Adults can be recognised within the Colymbetinae on the following combination of characters: pronotal beading absent; protibiae only weakly emarginate basoventrally; prosternal process medially rounded; anterior margin of metaventrite deeply incised for reception of prosternal process; metatarsomeres I-IV distinctly sinuate apically, with apicolateral lobes and metatarsal claws subequal in length, outer approximately two-thirds length of inner (Miller and Bergsten 2016). The sculpture of the elytra referred to by Miller and Bergsten (2016) is not a constant generic character of Meladema (see below). Larvae of two species (M. coriacea and M. lanio) have been described to date (Bertrand 1928, 1932a, 1932b, Falkenström 1938, Nilsson and Hilsenhoff 1991, Alarie and Hughes 2006). First instar larvae of Meladema can be distinguished from most other genera of Colymbetinae on: Additional setae present on dorsal margin of femur; short, spine-like mesofemoral seta FE5; additional setae on both ventral and dorsal margins of tibiae and relatively large size (Alarie and Hughes 2006). They can be separated from Bunites Spangler, 1972 on details of the setation of the coxae and abdominal segment 8 (Michat 2005). Third instar larvae are characterized by: relatively large size (head length > 4.40 mm cf. < 3.50 mm in other described Colymbetinae); urogomphus more elongate (> 1.7× dorsal length of last abdominal segment cf. < 1.5×); large number of secondary setae on legs and the presence of predominantly short and spine-like setae on outer and elongate setae with hair-like secondary setae along inner margin of urogomphus (see Alarie and Hughes 2006).

Compound eyes large, rounded, laterally somewhat protruding (Figure 3); anterior margin distinctly emarginate. Antennae long, slender, all segments elongate; segment 2 least so, 2.5× longer than broad; all other segments > 3× longer than broad; all segments broadening distally. Antennal insertions invisible dorsally, adjacent to anterior margins of compound eyes, below lateral margins of frons. Anterior margin of clypeus arcuate; anterior angles obtuse, weakly rounded. Clypeus with strongly transverse anterolateral foveae, each occupying approximately 0.25× length of anterior margin. Foveae shallow posteriorly, deep and abrupt anteriorly; furnished with stiff whitish or golden-yellow setae. Anterior margin of labrum with broad, semicircular emargination; central 0.4 furnished with stiff, close-set setae. Labium transverse, broadest anteriorly; anterior margin with brush of fine, close, elongate setae; lateral setae approximately 2× length of those at centre. Labial palpomeres elongate, particularly palpomere 2. Palpomere 3 with raised medial and apical tubercles ventrally, furnished with stout setae. Palpomere 3 expanded to apex, curved, convex ventrally and concave dorsally. Maxillary palpomere 1 slightly elongate; palpomeres 2–4 increasingly so. Palpomere 4 swollen, inner face almost straight, outer strongly curved; with setae on internal and ventral faces towards apex. Mentum setose anterolaterally; excavated in centre, with longitudinal ridges laterally. Pronotum strongly transverse, somewhat thickened laterally; without distinct lateral bead (Figure 3). Anterior angles acute, furnished with bunch of stiff, golden setae; posterior angles obtusely rounded. Anterior margin broadly arcuate around centre; posterior margin sinuate laterally or almost straight (Figure 3). Elytra elongate, broadest close to or behind middle (Figures 3, 4), with strong lateral bead; apices conjointly rounded. Each elytron with three rows of serial punctures (Figure 3). Each puncture of rows bearing 1–5 stout, recumbent to erect setae (e.g. Figures 3, 9, 10). Elytra of at least some individuals of all species with transverse, crescentic striolae (e.g. Figures 5, 6, 25, 26). Prosternum arched to tectiform; prosternal process lanceolate, bordered laterally, apex acuminately rounded. Metaventrite projecting anteriorly between mesocoxae; projection with elongate median groove to receive prosternal process; groove with marked central ridge, widening anteriorly. Discrimen and metacoxal suture strong, deep. Metacoxal lines strongly marked; metacoxal processes broadly rounded. Abdominal pleurite 2 without transverse ridges. Abdominal ventrites 2–6 with semicircular foveae laterally. Abdominal ventrite 6 with curved lateral wrinkles (Figures 11, 15).

Figure 3. 

Dorsal habitus of Meladema species males. A M. coriacea, Spain, Cáceres, nr. Plasencia B M. lepidoptera sp. n., Corsica, Francardo C M. imbricata, La Gomera, El Cedro D M. lanio, Madeira, Rabacal. Scale bar = 5 mm.

Figure 4. 

Meladema species males, colour pattern of isolated elytra (DNA voucher codes, where applicable). A M. coriacea, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., Corsica, Cap Corse (NHM-IRM12F) C M. imbricata, La Gomera, El Cedro (NHM-IRM3A) D M. lanio, Madeira, Ribeira dos Cedros (NHM-IRM8A). Scale bar = 5 mm.

Male. Foretarsi (Figure 12) with segments 1–3 strongly expanded and transverse, bearing four distinct rows of large articulo-setae ventrally; two on segment 1, one each on segments 2–3. Articulo-setal field bordered by dense, stiff, elongate, curved, hooked, golden setae. Segment 4 transverse; approximately 0.6–0.8 width of segment 3; with fields of stout, elongate setae of varying size laterally, inner setae curved anteriorly. Segment 5 elongate, with stout, elongate spines of varying size ventrally, close to lateral margins; basal spines curved interiorly. Foretarsal claws (Figures 12, 13) elongate, subequal, curved. Mesotarsi (Figure 12) with segments 1–3 expanded, transverse, bearing four distinct rows of large articulo-setae ventrally; two on segment 1, one each on segments 2–3. Articulo-setal field bordered by dense, stiff, elongate, curved, hooked, golden setae. Segment 4 elongate, with two rows of short, stout, stiff curved spines. Segment 5 strongly elongate, with two rows of stout spines ventrally, close to lateral margins. Mesotarsal claws elongate, subequal, curved. Abdominal ventrite 6 with weakly emarginated apex (Figure 11). Median lobe of aedeagus (Figure 14) elongate, strongly curved dorsally, with narrowly acuminate apex; lateral margins with transverse wrinkles in basal half; sinuate laterally towards apex in ventral view. Parameres (Figure 14) elongate, with dense setal fringe along internal margin, continued around apex, with scattered setae along external margin.

Female. Fore and mesotarsi simple, with ambulatory spines and setae only. Abdominal ventrite 6 (Figure 15) with bluntly pointed apex. Reproductive tract (Figure 16) with large, sclerotised bursa, with transverse corrugations and an elongate bursal gland; gland reddish in fresh material. Spermatheca elongate, fertilization and spermathecal ducts closely aligned, both relatively short. Gonocoxae (Figure 16) stout, elongate, with dense setation over entire surface and distinct apical penicil of setae. Laterotergites (Figure 16) elongate, with medial articulation. Gonocoxosternites (Figure 16) with bluntly pointed, setose apices. Female tract and genital structures do not appear to differ significantly between species and are not discussed further.

Both molecular and morphological data suggest a close relationship between Meladema and the Nearctic Hoperius Fall, 1927 and Neoscutopterus J. Balfour-Browne, 1943 (Morinière et al. 2015, 2016, Miller and Bergsten 2014). Female genital tract anatomy, here described for the first time, is similar to that described and figured for Neoscutopterus (Miller 2001).

Of these earlier works, only Aubé (1836) provides sufficient detail to demonstrate that he is referring to M. coriacea as redefined below. His description states: “entièrement couvertes de petites impressions demi-circulaires, plus profondes à la partie convexe; en avant elles sont assez bien isolées; en arrière elles sont un peu confondues”, details which allow the separation of M. coriacea from M. lepidoptera sp. n. (see below).

Laporte’s original description (1835) could refer to either M. coricaea as redefined here, or M. lepidoptera sp. n., the only reference to the unique elytral sculpture of these beetles being “corps couvert de points très serrés, presque chagriné”. As discussed by Evenhuis (2012), attempts to locate types of material described by Laporte between 1828 and 1840 have almost always proved fruitless, and it is generally assumed that this material is lost or destroyed. It seems Laporte donated his early collection (pre-1840 material) to the forerunner of the Smithsonian Institution in January 1842, and that these collections have been lost in a subsequent fire (Evenhuis 2012). As with other Laporte taxa, our attempts to locate the original type series of M coriacea have failed, no specimens being present in MNHN, despite Guignot (1961) suggesting that they may be located in this institution, nor in the BMNH or Melbourne Museum, Australia (S. Hinckley pers. comm.), the other two locations known to house some Laporte types (Sharp 1901, Horn et al. 1990). As a consequence, and in accordance with article 75.3 of the ICZN (1999), we here designate a neotype for M. coriacea, in order to establish its taxonomic identity and precise usage of the name. Laporte’s original description cites “Midi de la France”, a term used to describe a large part of the south of the country (not the centre, as suggested by Nilsson and Hájek, 2017a). All material we have genotyped, or examined, from southern France to date belongs to this same taxon.

“Midi de la France”.

Neotype ♀ (herein designated): “24/viii/2006 FRANCE Var// La Londe-les-Maures,// Vallon de Valcros, Les// Gaouby (ruines), pools in// Maravenne Torrent, 45m// P. Ponel leg.” “43°09'45.74"N 9°15'38.82"E” “DNA Voucher// NHM-IRM11C” “Meladema coriacea// Laporte, 1835// NEOTYPE// D T Bilton & I Ribera des. 2017” (NMW). Dry card mounted, tissue sample in ethanol and DNA aliquote, both with same data, retained in IBE. Sequence data from the neotype has been deposited in GenBank with accession numbers AF428207 (COI-3’) and AF428189 (16S ribosomal RNA).

