small residual water-bodies connected to hyporheic zones in otherwise dry riverbeds.

Etymology of Zwicknia ledoarei . This species is dedicated to Jacques Le Doaré, Châteaulin, France, in recognition of his important contribution to our knowledge of the distribution and ecology of the Plecoptera in France.

Figures

FIGURE 1. Zwicknia ledoarei sp. n., micropterous male, Bief du Ravinet brook, Torcey, Ain dpt, France. Photo B. Launay.

FIGURE 2. Zwicknia ledoarei sp. n., Valserine River, Mijoux (01), terminalia, lateral view. Photo B. Launay.

FIGURE 3. Zwicknia ledoarei sp. n., Valserine River, Mijoux (01), terminalia, latero-caudal view. Photo B. Launay.

FIGURE 4. Zwicknia ledoarei sp. n., Séran River, Ruffieu (01), terminalia, dorsal view. Photo B. Launay.

FIGURE 5. Zwicknia ledoarei sp. n., Séran River, Ruffieu (01), terminalia, ventral view. Photo B. Launay.

FIGURE 6. Zwicknia ledoarei sp. n., Dénériaz coomb, canton of Vaud, Switzerland, terminalia with eversible crest of epiproct everted, latero-dorsal view. Photo J. - P. G. Reding.

FIGURE 7. Phylogenetic tree of Zwicknia mitochondrial haplotypes based on 656 bp of COI. Bayesian inference tree. Support values: *, ** indicate Bayesian posterior probability> 0.95 and> 0.99 respectively; MP and NJ bootstrap percentages are shown in this order separated by a slash. Bootstrap values <50 not shown.

FIGURE 8. Median-joining network of 758 bp partial 28 S sequences of Zwicknia species.

FIGURE 9. Distribution map of Zwicknia ledoarei sp. n. Compiled by A. Ruffoni.