Algeria: 1 ♂ “24/viii/2006 ALGERIA// Aïn Damous 36 25.350N// 07 51.367E 523m V67// S. Bouzid leg.” “Meladema// coriacea Laporte// Fery det. 2007” “DNA voucher// MNCN-HI6” (IBE); 1 ♀ “24/iii/2006 ALGERIA// Oued Bagrat V28// S. Bouzid leg.” “Meladema// coriacea Laporte// Fery det. 2007” “DNA voucher// MNCN-HI4” (IBE). Chad: 1 ♀ “KOUDOU// VERS 2000 m.// 17 XI 49” “TIBESTI// MASSIF KOUSSI// PH. DE MIRÉ” “MUSÉUM PARIS” [blue label] “DNA voucher// IBE-AN739” (MNHN). France, mainland: 1 ♀ “24/viii/2006 FRANCE Var// La Londe-les-Maures,// Vallon de Valcros, Les// Gaouby (ruines), pools in// Maravenne Torrent, 45m// P. Ponel leg.” “Meladema coriaceum// FRANCE, Dept. du Var.// La Londe-les-Maures,// Vallon de Valcros,// Les Gaouby (ruines),// pools in Maravenne// torrent, alto 45 m// 5 ix 99 P. Ponel” [HW] “DNA Voucher// NHM-IRM11A” (IBE); 1 ♀ “24/viii/2006 FRANCE Var// La Londe-les-Maures,// Vallon de Valcros, Les// Gaouby (ruines), pools in// Maravenne Torrent, 45m// P. Ponel leg.” “DNA Voucher// NHM-IRM11B” (IBE). Italy, Sicily: 1 ♂ “SICILIA – PA// Bosco Ficuzzo// im beraio// 29.VII.88 le. M. ROMANO” [im beraio & date HW] “DNA voucher// IBE-AN691” (CTP). Malta: 1 ♀ “23/iv/2013 MALTA// between Mosta and L-Imtarfa// stony stream 35 53 60N// 14 24 14E A. Rudoy leg.” “DNA voucher IBE-RA1064” (IBE). Morocco: 1 ♂ “17/vii/1997 MOROCCO// Tazzeka Nat. Park/ 4, ca.// 4 km tras desvío Cedral// I. Ribera leg.” “DNA voucher// NHM-IRM1A” (IBE); 1 ♂ “17/vii/1997 MOROCCO// Tazzeka Nat. Park/ 4, ca.// 4 km tras desvío Cedral// I. Ribera leg.” “DNA voucher// NHM-IRM1C” (IBE); 1 ♂ “2i/vii/1997 MOROCCO// Anti-Atlas, Oued Massa// I. Ribera leg.” “DNA voucher// NHM-IRM2A” (IBE); 1 ♂ “2i/vii/1997 MOROCCO// Anti-Atlas, Oued Massa// I. Ribera leg.” “DNA voucher// NHM-IRM2B” (IBE); 1 ex. “4 MOROCCO Taza 22.3.2008// mountain stream in Tazzeka N.P.// 1448m N34°03'09.2" W4°10'27.0"// Ribera, Hernando & Aguilera leg.” “DNA Voucher// NHM-IR47” (IBE); 1 ex. “58 MOROCCO 19.4.2001// 30°47'507"N 7°31'351"W// Tachokchte: Assif Siroua// c.1500 m I.Ribera & A.Cieslak leg.” “DNA Voucher// NHM-IRM22a” (IBE); 1 ex. “58 MOROCCO 19.4.2001// 30°47'507"N 7°31'351"W// Tachokchte: Assif Siroua// c.1500 m I.Ribera & A.Cieslak leg.” “DNA Voucher// NHM-IRM22b” (IBE); 1 ex. “60 MOROCCO 21.4.2001// 30°39'456"N 9°21'134"W// Immouzèr-des-Ida-Outanane// Assif Tanit, 550 m I. Ribera & A. Cieslak” “DNA Voucher// NHM-IRM21a” (IBE); 1 ex. “60 MOROCCO 21.4.2001// 30°39'456"N 9°21'134"W// Immouzèr-des-Ida-Outanane// Assif Tanit, 550 m I. Ribera & A. Cieslak” “DNA Voucher// NHM-IRM21b” (IBE). Spain, mainland: 1 ♂, “1 ES Girona, Port Bou// Cami de la Riera, 28.6.2013// 42°25'34.5"N 3°8'9.3"E// 65m I.Ribera & A.Cieslak leg.” “DNA Voucher// IBE-DV293” (IBE); 1 ex. “Tarragona// Corbera d’Ebre// r. Gaia 10.1.2004 I.Ribera” “DNA Voucher// MNCN-AI104”(IBE); 1 ♂, “3 ES Huesca 6km S Bernués// Bco. de Bernués 6.9.2013// 42°26'38"N 0°36'51"W// 690m I.Ribera & A.Cieslak leg” “DNA Voucher// IBE-DV292” (IBE); 1 ex. “1 ES Castellón, Ballestar// r. Sénia, pools upstr. reservoir// 500m N40°41'41" E0°13'25.5"// I. Ribera leg. 2.6.2006” “DNA Voucher// MNCN-AI860” (IBE); 1 ex. “1 ES Castellón, Ballestar// r. Sénia, pools upstr. reservoir// 500m N40°41'41" E0°13'25.5"// I. Ribera leg. 2.6.2006” “DNA Voucher// MNCN-AI861” (IBE); 1 ♂ “8 ES Cáceres, PN Monfragüe// Arroyo de Trasierra, G. del Fraile// N39°50'07.2" W6°06'02.6"317m// I.Ribera & P.Abellán leg. 13.6.2008” “DNA Voucher// IBE-DV291” (IBE); 1 ♂ “19/ix/1999 SPAIN Murcia// Fte. Caputa, Rio Mula// A. Millán leg.” “DNA voucher NHM-IRM13a” (IBE); 1 ♂ “29/ix/1999 SPAIN Cordoba// Baena, Arroyo de las Beatas// 36 37 00N 04 20 00W// M. Baena leg.” “DNA voucher// NHM-IRM14A” (IBE); 1 ♀ “29/ix/1999 SPAIN Cordoba// Baena, Arroyo de las Beatas// 36 37 00N 04 20 00W// M. Baena leg.” “CColeopteraBA 9.99// M. BAENA” “DNA voucher// NHM-IRM14B” (IBE); 1 ♂ “29/ix/1999 SPAIN Cordoba// Baena, Arroyo de las Beatas// 36 37 00N 04 20 00W// M. Baena leg.” “DNA voucher// NHM-IRM14C” (IBE); 1 ♀ “26/vii/1998 SPAIN Cadiz// Facinas// I. Ribera leg.” “CADIZ 98// 10a” [HW] “DNA voucher// NHM-IRM10A” (IBE). Spain, Mallorca: 1 ex. “4 MALLORCA Mortixet// Rd. C710 tributary Te. Son March// I.Ribera & A.Cieslak 12.11.2000” DNA Voucher// NHM-IRM23a” (IBE); 1 ex. “4 MALLORCA Mortixet// Rd. C710 tributary Te. Son March// I.Ribera & A.Cieslak 12.11.2000” DNA Voucher// NHM-IRM23b” (IBE); 1 ex. “5 MALLORCA Els Casals// Rd. C710 Te. Son March// I.Ribera & A.Cieslak 12.11.2000” “DNA Voucher// NHM-IRM24a” (IBE). Spain, Canary Islands: 1 ex. “2 Gran Canaria 14.4.2001// S. Nicolas de Tolentino// Risco: bco. Guy Guy grande// I.Ribera & A.Cieslak leg.” “DNA Voucher// NHM-IRM20a” (IBE); 1 ♀ “12/i/2000 Tenerife// Bco. Infierno 900m// coriacea” “DNA voucher// NHM-IRM18a” (CBP); 1 ♂ “14/i/2000 SPAIN Tenerife// Bco. de Masca// D. T. Bilton leg.” “DNA voucher// NHM-IRM19A” (CBP); 1 ♂ “14/i/2000 SPAIN Tenerife// Bco. de Masca// D. T. Bilton leg.” “DNA voucher// NHM-IRM19B” (CBP); 1 ex. “Tenerife, Chamorga// Bco. Roque Bermejo// 20.7.2006 A. Castro” “DNA Voucher// MNCN-AI1095” (IBE). Tunisia: 1 ♀ “13/iii/2005 TUNISIA// Cite el Morjne// 36 56 17.1N 8 47 29.8E// A. Castro leg.” “Cite-el-Morjene// N36°56'17.1// E8°47'29.8// 13-03-2005” reverse “Meladema” [HW] “MNCN AI-84// DNA spare ex.” [HW] “DNA voucher// MNCN-AI84” (IBE). Turkey: 1 ♀ “26/vii/2014 TURKEY Izmir// 6 km E of Phoca, head of// reservoir 38 39 37.3N// 26 49 23.9E 40m// I. Ribera & A Cieslak leg.” “TR 15” [HW] “DNA voucher// IBE-DV294” (IBE). All with “Meladema// coriacea Laporte, 1835// D T Bilton [or I Ribera] det. 2017”.

Additional material examined (non-genotyped specimens). Algeria: 8 ♂♂, 15 ♀♀ “Algerie// Yakouren// J Dayren// VI.VII. 1909” “MUSÉUM PARIS// 1952// COLL R OBERTHUR” [blue label] (MNHN); 1 ♂, 1 ♀ “Algerie// Yakouren// J Dayren// VI.VII. 1909” “MUSÉUM PARIS// 1952// COLL R OBERTHUR” [blue label] “Meladema// coriaceum// Cast.// de. M. Brancucci ‘82” [Latin name, describer & ’82 HW] (MNHN); 1 ♀ “Algerie// Yakouren// J Dayren// VI.VII. 1909” “MUSÉUM PARIS// 1952// COLL R OBERTHUR” [blue label] “Meladema// coriacea// Cast.” [HW] (MNHN); 1 ♀ “Meladema// coriaceum// O. Kaïrous, Mouz.// 8 avríl 06” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♀ “O. Kaïrous// mouzaïa// 8 avríl 06” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ “Meladema// coriaceum// Djurdj. 9” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ “Djurdjura// 14.VIII.1947” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♀ “Aïn Takrarat// Mouzaïa// 28.VII.1934” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ “Oued In Dalay// Kouddu// Hoggar// 19.3.1925” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂, 1 ♀ “Aguelmane// Sekkarasen// Hoggar 206// 29.3.1928” [HW] “MISSION DU// HOGGAR// FÉVRIER-MAI// 1928” “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ “Oued Tinikert// Mt. Oudan, 1160// 16 avril 1928” [HW] “MISSION DU// HOGGAR// FÉVRIER-MAI// 1928” “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♀ “Tassili des Ajjers// Gta. de Tikkal// 24 mars 1961” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ [green circle, illegible HW] “Meladema// coriaceum// Bône” [HW] (ISNB); 1 ♂ “C. coriaceus// Alger” [HW] “G. C. Champion Coll.// B. M. 1927-409” (BMNH); 1 ♂ “27// 95” “Setif.// Algerien” [HW] “Meladema// coriacea Lap.// det. H. Shaverdo 2015” [Latin name, describer & 15 HW] (NMW). Chad: 1 ♂ “KOUDOU// VERS 2000 m.// 17 XI 49” “TIBESTI// MASSIF KOUSSI// PH. DE MIRÉ” “Meladema// coriaceum Cast.// C. Legros det.” [Latin name & describer HW] (MNHN); 2 ♀♀ “KOUDOU// VERS 2000 m.// 17 XI 49” “TIBESTI// MASSIF KOUSSI// PH. DE MIRÉ” (MNHN); 1 ♀ “Bassin de Gorrom// 2500m.// 24 déc. 1958” “TIBESTI// Emi Koussi// Bruneau de Miré” “DNA voucher// IBE-AN740” [not possible to amplify any DNA sequences] (MNHN); 1 ♀ “Tibesti// S. O du Kohoz// 2000m// 6.XI.49 Miré” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN). France, mainland: 1 ♂, 1 ♀ “S. France,// Cerbere.// C. Thomas &// R. L. Pocock.// 1900-215.// 18.IV.00” [date HW] (BMNH); 1 ♂ “S. France,// Cerbere.// C. Thomas &// R. L. Pocock.// 1900-215.// 18.IV.00” [date HW] “Scutopterus// coriaceus Aub,” [HW] (BMNH); 1 ♀ “S. France,// Cerbere.// C. Thomas &// R. L. Pocock.// 1900-215.// 18.IV.00” [date HW] “Meladema// coriacea Cast.// C. R. Smith det. 1982” [Latin name, describer & 2 HW] (BMNH); 1 ♂, 3 ♀♀ “ BANYULS// PYR. OR.// 12.8.1950” [12.8. HW] “BACHUFER” (NMW); 2 ♂♂, 1 ♀ “Banyuls s. Mer// 17.25 – V – 1951// F. G. Overlaet” “R, Mouchamps det.// Meladema coriacea Cast.” [Latin name & describer HW] (ISNB); 1 ♂, 2 ♀♀ “Banyuls s. Mer// 17.25 – V – 1951// F. G. Overlaet” (ISNB); 1 ♂, 1 ♀ “21.8.84 France// Pyr. Or., ung. Banyuls// Bach Fery leg.” [HW] “Mel.// coriacea// Cast. Fery det.” [HW] (NMW); 1 ♂ “meladema// coriaceum” [HW] “Foix” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 1 ♂ “AVEYRON FR.// ST. ROME CERN.// VII-VIII 1940” “Coll. R. Van Dorsselaer” (ISNB); 1 ♀ “Provence” “Coll. P. Boppe// Le Moult vendit” (ISNB); 3 ♀♀ “France. 10.10.55.// Provence.// leg. Weygoldt.’’ reverse “Coriaceum// Budberg” [HW] (NMW); 1 ♂ “Carcassonne// L. GAVOY” “♂” “MELADEMA// CORIACEUM// LAP.” (ISNB); 1 ♀ “Carcassonne// Aude” [HW] “Ex. Coll. Bettinger” (ISNB); 1 ♀ “Toulon.” [HW] “Gallia// mer.” “Collectio// Kaufmann” (NMW); 1 ♂, 2 ♀♀ “Toulon// S. Frankreich” reverse “A// 7598” [HW] (NMW); 1 ♂, 1 ♀ “Gard” [HW] “ex coll R P David// ex coll Peres Jesuites// (le moult vendit)” (ISNB); 1 ♂ “La Sauzette” “Meladema// coriaceum” [HW] (ISNB); 1 ♂ “St. Etienne Valléefranc. de// s// Rogerie” [HW – Villefranche de Rougerie?] “Coll. A. Fauvel” “Coriacea// Lap.” [HW] (ISNB); 1 ♂ “CAUX// HERAULT// IX 1928” [IX & 8 HW] “Collection// Dr. Guignot” “♂” “Guignot det// MELADEMA// coriacea Cast.” [Latin name & describer HW] (ISNB); 1 ♂ “CAUX// HERAULT// IX 1928” [8 HW] “♂” “MUSÉUM PARIS// 1960// Coll F. Guignot” “coriacea Cast.” [HW] (MNHN); 2 ♀♀ “CAUX// HERAULT// IX 1928” [8 HW] “♀” “MUSÉUM PARIS// 1960// Coll F. Guignot” (MNHN); 2 ♂♂ “neffies// HERAULT// H. LAVAGNE” [neffies HW] “Coriaceum” [HW] “Ex Coll. Bettinger” (ISNB); 1 ♀ “St GUILHEM// HERAULT// H. LAVAGNE” “Coriaceum” [HW] “Ex. Coll. Bettinger” (ISNB); 1 ♀ “Meladema// coriaceum” [HW] “Montpellier// (Hérault)” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 1 ♀ “Béziers// (He^lt.)” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 2 ♂♂ “Lozère” [HW] “Meladema// coriaceum” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 2 ♂♂ “Lozère” [small, circular brown label, no text] “Coll. Odier.// B. M. 1921-288” (BMNH); 1 ♀ “♀// 5188” [HW] “Coll. R. I. Sc. N. B.// France Marseille// Coll. Fairmaire// Coll. L. Pandellé” [Marseille & Coll. Fairmaire HW] “Coll. A. Fauvel// (ex. coll. Pandellé)// Meladema// coriacea Lap.// R.I.Sc.N.B. 17.819” [Latin name & describer HW] (ISNB); 1 ♀ “♀” [HW] “Coll. R. I. Sc. N. B.// France Marseille// Coll. Fairmaire// Coll. L. Pandellé” [Marseille & Coll. Fairmaire HW] “Coll. A. Fauvel// (ex. coll. Pandellé)// Meladema// coriacea Lap.// R.I.Sc.N.B. 17.819” [Latin name & describer HW] (ISNB); 1 ♂ “♂// 5188” [HW] “Coll. R. I. Sc. N. B.// France Marseille// Coll. Fairmaire// Coll. L. Pandellé” [Marseille & Coll. Fairmaire HW] “Coll. A. Fauvel// (ex. coll. Pandellé)// Meladema// coriacea Lap.// R.I.Sc.N.B. 17.819” [Latin name & describer HW] (ISNB); 1 ♂ “♂// 5188” [HW] “Coll. R. I. Sc. N. B.// France Marseille// Coll. Fairmaire// Coll. L. Pandellé” [Marseille & Coll. Fairmaire HW] “Coll. A. Fauvel// (ex. coll. Pandellé)// Meladema// coriacea Lap.// R.I.Sc.N.B. 17.819” [Latin name & describer HW] “Colymbetes Clairville// coriaceus, Hoffm.// Marseilles” [HW] (ISNB); 6 ♂♂, 2 ♀♀ “St Cyr// Ravindu// Degoutant” [HW] (MNHN); 1 ♀ “Le Beausset.// Octobre” “Meladema// coriaceum Lap.” [HW] (NMW); 1 ♀ “Le Beausset.// Mol. de Boissy” (NMW); 1 ♀ “M. coriaceum” [HW] “M. de Boissy// Le Beausset Var” [HW] (NMW); 1 ♀ “Rhonetal// mont. ol. Maures// östl. Aixen Provence” [HW] “M. J. MAURES// Det. VI/81//511” [HW] (NMW); 1 ♂, 1 ♀ “Nice” [HW] “coll. P. J. Roeflofs” (ISNB); 1 ♀ “Gallia merid// Agay (Var)// 15 Juli 14// W. Liebmann// Arnstadt” [15 HW] (NMW); 2 ♂♂ “Gallia merid// Agay (Var)// 15 Juli 14// W. Liebmann// Arnstadt” [15 HW] “Collectio// Paganetti” (NMW); 1 ♂, 1 ♀ “St. Maxime// Var// G. Audran” [HW] “R. Mus. Hist. Nat.// Belg. I.G. 14.406” (ISNB); 1 ♀ “Hyères” [HW] “coll. A. Fauvel” (ISNB); 1 ♂ “Nyons” [HW] “Colymbetes// coriaceus// 22” [HW] “coll. Delgrange” (ISNB); 1 ♀ “Colymbetes// coriaceus// Capt. P. Bauret” [HW] “Trinité-Victor// Juillet 1942” [HW] (ISNB); 1 ♂ “Colymbetes// coriaceus// Lap.” [HW] “Oct. 46// Le Laghet// Trinite Victor” [HW] (ISNB); 1 ♀ “Colymbetes// coriaceus// Lap.” [HW] “October 1945// Le Laghet// Trinité Victor” [HW] (ISNB); 1 ♂ “Cavalaire” [HW] “f. 532 Coll.//Dr. Mouchamps” [32 HW] “Pavel Riha leg.// MELADEMA// coriacea Cast.” [Latin name & describer HW] (ISNB); 3 ♀♀ “261// 131” [blue, circular label, HW] reverse “Savoy” [HW] (BMNH); 1 ♂ “[illegible]// Coll. Chevrolat// Det. Sharp 82.” [HW] “3306” “Colymbetes// coriaceus// Ex Aubé 220,// Gall. mer.” [HW] (ISNB); 1 ♂ “Gallia m.” (NMW); 1 ♂ “8367” [blue-green label, HW] “Gallia” [HW] “Fry Coll.// 1905-100” (BMNH); 1 ♀ “Gallia” [HW] “Fry Coll.// 1905-100” (BMNH); 1 ♀ “Colymbetes// coriaceus. Hoffm.// Gehin France” [HW] (BMNH); 1 ♀ “Coriaceus// Gallia” [blue label, HW] “Sharp Coll.// 1905-31.” (BMNH); 1 ♀ “Gallia 10” “Coll. Liepolt” (ISNB); 1 ♂ “coriaceus Lap.// Gallia// Bien Bacher” [HW] “c. Epplsh.// Steind. d.” (NMW). Greece, mainland: 1 ♂ “Attica// Dr. Kinper” [HW] “Collect.// Hausser” (NMW); 1 ♀ “Attica// Dr. Kinper” [HW] “punctulatus” [HW] “Collect.// Hausser” (NMW). Greece, Aegina: 1 ♀ “Griechenland// Aegina// F. Werner, 13 V. 37” [Aegina & 13 HW] (NMW). Greece, Amorgos: 1 ♀ “Graecia, KYKL.// Amorgos// P. Velensky lgt.// 13.-16.5.1984” “Coll.// HENDRICH// Berlin” (ZSM); 1 ♀ “Graecia, KYKL.// Amorgos// 13.-16.5.1984// P. Velensky lgt.” “Coll.// HENDRICH// Berlin” (ZSM). Greece, Chios: 1 ♀ “GR: Chios// Mai 1980// lg. Bilak, Kritscher” (NMW). Greece, Corfu: 1 ♂, 1 ♀ “GREECE: NE Corfu (GR3)// ca. 1.5 km W Zigos// 200 m a.s.l., 19.VII.08// leg. M. A. Jäch” “small shallow residual// pool in river bed// 39°43'44.4"N// 19°46'38.3"E” “Meladema// coriacea Laporte// Fery det. 2010” (NMW); 1 ♂, 2 ♀♀ “28/v/2017 Greece Corfu// nr. Zigos 233 m// 39 43 40.6N 19 46 41.0E// C R Turner leg.” (CBP). Greece, Ios: 1 ♂ “Kykladen// Ios, Buch von// Milopotamos” “leg. Schönmann// 9. April 1981” [first 9 HW] (NMW); 1 ♀ “Kykladen// Ios, Buch von// Milopotamos” “leg. Schönmann// 8. April 1981” [first 8 HW] (NMW). Greece, Kithira: 1 ♀ “GR-Kithira (4)// Arlemonas 8.V.76// leg. Malicky” “Melaema// coriacea Cast.// det. Wewalka 76” [HW] (NMW). Greece, Milos: 3 ♂♂, 1 ♀ “GR-MILOS 19.9.90// 1km E. Sider ianos.// leg. M. Jäch (5)” (NMW). Greece, Naxos: 1 ♀ “GR-Naxos V.1988// leg. Bilak// et Kritscher” (NMW); 2 ♀♀ “GR-Naxos (13)// Potamia 26.V.76// leg. Malicky” (NMW). Greece, Poros: 1 ♂ “GR-Poros (2)// Kampos 17.V.74// leg. Malicky” (NMW). Greece, other: 1 ♀ “Griechenland// Kylhima// F. Werner, 26.V.37” [Kylhima & 26 HW] (NMW); 1 ♂ “Greece// Merlin Coll.// 96-275” (BMNH); 1 ♂ “Greece// Merlin Coll.// 96-275” “Spec. nov.// Hymettus” [pink label, HW] “Received with// this name// from Merlin// C. O. W.” [Merlin HW] (BMNH); 1 ♂ “Graecia” “Meladema// coriaceus” [HW] “Coll. Plason” (NMW). Italy, mainland: 1 ♂ “Grotta di// Pastena// Italie merid.” [HW] “Meladema// coriacea Cast.” [HW] “Brit. Mus.// 1963-344.” (BMNH); 1 ♂, 2 ♀♀ “Meladema// coriaceum” [HW] “Italie” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 1 ♂ “Coriaceus// Italia F” [HW] “R. Mouchamps det.// Meladema// coricaea Cast.” [Latin name & describer HW] (NMW); 1 ♀ “Meladema// coriaceum” [HW] “Calabre” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 1 ♀ “Gerace, Calab.// Paganetti” “Collectio// Paganetti” (NMW); 1 ♀ “Gerace, Calab.// Paganetti” “Collectio// Paganetti” “Meladema// coriaceum” [HW](NMW). Italy, Pontine Islands: 2 ♂♂, 3 ♀♀ “Ins. Pont. - @ nos// auf Tamanis// 12-4-57// Dr. Eckerlein leg. [HW] (NMW). Italy, Sicily: 1 ♂ “Madricia// Ragusa” [HW] “3306” (ISNB); 1 ♀ “Monts Madonie 74” “E. Ragusa” (BMNH); 1 ♀ “Castelbueno// Ragusa” [HW] “3307” (ISNB); 2 ♂♂, 1 ♀ “Sicil.” “Collectio// Kaufmann” (NMW); 1 ♀ [circular label, black margin, no text] “Sicilia” [HW] “alte// Sammlung” (ZSM); 1 ♀ “Sic.” “Sammlung// C. Müller” (ZSM); 1 ♀ “Italia: Sicilia – PA// MTE. Maganoce// 900 M. Ü. N.” [HW] (ZSM). Malta: 2 ♀♀ “Prof. Barthet// Malta” “29.8.25// Malta” [HW] “R.L.S.N.B.//I.G. 17.619” (ISNB); 1 ♀ “Malta” [HW] “Ex. Coll. Bettinger” (ISNB); 1 ♂ “Malta” “G.C. Champion Coll.// B. M. 1929-409” “Meladema// coriaceum Lap.// Malta” [HW] (BMNH); 1 ♂ “Buskett Gdns.// Malta// 12.XII.55.// G. V. P. Sewell” [HW] “Meladema// coriacea Cast// C. R. Smith det. 182” [Latin name, describer & 2 HW] (BMNH); 1 ♂ “MALTA// Chadwick Lakes// Mars ‘85// D. Johnson” [HW] (NMW). Morocco: 1 ♂, 1 ♀ “Tachdirt// Marocco// Schwingenschuss” [HW] “Coll. Liepolt” (ISNB); 7 ♂♂, 5 ♀♀ “Maroc: Azrou// 6-12-vii-1934 (1350m)// A. Ball 56 M 22” “R. Mus. Hist. Nat.// Belg. I.G. 10.417” “A. Ball det. 1935// Meladema// coriacea Cast.” [1935, Latin name & describer HW] (ISNB); 1 ♀ “Middle Atlas Mts// Oviduane// 2,000ft.” “MOROCCO:// K. Chapman &// J.W.S. Pringle.// B.M. 1934-554” ‘MELADEMA// coriacea// Laporte” [HW] “det. J. Hajek// xi. 2008” (BMNH); 4 ♂♂, 4 ♀♀ “Middle Atlas Mts// Oviduane// 2,000ft.” “MOROCCO:// K. Chapman &// J.W.S. Pringle.// B.M. 1934-554” (BMNH); 1 ♂ “Middle Atlas Mts// Oviduane// 2,000ft.” “MOROCCO:// K. Chapman &// J.W.S. Pringle.// B.M. 1934-554” “Oum en Rhio Aug 20^th// large beetle 400” [folded, HW, 400 in a box] (BMNH); 1 ♀ “MOROCCO// Dayet Ifrah (lake)// 16 km. E.N.E. Ifrane// 28.V.1961.” [all except Morocco HW] “1334// P. N. Lawrence// B. M. 1961-328” [HW] (BMNH); 1 ♀ “MOROCCO// Great Atlas Mts.// Ijoukak. 3,900ft.// 3.vi.1936” [3 HW] “K. H. Chapman// & G. A. Bisset.// B. M. 1936-527.” (BMNH); 1 ♀ “S. MORROCCO// TIZI-N-BACHKOUM env.// 25.4.1995// P. Prüdek leg.” “Coll.// HENDRICH// Berlin” “Meladema// coriacea// Cast.// HENDRICH det. 1996” [name & describer HW] (ZSM); 1 ♂ “Imi n’ Ouaka// 1500m.” “Maroc// 1-15 Sept.” “♂” “♂” “MUSÉUM PARIS// 1960// Coll F. Guignot” (MNHN); 2 exx. “71 MOROCCO 8.4.2007// Tizi-n’Rechou, Kerrouchèn// stream on rock N pass, rd. 3437// 1570m N32°48'22.1" W5°16'13.9"// Aguilera Hernando & Ribera leg.” (IBE); 1 ♀ “99 MOROCCO Chefchauen// 13.2.2015 afl. Oued Laou 150m// 35°17'57.0"N 5°12'55.5"W// Ribera, Millán & Velasco leg.” (IBE); 1♂, 1♀ “Morocco// Âit-Iftene// 29, Oued Ait-Baha// 220797 Ribera, Aguilera, Hernando, Millán” (IBE); 1♂, 1♀ “Morocco// Sidi-Ibrahim// 17, 77.5 km S Guercif// 150797 Ribera, Aguilera, Hernando, Millán” (IBE); 1 ex. “MOROCCO Haut Atlas// NE Tizi-n-Test, 1710m No.18// 30°54'12N 08°18'39W// 30.12.2002 V.Assing & P.Wunderle” (IBE). Portugal, mainland: 1 ♂ “Portugal// Oliveira” [HW] “3306” (ISNB); 3 ♀♀ “Portugal// Oliveira” [HW] “3307” (ISNB); 1 ♀ “Portugal// Oliveira” [HW] “♀” (ISNB); 1 ♂ “Portugal// Oliveira” [HW] (ISNB); 8 ♂♂, 10 ♀♀ “PORTUGAL:// Serra do Malhao.// 8.v.1966// J. Abraham &// M. E. Bacchus.// B. M. 1966-296” “M. E. Bacchus// B. M. 1966-296” (BMNH); 1 ♀ “PORTUGAL:// Algarve. Rio Seco.// 250m ca. 10km.// W. Barranco de// Velho. 10.v.1966.” (BMNH); 1 ♀ “PORTUGAL:// Algarve. Rio Seco.// 250m ca. 10km.// W. Barranco de// Velho. 10.v.1966.” “Meladema// coriacea cast.// J. Balfour-Browne det.// xii. 1972” (Latin name, describer & 72 HW) (BMNH); 1 ♀ “5/v/93 Portugal// Viana do Castelo// Serra do Minho// stream @ ca. 700m// D. T. Bilton leg.” [HW] (CBP); 1 ♀ “PORTUGAL: Traz os Montes// Carrazeda de Anciais// 745m., 7.viii.1966” “slow weedy// stream” “Brit. Mus.// 1973-562” “Meladema// coriacea Cast.// J. Balfour-Browne det.// IX 1969” [Latin name, describer, IX & last 9 HW] (BMNH). Spain, mainland: 1 ♂, 2 ♀♀ “Cataluna// Vall. x-42// Español” [HW] “R. Mouchamps det 1948// MELADEMA// coriacea Cast.” [8, Latin name & describer HW] (ISNB); 2 ♂♂ “Vallvidrera// Barcelona// VIII.1948// Español” [HW] “R. Mouchamps det.// Meladema// coriacea Cast.” [Latin name & describer HW] (ISNB); 2 exx. “Barcelona// Montseny d’Amunt// Riachuelo: remanso// 290988 I. Ribera” (IBE); 1 ♂ “Girona// Capmany// Estanys: alberca// 041189 I.Ribera” (IBE); 3 ♂♂, 1 ♀ “Girona// Capmany// Estanys: alberca// 110389 I.Ribera” (IBE); 4 ♂♂, 3 ♀♀ “Girona// Capmany// Estanys: alberca// 281090 I. Ribera” (IBE); 6 ♂♂, 3 ♀♀ “Girona// Capmany// Estanys: alberca// 251190 I. Ribera” (IBE); 4 ♂♂, 3 ♀♀ “Girona// Capmany// Estanys: alberca// 120191 I.Ribera” (IBE); 2 ♀ “Girona// Capmany// Estanys: alberca// 171294 I. Ribera & P. Aguilera” (IBE);1 ♂ “Girona// Capmany// Estanys: alberca// 100994 I.Ribera & P.Aguilera” (IBE); 2 exx. “1 GIR La Junquera 4.x.2014// estanys de Capmany: alberca// 42°24'20.5"N 2°54'8.0"E// 188m I. & B.Ribera leg.” (IBE); 1 ♂ “Girona// Capmany// Querafumat: abrevadero 1// 251190 I. Ribera” (IBE); 2 ♂♂, 1 ♀ “2 ES Girona, Port Bou// Cami de la Riera, 10.5.2012// N42°25'38.2" E3°08'13.4"// 81m I.Ribera & A.Cieslak leg.” (IBE); 2 ♂♂, 2 ♀♀ “Girona// Sta. Pau// Rio Ser: remanso 1b// 260890 I. Ribera” (IBE); 1 ♀ “Girona// Sta. Pau// Rio Ser: remanso 2// 260890 I. Ribera” (IBE); 2 ♂♂, 1 ♀ “1 ES Girona, Port Bou// Cami de la Riera, 28.6.2013// 42°25'34.5"N 3°8'9.3"E// 65m I.Ribera & A.Cieslak leg.” (IBE); 1 ex. “Tarragona// Vila-rodona// Jardín: alberca// 110985 M.Galán” (IBE); 1 ex. “Tarragona// Vila-rodona// Jardín: alberca// 201085 I.Ribera” (IBE); 2 ♂♂, 2 ♀♀ “16/v/1994// SPAIN Tarragona// stream 2km SW of// Esblada D. T. Bilton leg.” (CBP); 1 ♂, 3 ♀♀ “3 ES Huesca 6km S Bernués// Bco. de Bernués 6.9.2013// 42°26'38"N 0°36'51"W// 690m I.Ribera & A.Cieslak leg” (IBE); 3 exx. “1 ES Castellón, Ballestar// r. Sénia, pools upstr. reservoir// 500m N40°41'41" E0°13'25.5"// I. Ribera leg. 2.6.2006” (IBE); 1 ♂, “Teruel// Calaceite// Riu Matarranya// 240794 I.Ribera & P.Aguilera” (IBE); 1 ♂, 1 ♀ “19/ix/1999 SPAIN Murcia// Fte. Caputa, Rio Mula// A. Millán leg.” (IBE); 1 ♂ “19/ix/1999 SPAIN Murcia// Fte. Caputa, Rio Mula// A. Millán leg.” “Fte Caputa// R. Mula 19-9-99// A. Mellado” [HW] (IBE); 1 ♂ “19: SPAIN: CACERES:// 21 km S of Aldeanuo// del Camino: granite// stream 18 April 1985” (CBP); 1 ♂, 1 ♀ “12/v/1990 SPAIN Cáceres// stream with pools 7km N of// Plasencia on N630 road” (CBP); 2 ♀♀ “27/iv/1993// Spain Extremadura// Cáceres stream by N630// 15km N of Plasencia” (CBP); 1 ex. “3 ES Cáceres, PN Monfragüe// Arroyo de Malvecino// N39°51'04.0" W6°02'22.3" 284m// I.Ribera & P.Abellán leg. 12.6.2008” (IBE); 1 ♀ “SPAIN: Cordoba.// Posadas – Villaviciosa Rd.// 20 Km N. of Posadas// Arroyo Calderas. 500m.// 14.V.1967” “Meladema coriacea// CAST. I. RIBERA det 1990” [HW] (BMNH); 1 ♂ “MALAGA// H. Clark.// May 1856” “Fry Coll.// 1905-100” (BMNH); 2 ♀♀ “MALAGA// H. Clark.// May 1856” (BMNH); 1 ex. “1 ESP. Malaga 3.1.03// Villanueva del Trabuco// r. Guadalhorce, cta. MA156// I.Ribera & A.Cieslak leg.” (IBE); 1 ♂, 1 ♀ “Almería// Tabernas// Rbla. de Tabernas: charca 1// 110597 I.Ribera & A.Millán” (IBE); 1 ♂ “Nerja” [HW] (MNHN); 1 ♂ “24/iv/1993// Spain Andalucia Cadiz// Drying stream by N340// 28km N of Tarifa” (CBP); 1 ♀ “17: SPAIN: HUELVA: 1 km// S. of Santa Ollala del// Cala: weedy stream:// 18 April 1985// R.B. Angus & G.N. Foster” “Meladema corac” [HW] (BMNH); 2 ♂♂, 1 ♀ “17: SPAIN: HUELVA: 1 km// S. of Santa Ollala del// Cala: weedy stream:// 18 April 1985// R.B. Angus & G.N. Foster” “R. B. Angus// BMNH(E) 2010-22” (BMNH); 1 ♂, 1 ♀ “Hispania” [HW] “Meladema// Lap.// coriacea// Cast.” [HW, genus name on green label, glued on top of original] “Coll. Martin// Le Moult vendit” (ISNB); 1 ♂, 1 ♀ “Hispan” [HW] “Fry Coll.// 1905-100” (BMNH); 1 ♂ “Hispa// nia” [HW] “57.// 107.” [blue, circular label, HW] (BMNH). Spain, Mallorca: 2 exx. “1 Mallorca, Lloseta 13.5.2007// te. des Estorall, potabilizadora// 170m N39°43'49.7" E2°50'04.6"// I.Ribera & A.Cieslak leg.” (IBE); 1 ♂, 1 ♀ “2 Mallorca, Bunyola 13.5.2007// te. de Bunyola, Ma 2100 pk 5.5// 455m N39°43'24.7" E2°43'15.7"// I.Ribera & A.Cieslak leg.” (IBE) ; 1 ♂ “Inca, Majorca.// March.// O. Thomas &// R. I. Pocock.// 1900-215” (BMNH). Spain, Canary Islands: 2 ♀♀ “Canary Is.// Gran Canaria,// Sta Brigida.// 5.ix.1927.// E. Appenhagen.// B. M. 1928-20” [5 & ix HW] (BMNH); 1 ♂ “Canary Is.// Gran Canaria,// Sta Brigida.// 5.ix.1927.// E. Appenhagen.// B. M. 1928-20” [5 & ix HW] “Colymbetes// coriaceus, Cast.// 5-9-27.// 206” [HW] (BMNH); 1 ♀ “CANARY IS.// Gran Canaria// 29.xii.92-11.i.93” “Bco de// Ayagaures// ca 100 m.” [HW] “P. M. Hammond// B. M. 1993-11” “coriaceus” [HW] (BMNH); 1 ♂ “Barranco de// Temise près Haria// (Lanzarote) 2” [HW] “coll. A. Fauvel” (ISNB); 1 ♀ “Colymbetes// coriaceus Hoffm.// Bajamar XI. 1909” [HW] “Le Moult vend.” (ISNB); 1 ♂ “Teneriffa// geibelez” [HW] “DON// P DUPUIS” (ISNB); 1 ♀ “Colymbetes Clairv.// 203 coriaceus Lap.// 6 ex Teneriffe 1904 (Prevost):Elliott Coll.” [HW, folded] “Claude Morley// Collection// B.M. 1952-159” “Meladema// coriacea Cast.// C. R. Smith det. 1982” [Latin name, describer & 2 HW] (BMNH); 2 ♂♂, 1 ♀ “Claude Morley// Collection// B.M. 1952-159” [part of same series as previous] (BMNH); 1 ♂ “Claude Morley// Collection// B.M. 1952-159” “Meladema// coriacea Cast.// C. R. Smith det. 1982” [Latin name, describer & 2 HW] [part of same series as previous] (BMNH); 2 ♂♂, 2 ♀♀ “Canary Is.// Teneriffe,// Guajonje,// 17.iii.1927// E. Appenhagen” [Guajonje & date HW] (BMNH); 1 ♀ “Canary Is.// Teneriffe,// Guajonje,// 17.iii.1927// E. Appenhagen” [Guajonje & date HW] “Colymbetes// coriaceus, Lap.// G. J. Arrow det.” [Latin name & describer HW] (BMNH); 1 ♂ “Tenerife// Barr. Bufadno” [HW] “6.2.1949// Fernandez” [6, 2 & last 9 HW] “Brit. Mus.// 1951-318” “Meladema// coriaceum Lap.// Harald Lindb. det” [Latin name & describer HW] (BMNH); 1 ♂ “ES: Tenerife// Masca 12 April// 1991 AN Nilsson” “Meladema coriacea// Laporte, 1835// Det AN Nilsson 1991” (CBP); 1 ♂ “April 1998 SPAIN// Islas Canarias Tenerife// Bco. del Infierno Pools// D.T. Bilton leg.” (CBP); 1 ♂, 1 ♀ “April 1998 SPAIN// Islas Canarias Tenerife// Bco. de Masca Pools// D.T. Bilton leg.” (CBP); 1 ♀ “29/xi/97 Tenerife// Anaga Massif – stream// @ Iguana D.T. Bilton” [HW] (CBP); 1 ♀ “CANARIES” “Coll. R. Van Dorsselaer” (ISNB); 1 ♀ “Canary Is.// 1903 (Prevost)// coll. Elliott . 36” [HW] “Claude Morley// Collection// B.M. 1952-159” (BMNH). Tunisia: 3 ♀♀ “TUN.: 3.8.91(7)// Beni Melir SE// A. Draham, Schödl” (NMW). Turkey: 1 ♀ “Asia min. occ.// Izmir (H. Mudja)// 20 m. St. 71// 8-V-’31 (Orch.)” “Reg. Mus. Hist. Nat.// Belg. I.G. 9642” “A. Ball det. 1935// Meladema// coriacea Cast.” [1935, Latin name & describer HW] (ISNB). Without locality data: 1 ♂ “Coll Abcedal” [HW] “Colymbetes// coriaceus” [HW] “R.I.Sc.N.B. 21.418// Coll. P. de Moffarts” “Meladema// coriaceum// Lap.” [green label, HW] (ISNB); 1 ♂ “Coll Abcedal” [HW] “Coll. P. de Moffarts” (ISNB); 1 ♀ “Meladema// coriaceum// Coll. Hevon” [HW] “Ex coll. Bettinger” (ISNB); 1 ♂ “121” “Meladema// coriaceum” [HW] “Ex. Coll. Bettinger” (ISNB); 1 ♂ “ex coll R P David// ex coll Peres Jesuites// (Le Moult vendit)” “coriaceus Cast.// Region mediterranee” [HW] (ISNB); 1 ♂ “Coll. P. Boppe// Le Moult vendit” “Meladema// coriaceum// Cast.” [HW] (ISNB); 1 ♂ “Meladema// coriaceum” [HW] “Coll. E. Dongé// Le Moult vendit” (ISNB); 1 ♂ “H Guyon” [HW] “Coll. E. Traizet// Le Moult vendit” (ISNB); 7 ♂♂, 4 ♀♀ “coll. A. Fauvel” (ISNB); 1 ♂ “coll. Delgrange” (ISNB); 1 ♂ “G. C. Champion Coll.// B. M. 1927-409” (BMNH); 1 ♀ “Coll. Odier.// B. M. 1921-288” (BMNH); 2 ♀♀ [small, circular brown label, no text] “Coll. Odier.// B. M. 1921-288” (BMNH); 1 ♂ “Alono-// dorsar” [? HW] “Meladema// coriacea” [HW] (BMNH); 1 ♂ “Europa// B. Heyne” [HW] (BMNH); 1 ♂ “coriaceus Lap.” [folded, HW] (BMNH); 1 ♂ “Dytiscuscoriaceus// no. in list. 19.” [HW] reverse “Presented by// Signor Passerini” [name HW] (BMNH); 1 ♂ “1911” [round label, HW] (BMNH). All with “Meladema// coriacea Laporte, 1835// D T Bilton [or I Ribera] det. 2017”.

Size: Neotype TL = 22.66 mm; EL = 16.90 mm; MW = 11.52 mm. Other material examined TL = 18.56–23.17 mm; EL = 14.34–16.90 mm; MW = 8.45–11.14 mm.

Colour. Dorsum dark reddish brown to black (Figure 3A); lateral margins of pronotum, labrum and anterior half of clypeus somewhat paler, sometimes with diffuse lateral maculae. Elytra unicolorous, without distinct mottling even when lifted (Figure 4A). Head with a pair of oval, reddish yellow medial interocular patches, slightly elongated apicolaterally. Antennae and maxillary and labial palpi reddish yellow. Legs dark reddish brown to black with golden yellow setae; large spines somewhat paler. Venter reddish brown to black; gula, meso and metacoxae and trochanters paler.

Head. Labrum shining, with moderate to coarse, sparse punctures. Reticulation absent in apical half, becoming increasingly more evident basally, here forming weakly impressed, transverse meshes. Clypeus and anterior half of frons shining, doubly punctate, without reticulation and with very close, fine and very sparse, coarse punctures. Coarse punctures approximately 5–8x diameter of fine; without visible reticulation. Paired epicranial foveae, one immediately behind the other, on each side of frons, close to lateral margins and immediately behind lateral remnants of frontoclypeal suture. Anterior epicranial foveae transverse, posterior slightly elongate oval; both with cluster of stout, yellow recumbent to decumbent setae. Areas between anterior and posterior foveae with coarse wrinkles. Posterior frons with open, elongate, wrinkled reticulation, especially alongside lateral margins of compound eyes and onto vertex; meshes tumid, with rugose appearance. Internal and posterior borders of compound eyes distinct, raised relative to level of adjacent cuticle. Lateral margins bordered by distinct narrow channel; deeper anteriorly than posteriorly and continuing behind posterior margin of eye onto vertex. Channel with dense punctures, bearing long, stiff, yellow recumbent to decumbent setae.

Pronotum. Posterior margin strongly sinuate laterally (Figure 3A). Surface somewhat shining, strongly rugose. Reticulation meshes large, open, almost isodiametric and relatively flat either side of mid-line on disc; smaller, tumid and more uneven in size and shape towards all margins. Transverse irregular row of medium punctures bearing long, yellow recumbent to decumbent setae 1/5 behind anterior pronotal margin; interrupted briefly in centre, continuing inside lateral margins and inside lateral third of posterior margin. Centre of disc with elongate, narrow, slit-like fovea, sometimes partially interrupted in mid-length. Lateral margins slightly raised, shining, without rugose sculpture and with fine, scattered punctures.

Elytra. Somewhat shining, with dense, transverse, crescentic striolae (Figures 5A, 6A) giving a scaly appearance (Figure 3A). Striolae less dense anteriorly and medially, here mostly distinctly separated from each other laterally (e.g. Figures 5A, 7A, C, 8A, C, 9, 10). Size of crescentic striolae variable, especially on disc close to suture (e.g. Figures 9, 10). Crescentic striolae becoming denser and somewhat continuous laterally and posteriorly (Figure 3A). Surface between crescentic striolae (Figures 5A, 6A) doubly punctate, with very fine, close punctures and medium, very sparse punctures (the latter bearing short, peg-like setae); also with fine, obsolete, open reticulation, usually more evident in apical two thirds, and more evident in some specimens than others; sometimes apparently absent. Puncture rows well-marked, continuous almost to elytral apices; punctures shallower posteriorly than anteriorly.

Figure 5. 

Meladema species males, elytral shoulder sculpture SEMs (DNA voucher codes, where applicable). AM. coricaea, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., Corsica, Cap Corse (NHM-IRM12F) C M. imbricata, La Gomera, El Cedro (NHM-IRM3A) D M. lanio, Madeira, Ribeira dos Cedros (NHM-IRM8A).

Figure 6. 

Details of elytral shoulder sculpture SEMs (DNA voucher codes where applicable). A M. coriacea, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., Corsica, Cap Corse (NHM-IRM12F).

Figure 7. 

Meladema species elytral shoulder sculpture SEMs (DNA voucher codes where applicable). A M. coriacea, male, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., male, Corsica, Cap Corse (NHM-IRM12F) C M. coriacea, female, Spain, Murcia, Fte. Caputa D M. lepidoptera sp. n., female, Corsica, Porto-Vecchio (NHM-IRM12A).

Venter. Prementum shining, tumid in centre, with fine and medium, sparse punctures. Mentum shining; central projection with shallow median emargination. Lateral lobes with medium, sparse punctures, and scattered, whitish recumbent to decumbent setae. Submentum shining, with transverse wrinkles centrally, and elongate wrinkles laterally. Central 1/4 with medium, sparse punctures bearing long, white-yellowish, erect setae. Gula shining, with sparse, shallow, transverse wrinkles; patch of medium-coarse punctures posterolaterally. Genae shining, with obsolete, open, elongate reticulation. Prosternum shining, with irregular transverse ridges laterally. Strongly arched in centre and with fine, moderate to close punctures laterally, bearing long, white-yellowish, recumbent to erect setae; punctures and setae extending in a sparse, irregular row onto process, just below arch. Process lanceolate, tectiform; apex acuminately rounded. Centre of prosternum and process with double punctation of very fine, moderate and medium, very sparse punctures. Pronotal hypomeron shining, impunctate. Elytral epipleurs shining, with fine wrinkles; irregular puncture row close to internal margin, from centre to close to apex; punctures bearing fine, whitish, erect setae. Metaventrite shining, central portion with sparse, transverse scratches and fine to very fine, sparse to very sparse punctures; not clearly forming two size classes. Metaventral process strongly reticulate, with transverse, rugose meshes and traces of fine, sparse punctures; small, central patch at base with very small reticulation meshes. Metaventral process relatively broad; apex acuminate and upturned slightly anterolaterally. Metacoxal lines almost reaching anterior border of metacoxae; shallow and interrupted in anterior 1/5. Internal laminae of metacoxae shining, sculpture as on centre of metaventrite. Metacoxal lobes sculptured as internal laminae, strongly rounded, with irregular, elongate field of medium to coarse punctures close to lateral margins, bearing fine, white, recumbent to erect setae. External laminae of metacoxae shining, smooth close to process, but with strong reticulation elsewhere; reticulation meshes very elongate posteromedially, to transverse anteriorly. Abdominal ventrites shining. Ventrites 3–5 with cluster of golden, erect setae anteromedially. Ventrite 1 with elongate reticulation throughout. Ventrite 2 with similar reticulation; absent close to centre. Ventrite 3 with elongate reticulation laterally, becoming transverse close to smooth central 1/5. Reticulation of ventrites 4 and 5 restricted to lateral third and with superimposed elongate furrows. Ventrites 2–5 doubly punctate; very fine, moderate and fine, sparse to very sparse punctures; punctures most evident in areas without reticulation. Ventrites 3–5 with transverse irregular row of long, yellowish, recumbent to decumbent setae laterally. Ventrite 6 (Figure 11A) with very fine, moderate punctures and medium to coarse, sparse to moderate punctures; punctures coarser close to apex. Elongate, semicircular wrinkles and channels apicolaterally. Some punctures in channels bearing elongate, whitish, erect setae.

Male. Foretarsi (Figure 12A) with articulo-setal counts as follows (base to apex): row 1, 7; row 2, 8–10; row 3, 8–10; row 4, 6-8. Number of setae in rows may differ from right to left tarsus in same beetle. Curved, golden setae bordering articulo-setal field dense, particularly basally. Foretarsal claws (Figures 12A, 13A) elongate, curved; interior margin straight or somewhat raised in basal two thirds, strongly curved in apical third. Mesotarsi (Figure 12E) with articulo-setal counts as follows (base to apex): row 1, 7–8; row 2, 8; row 3, 7; row 4, 4 (2 clusters of 2, situated laterally). Curved, golden setae bordering articulo-setal field dense, particularly basally. Mesotarsal claws (Figure 12E) elongate, strongly curved. Abdominal ventrite 6 (Figure 11A) with apex rounded, with very shallow median emargination. Median lobe asymmetrical (Figure 14A), sinuation weak, approximately 1/5–1/4 from apex; ventral margin of apical portion relatively straight in lateral view. Parameres (Figure 14A) with basal portion of inner margin relatively evenly curved; outer and inner margins almost straight over apical two thirds.

Female. As male, except for simple fore and mesotarsi, differently shaped abdominal ventrite 6 (with bluntly pointed apex, Figure 15A). The density of crescentic striolae on the elytra does not differ consistently between sexes, contrary to the statement of Balfour-Browne (1948).

Variation. Variation is evident in a number of characters. The size and density of the crescentic striolae on the elytra differs between individuals (e.g. Figures 5A, 7A, C, 8A, C, 9, 10, 17, 18D, 19), and shows some geographical structure. Specimens of both sexes from the Iberian Peninsula (e.g. Figures 7A, C, 8A, C, 9B, 10B) have relatively sparse crescentic striolae, with individual striolae relatively small in size, features also seen in most specimens from the Balkans and Anatolia (e.g. Figure 10A). Some specimens from France, North Africa and southern Italy (e.g. Figures 9D, 10C, D, 19) have slightly denser striolae, with some of these slightly larger. Specimens from massifs in the central Sahara (Hoggar and Tassili n’Ajjer (Algeria), Tibesti (Chad)) have very dense crescentic striolae (Figure 17). In these beetles the striolae are relatively small and strongly curved, giving a distinctly tuberculate appearance to the elytra, even close to the shoulders. In no cases do the elytral striolae approach the condition seen in M. lepidoptera sp. n., however (see below). The degree of curvature of male foretarsal claws differs between individuals, even within the same population. Some specimens have claws which are shorter, and broader at their apices that in Figure 13A, these almost certainly being older insects, whose claws have become worn down during life. The size and shape of the median lobe also varies slightly between individuals and populations (e.g. Figure 20). Specimens from North Africa, for example, (Figure 20H–K) have narrower apical portions, whilst the largest median lobes are seen in Greek animals (Figure 20M–N).

Morphologically, this species is almost identical to M. lepidoptera sp. n., something which has prevented the latter’s formal description until now. The two species can be reliably separated only on details of their elytral sculpture, M. coriacea having smaller, less dense crescentic striolae than M. lepidoptera sp. n., this being particularly evident at the elytral base, close to the scutellum, and in the middle, close to the suture (see Figure 1 and e.g. Figures 7A, C, 8A, C, 9, 10 vs. 7B, D, 8B, D, 21, 22). In M. coriacea, the striolae are largely well separated from each other in both regions (see comment by Aubé 1836, quoted above) whereas in M. lepidoptera sp. n. they are typically much more contiguous (see below). In the mid-elytra of M. coriacea there is also typically a distinct gap between the elytral suture and the first crescentic striolae (see e.g. Figure 8A, C), whereas in M. lepidoptera sp. n. the striolae begin much closer to the suture (e.g. Figure 8B, D). Whilst there is some variation, as illustrated in the figures, these features appear consistent throughout the wide geographical range of M. coriacea. Other characters, including male genitalia, claws, and the shape and sculpture of abdominal ventrite 6 in both sexes, do not differ consistently between the two species, there being as much intraspecific as interspecific variation.

The genetic differences between M. coriacea and M. lepidoptera sp. n. are well defined and comparable to those seen between M. imbricata and M. lanio, although mostly seen in mitochondrial markers (Sýkora et al. 2017). Of the two nuclear markers sequenced, H3 was identical in M. coriacea and M. lepidoptera sp. n. (but different from M. lanio and M. imbricata), only wingless having some diagnostic positions separating the two species (see Sýkora et al. 2017).

Even as redefined here, this is by far the most widespread species of the genus, distributed from the Canary Islands to Turkey, and south to massifs of the central Sahara (Figure 23). We have examined material from all countries from which this species has been reported, with the exception of Bulgaria (Guéorguiev 1987). Despite being listed from Lybia in the Palaearctic and World Catalogues of Dytiscidae (Nilsson and Hájek 2017a, 2017b), all Saharan records which we have traced emanate from Algeria and Chad. As discussed by Sýkora et al. (2017), basal splits within the M. coriacea lineage apparently took place in north-west Africa, with subsequent range expansion across the western Mediterranean, and several apparently independent, recent, colonisations of the Canary Islands from Morocco. The most easterly specimens included in genetic analyses (from Malta and Izmir, Turkey) were more closely related to beetles from north Africa than those from Iberia or southern France, suggesting that colonisation of the eastern part of the species range occurred via dispersal from Algeria/Tunsia, likely through Malta, Sicily and southern Italy. Genotyped individuals have been studied from throughout the wide range of M. coriacea, as well as a large number of additional non-genotyped specimens (Figure 23). The exact limits of M. coriacea in the Italian Peninsula remain unclear, and can only be resolved through combined genetic and morphological study of additional, fresh material. It is apparent, however, that M. coriacea is present in at least Sicily and that M. lepidoptera sp. n. occurs in the north and centre of the peninsula (Toscana and Lazio – as confirmed from genotyped specimens). Beetles morphologically intermediate between M. coriacea and M. lepidoptera sp. n. have been seen from Apulia and Campania (Figure 18B, C), suggesting possible hybridization between the two species where their ranges come into contact. Such a process is already established to have occurred between M. coriacea and M. imbricata on Tenerife (see below and Ribera et al. 2003). Meladema coriacea is a species typical of seasonally fluctuating or intermittent Mediterranean stream systems, being particularly characteristic of localities with winter-spring flow, which dry down to pools in summer. It is associated with drier and more seasonally variable conditions than other species of the genus, something which may have facilitated its range expansion (Sýkora et al. 2017).

France, Corsica, Cap Corse, stream nr. Bettolacce, 42°58'2.4"N 9°24'42.4"E.

(genotyped specimens only). Holotype ♂: “11 FR Corsica 21.ix.1999// Cap Corse: Bettolacce// 42°58'2.4"N 9°24'42.4"E 250m// I.Ribera & A. Cieslak leg.” “DNA voucher// NHM-IRM12E” “Meladema lepidoptera// Bilton & Ribera, 2017//HOLOTYPE” (NMW). Dry card mounted, tissue samples and DNA aliquotes, with same data, in IBE. Sequence data from the holotype has been deposited in GenBank with accession numbers AF428206 (COI-3’) and AF428188 (16S ribosomal RNA). Paratypes (13): 1 ♂ “11 FR Corsica 21.ix.1999// Cap Corse: Bettolacce// 42°58'2.4"N 9°24'42.4"E 250m// I.Ribera & A. Cieslak leg.”“DNA voucher// NHM-IRM12F” (NMW); 1 ♀ “11 FR Corsica 21.ix.1999// Cap Corse: Bettolacce// 42°58'2.4"N 9°24'42.4"E 250m// I.Ribera & A. Cieslak leg.” “DNA voucher// NHM-IRM12C” (BMNH); 1 ♀ “5 FR Corsica 19.ix.1999// Porto-Vecchio, l’Ospedale// 41°39'13.7"N 9°12'41.0"E 690m// I.Ribera & A. Cieslak leg.” “DNA voucher// NHM-IRM12A” (CBP); 1 ♀ “5 FR Corsica 19.ix.1999// Porto-Vecchio, l’Ospedale// 41°39'13.7"N 9°12'41.0"E 690m// I.Ribera & A. Cieslak leg.” “DNA voucher// NHM-IRM12D” (MNHN); 1 ♀ “5 FR Corsica 19.ix.1999// Porto-Vecchio, l’Ospedale// 41°39'13.7"N 9°12'41.0"E 690m// I.Ribera & A. Cieslak leg.” “DNA Voucher// NHM-IRM12g” (IBE); 1 ♀ “9 FR Corsica 19.ix.1999// Ghisoni, road to Campannella// 42°4'8.7"N 9°11'6.0"E 830m// I.Ribera & A. Cieslak leg.” “DNA Voucher// NHM-IRM12b” (IBE); 1 ♂ “Toscana (PI) S. Luce// ‘Boso’ de Castagni, s tr. s.// Luce-Castellina Marittima// 3.X.2007 leg. M. Toledo” “DNA voucher// IBE-AN693” (CTP); 1 ♀ “April 2015 Italy Monti della Tolfa// Rio Ippovia della Cicugnola// (pozze ruscellamento)// 42°4'26.27"N 11°56'12.25"E// V. Buono leg.” “DNA voucher// IBE-AN760” (CVR); 1 ♂ “I. D’ELBA – POMONTE// Fosso BARIONE// m 250-300// 29.V.94 TOLEDO LGT.” [HW] “DNA voucher// IBE-AN692” (CTP); 1 ♂ “3/vi/2014 ITALY Montecristo// 42.334N 10.308E 260m// R. Vila leg.” “DNA voucher// IBE-DV289” (IBE); 1 ♂ “3/vi/2014 ITALY Montecristo// 42.334N 10.308E 260m// R. Vila leg.” “DNA voucher// IBE-DV290” (ISNB); 1 ♂ “27/v/2009 ITALY Sardinia// Nuoro prov.(Ogliastra historical region)// brook 562m ESE Villagrande Strìsaili (WNW Tortoli)// 39.95084N 9.51912E H. Fery & M. Toledo leg.” “DNA voucher// IBE-RA5” (MNCN); 1 ♀ “27.5.2009 Italy, Sardinia// Ogliastra prov., ca. 5 km// WNW Tortoli, (on road// Tortoli – Villagrande Stris.)” “39.93982N 9.59280E// ca 80m, brook// Fery & Toledo leg.” “DNA voucher// IBE-RA18” (MNCN). Each with red label “Meladema lepidoptera// Bilton & Ribera, 2017//PARATYPE”. All dry card mounted, tissue samples and DNA aliquotes, with same data, in IBE.

(non-genotyped specimens). France, Corsica: 2 ♂♂, 1 ♀ “11/iv/1993// Corsica Francardo// Mediterranean stream// D. T. Bilton leg.” (CBP); 1 ♀ “Calvi// 29.VIII.” “Pietra// Maggiore” “KORSIKA// VIII.1955” “Meladema// coriacea Cast.// M/ Balke det. 1990” [Latin name, describer & 90 HW] (NMW); 1 ♂, 2 ♀♀ “12.4.79 Korsika// Pinito, ca 700m// Bach” [HW] reverse “Fery leg.” “Mel.// coriacea// Cast.” [HW] (NMW); 1 ♀ “12.4.79 Korsika// Pinito, ca 700m// Bach” [HW] reverse “Fery leg.” “Mel.// coriacea// Cast.” [HW] “coll. Shaverdo” (NMW); 1 ♀ “Korsika// 7.80” [HW] “Meladema// coriacea Cast.// M. Balke 1990” [Latin name, describer & 90 HW] (NMW); 1 ♂ “Corse” [HW] “Reveliere” [yellow, square label, HW] “Coriaceus// Corsica” [HW] “C. Epplsh.// Steind. d.” (NMW); 1 ♂ “Corsica” (NMW); 1 ♀ “12.4.79 Korsika// Pinito, ca. 700m// Bach” [HW] “Mel.// coriacea// Cast.” [HW] “Coll.// HENDRICH// Berlin” (ZSM). Italy, Sardinia: 5 ♂♂, 3 ♀♀ “Sardinia (CA) Dolianova// Rio Flumini 30.VI.91// leg. Meloni” (CBP, CTP); 3 ♂♂, 1 ♀ “SARDINIEN// Bosa// 24.5-24.6.//1963, Budberg” reverse “Meladema// coriaceum Lap.” [HW] (NMW); 1 ♀ “coll. Win-// gelmüller” “coriaceum Lap.// Sardinien” [HW] (NMW); 1 ♂ “Villasimius// Sard. m. 19.9.59// E. Jünger” [19 HW] “A7598// coricae Cast.// det. K. Hoch 1959” [number, Latin name, describer & 9 HW] (ZSM); 1 ♀ “Sardegna// Lode// 28.7.79// S. Gottwalt” [HW] “coll.//HENDRICH// Berlin” (ZSM); 2 ♀♀ “Sardinien: bei Nuvafus,// 09.IX.1978” “Meladema// coriacea// Cast.” [HW] “entnommen// 1 Exempl.” [HW] “Sardinien// S. Nuvafus// 9.9.78” [HW] (ZSM); 1 ♂, 1 ♀ “Sardinien: S Monte// Limbara. Tempio, 650m,// 28.VIII.1978” “Meladema// coriacea// Cast.” [HW] “Entnommen// 1 Exempl.” [HW] “Sardinien// Südl. Monte. Liombara// Tempio 650m// 28.8.78” [HW] “Fuss Monte// Limbara 650m// 28.8.78” [HW] (ZSM); 1 ♀ “Sardinien: Lagoatto di// Flumendosa, Villanova// Strisaili, 05.IX.1978” “Meladema// coriacea// Cast.” [HW] “Sardinien// Lagoatto di Flumendosa// Villanova Strisaili// 5.9.1978” [HW] (ZSM); 6 ♂♂, 3 ♀♀ “Meladema// coriacea Cast.// Italien/ Sardinien// Giara di Gesturi// - 9.1980// Coll./ leg.// Burmeister” “Sardinien// Giara di Gesturi// 9.1980// leg. Burmeister” [HW] “Meladema// coriacea// Cast.” [HW] (ZSM); 1 ♂ “Meladema// coriacea Cast.// Italien/ Sardinien// b. Tempio// Bergbach am// Monte Limbarra// - 9.1980// Coll./ leg.// Burmeister” “Sardinien// b. Tempio// Bergbach am// Monte Limbarra// 9. 1980” [HW] “Meladema// coriacea// Cast.” [HW] (ZSM); 2 ♂♂, 2 ♀♀ “Meladema// coriacea Cast.// Italien/ Sardinien// Pass b. Genne// Cruxi// 1.9.1980// Coll./ leg.// Burmeister” “Sardinien// Pass b. Genne Cruxi// 10. [circled]// leg. Burmeister” [HW] (ZSM); 2 ♀♀ “Meladema// coriacea Cast.// Italien/ Sardinien// b. Nuragus// (Viehtränke)// 3.9.1980// Coll./ leg.// Burmeister” “Sardinien// b. Nuragus// Viehtränke// 3.9.1980 23. [circled]// leg. Burmeister” [HW] “Meladema// coriacea// Cast.” [HW] (ZSM); 1 ♂ “Meladema// coriacea Cast.// Italien/ Sardinien// b. Tempio// Bergbach// b. Mnte Limbarra// - 9.1980// Coll./ leg.// Burmeister” “Sardinien// b. Tempio, Bergbach// b. Monte Limbarra// 9/1980 39. [circled]// leg. Burmeister” [HW] ‘Meladema// coriacea// Cast.” [HW] (ZSM); 1 ♂ “Glasgow University// HUNTERIAN MUSEUM// ex. coll. G.N. Foster” “Sardinia 32T UTH// 04602/44750// S of Mont Minerva// G N Foster 16.10.2006” (CFA); 1 ♀ “27.5.2009 Italy, Sardinia// Ogliastra prov., ca. 5 km// WNW Tortoli, (on road// Tortoli – Villagrande Stris.)” “39.93982N 9.59280E// ca 80m, brook// Fery & Toledo leg.” (IBE). 1 ♂ “9 Sardinia, Calangianus 11.iv.2017// stream, way to Pascaredda tomb// 40°54'35.6"N 9°10'12.8"E 435m// I.Ribera & A.Cieslak leg.” (IBE); Italy, Elba: 1 ♂ [no head] “Ins. Elba// 1908// Paganetti” (NMW); 2 ♂♂ “Elba: bei Marciana, Bach in// Kastanienwald, 17.IX.1975,// leg.: Schmalfuss” “17.9.75 Elba, bei Murciana// Bach in Kastanienwald// Schmalfuss leg.” [HW] “Meladema// coriaceum// Lap.” [HW] “Meladema coriacea Cast.” [HW] (ZSM). Italy, mainland: 5 ♂♂, 3 ♀♀ “Levante,// Liguria, Italy.” “Coll. Odier.// B.M. 1921-288.” (BMNH); 1 ♂, 1 ♀ “Levante,// Liguria, Italy.” “Coll. Odier.// B.M. 1921-288.” “Meladema// coriacea Cast// C.R. Smith det. 1982” [Latin name, describer & 2 HW] (BMNH); 1 ♀ “ITALY: Liguria.//Torr. 2km W. of Pogli,// trib. of Torr. Arroscia, nr. Albenga, 7.iv.1958// J. Balfour-Browne.” “Brit. Mus.// 1960-482.” [482 HW] “Meladema// coriacea Cast// C.R. Smith det. 1982” [Latin name, describer & 2 HW] (BMNH); 3 ♂♂, 1 ♀ “Italia// Ruta// 2-6. VII.95// A. Fiori” [HW, A. Fiori printed] (NMW); 1 ♀ “coriaceus// Toscana// V. Heyden” [HW] “c. Epplsh.// Steind. d.” (NMW); 1 ♂ “S. gimignano// Sienna → Florenz// 3.8.36 Eiselt” [HW] “Meladema// coriaceum” [HW] (NMW); 1 ♂ “San Gimignano// IX 36 EISELT” “Meladema// coriaceum Lap.” [HW] (NMW); 1 ♀ “San Gimignano// IX 36 EISELT” (NMW); 1 ♂ “Nervi// 27.III.05” [HW] (ZSM); 2 ♀♀ “Italia// 26.3.05” [date HW] “Nervi” (ZSM); 1 ♀ “Italien// 11.xii.05” [HW] “♀” “Samml. A.// Zimmermann” (ZSM); 1 ♀ [small, circular brown label, no text] “Pisa” [HW] “alte// Sammlung” (ZSM); 1 ♀ “Italia: Umbria, NE Gosparini,// 550m, dry stream, 09.IX.2015// 43°14'31.73"N, 12°6'19.69"E,// leg. Komarek, Beutel (TM7)” “Meladema// coriacea Lap.// det. H. Shaverdo 2016” [Latin name, describer & 16 HW] (NMW). Without locality data: 1 ♀ [small, circular brown label, no text] “Coll. Odier.// B.M. 1921-288.” (BMNH); 1 ♀ “coriaceus” [HW] “E Coll.// Curtis” [HW] (BMNH); 1 ♂ “40// 4 2// 2135” [round label, HW] (BMNH). All with “Meladema lepidoptera// Bilton & Ribera// D T Bilton [or I Ribera] det. 2017”.

Size: Holotype TL = 20.74 mm; EL = 15.74 mm; MW = 10.50 mm. Other material examined TL = 19.20–20.99 mm; EL = 14.98–16.38 mm; MW = 9.73–11.39 mm.

Colour. Dorsum dark reddish brown to black (Figure 3B); lateral margins of pronotum, labrum and anterior half of clypeus somewhat paler, sometimes with diffuse lateral maculae. Elytra unicolorous, without distinct mottling even when lifted (Figure 4B). Head with a pair of oval, reddish yellow medial interocular patches, slightly elongated apicolaterally. Antennae and maxillary and labial palpi reddish yellow. Legs dark reddish brown to black with golden yellow setae; large spines somewhat paler. Venter reddish brown to black; gula, meso and metacoxae and trochanters paler.

Head. Labrum shining, with moderate to coarse, sparse punctures. Reticulation absent in apical half, becoming increasingly more evident basally, here forming weakly impressed, transverse meshes. Clypeus and anterior half of frons shining, doubly punctate, without reticulation and with very close, fine and very sparse, coarse punctures. Coarse punctures approximately 5–8x diameter of fine; without visible reticulation. Paired epicranial foveae, one immediately behind the other, on each side of frons, close to lateral margins and immediately behind lateral remnants of frontoclypeal suture. Anterior epicranial foveae transverse, posterior slightly elongate oval; both with cluster of stout, yellow recumbent to decumbent setae. Areas between anterior and posterior foveae with coarse wrinkles. Posterior frons with open, elongate, wrinkled reticulation, especially alongside lateral margins of compound eyes and onto vertex; meshes tumid, with rugose appearance. Internal and posterior borders of compound eyes distinct, raised relative to level of adjacent cuticle. Lateral margins bordered by distinct narrow channel; deeper anteriorly than posteriorly and continuing behind posterior margin of eye onto vertex. Channel with dense punctures, bearing long, stiff, yellow recumbent to decumbent setae.

Pronotum. Posterior margin strongly sinuate laterally (Figure 3B). Surface somewhat shining, strongly rugose. Reticulation meshes large, open, almost isodiametric and relatively flat either side of mid-line on disc; smaller, tumid and more uneven in size and shape towards all margins. Transverse irregular row of medium punctures bearing long, yellow recumbent to decumbent setae 1/5 behind anterior pronotal margin; interrupted briefly in centre, continuing inside lateral margins and inside lateral third of posterior margin. Centre of disc with elongate, narrow, slit-like fovea, sometimes partially interrupted in mid-length. Lateral margins slightly raised, shining, without rugose sculpture and with fine, scattered punctures.

Elytra. Somewhat shining, with dense, transverse, sometimes contiguous, crescentic striolae, giving a very scaly appearance (Figure 3B). Striolae relatively dense throughout, frequently contacting each other laterally on shoulder close to suture (e.g. Figures 5B, 7B, 18A, 21, 22, and relatively dense in mid-elytra close to suture (e.g. Figures 7D, 8D, 18A, 21, 22)). Size of crescentic striolae relatively large, both in shoulder and mid-elytral regions (e.g. Figures 7B, D, 8B, D, 18A, 21, 22). Crescentic striolae becoming denser and somewhat continuous laterally and posteriorly. Surface between crescentic striolae (Figures 5B, 6B) doubly punctate, with very fine, close punctures and medium, very sparse punctures (the latter bearing short, peg-like setae); also with fine, obsolete, open reticulation, usually more evident in apical two thirds, and more evident in some specimens than others. Puncture rows well-marked, continuous almost to elytral apices; punctures shallower posteriorly than anteriorly.

Venter. Prementum shining, tumid in centre, with fine and medium, sparse punctures. Mentum shining; central projection with shallow median emargination. Lateral lobes with medium, sparse punctures, and scattered, whitish recumbent to decumbent setae. Submentum shining, with transverse wrinkles centrally, and elongate wrinkles laterally. Central 1/4 with medium, sparse punctures bearing long, white-yellowish, erect setae. Gula shining, with sparse, shallow, transverse wrinkles; patch of medium-coarse punctures posterolaterally. Genae shining, with obsolete, open, elongate reticulation. Prosternum shining, with irregular transverse ridges laterally. Strongly arched in centre and with fine, moderate to close punctures laterally, bearing long, white-yellowish, recumbent to erect setae; punctures and setae extending in a sparse, irregular row onto process, just below arch. Process lanceolate, tectiform; apex acuminately rounded. Centre of prosternum and process with double punctation of very fine, moderate and medium, very sparse punctures. Pronotal hypomeron shining, impunctate. Elytral epipleurs shining, with fine wrinkles; irregular puncture row close to internal margin, from centre to close to apex; punctures bearing fine, whitish, erect setae. Metaventrite shining, central portion with sparse, transverse scratches and fine to very fine, sparse to very sparse punctures; not clearly forming two size classes. Metaventral process strongly reticulate, with transverse, rugose meshes and traces of fine, sparse punctures; with small central patch of reticulation with very small meshes. Metaventral process relatively broad; apex acuminate and upturned slightly anterolaterally. Metacoxal lines almost reaching anterior border of metacoxae; shallow and interrupted in anterior 1/5. Internal laminae of metacoxae shining, sculpture as on centre of metaventrite. Metacoxal lobes sculptured as internal laminae, strongly rounded, with irregular, elongate field of medium to coarse punctures close to lateral margins, bearing fine, white, recumbent to erect setae. External laminae of metacoxae shining, smooth close to process, but with strong reticulation elsewhere; reticulation meshes very elongate posteromedially, to transverse anteriorly. Abdominal ventrites shining. Ventrites 3–5 with cluster of golden, erect setae anteromedially. Ventrite 1 with elongate reticulation throughout. Ventrite 2 with similar reticulation; absent close to centre. Ventrite 3 with elongate reticulation laterally, becoming transverse close to smooth central 1/5. Reticulation of ventrites 4 and 5 restricted to lateral third and with superimposed elongate furrows. Ventrites 2–5 doubly punctate; very fine, moderate and fine, sparse to very sparse punctures; punctures most evident in areas without reticulation. Ventrites 3–5 with transverse irregular row of long, yellowish, recumbent to decumbent setae laterally. Ventrite 6 (Figure 11B) with very fine, moderate punctures and medium to coarse, sparse to moderate punctures; punctures coarser close to apex. Elongate, semicircular wrinkles and channels apicolaterally. Some punctures in channels bearing elongate, whitish, erect setae.

Male. Foretarsi (Figure 12B) with articulo-setal counts as follows (base to apex): row 1, 7; row 2, 8–10; row 3, 8–10; row 4, 6–8. Number of setae in rows may differ from right to left tarsus in same beetle. Curved, golden setae bordering articulo-setal field dense, particularly basally. Foretarsal claws (Figures 12B, 13B) elongate, curved; interior margin straight or somewhat raised in basal two thirds, strongly curved in apical third. Mesotarsi (Figure 12F) with articulo-setal counts as follows (base to apex): row 1, 7–8; row 2, 8; row 3, 7; row 4, 4 (2 clusters of 2, situated laterally). Curved, golden setae bordering articulo-setal field dense, particularly basally. Mesotarsal claws (Figure 12F) elongate, strongly curved. Abdominal ventrite 6 (Figure 11B) with apex rounded, with very shallow median emargination. Median lobe asymmetrical (Figure 14B), sinuation weak, approximately 1/5–1/4 from apex; ventral margin of apical portion relatively straight in lateral view. Parameres (Figure 14B) with basal portion of inner margin relatively evenly curved; outer and inner margins almost straight over apical two thirds.

Female. As male, except for simple fore and mesotarsi, differently shaped abdominal ventrite 6 (with bluntly pointed apex - Figure 15B). As with M. coriacea, no consistent differences between males and females are evident in terms of elytral sculpture.

Variation. The size and density of the crescentic striolae on the elytra differs somewhat between individuals and localities (Figures 5B, 7B, D, 8B, D, 18A, 21, 22), these being relatively dense in mainland Italy, Corsica and the Tuscan archipelago, and less so in most Sardinian specimens (e.g. Figure 22). The combination of size and density of these striolae is always greater than seen in M. coriacea, however (see above). The degree of curvature of male foretarsal claws differs between individuals, as in M. coriacea, as does the size and shape of the median lobe (e.g. Figure 20O–U).

Morphologically almost identical to M. coriacea (see above). Only distinguishable on the size, shape and density of crescentic striolae on the elytra, which give M. lepidoptera sp. n. a very scaly appearance, evident even at relatively low magnification (e.g. Figure 3A vs. 3B). See above for genetic differences between this species and M. coriacea.

From the ancient Greek “lepidos” (λεπίδος, scale, but also referring to roof tiles) and “pteron” (πτερόν, wing). The specific epithet is a noun in the nominative plural.

On the basis of current data, found on Corsica and Sardinia, islands of the Tuscan Archipelago (Elba, Montecristo) and parts of peninsular Italy, from Liguria to Umbria (Figure 23). M. lepidoptera sp. n. is apparently the only species of the genus found on Corsica, Sardinia, Elba and Montecristo (past records of M. coriacea from these islands - e.g. Poggi 1976, Franciscolo 1979, Dettner 2007 - almost certainly referring to this species), but co-occurs with M. coriacea in the Italian peninsula. The exact limits of the distribution of the two species in peninsular Italy remain unclear (see above), but there is morphological evidence suggesting hybridization where they meet, at least in the south (see below). The contact zone between the two species in the north appears to be situated on the Mediterranean coast, somewhere close to the French-Italian border, but to date no intermediate specimens have been seen from this area. Clearly future work, using both genetic and morphological approaches, would be illuminating in understanding the location and dynamics of these contact zones. As with other extant Meladema lineages, this species appears to have originated in the early Pleistocene, colonisation of the Tyrrhenean islands occurring long after the Messinian Salinity Crisis (Ribera et al. 2003, Sýkora et al. 2017). Sýkora et al. (2017) suggest that M. lepidoptera sp. n. may have originated following the colonization of the Tyrrhenian islands, a hypothesis which should be tested in the future through genetic study of more individuals from peninsular Italy. Sýkora et al. (2017) additionally suggest that this species is characteristic of sites with lower seasonality than is typical for M. coriacea, based on MaxEnt modelling. They note that many of the peninsular Italian localities (obtained from the literature) included in their analyses (from which specimens were not studied) fitted into the climatic space occupied by M. lepidoptera sp. n. (as ‘coriaceaCSM’), not surprising given our finding that this species does indeed occur on the Italian mainland. Clearly it would be interesting to repeat Sýkora et al.’s (2017) analyses in the future, once the range limits of these taxa are better established. On the basis of current evidence, this species occurs in similar habitats to those occupied by M. coriacea, although the two taxa have not been detected to date in the same locality.

Figure 8. 

Meladema species elytral middle sculpture SEMs (DNA voucher codes where applicable). A M. coriacea, male, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., male, Corsica, Cap Corse (NHM-IRM12F) C M. coriacea, female, Spain, Murcia, Fte. Caputa D M. lepidoptera sp. n., female, Corsica, Porto-Vecchio (NHM-IRM12A).

Figure 9. 

Meladema coriacea female elytral sculpture; shoulder and middle left and right, respectively (DNA voucher codes). A neotype, France, Var, La-Londe-les-Maures (NHM-IRM11C) B Spain, Córdoba, Baena (NHM-IRM14B) C Morocco, Tazzeka (NHM-IRM1A) D Algeria, Oued Bagrat (MNCN-HI4). Scale bar = 0.5 mm.

Figure 10. 

Meladema coriacea elytral sculpture; shoulder and middle left and right, respectively (DNA voucher codes). A Female, Turkey, Izmir (IBE-DV294) B male, Spain, Córdoba, Baena (NHM-IRM14A) C male, Morocco, Oued Massa (NHM-IRM2A) D male, Algeria, Aïn Damous (MNCN-HI6). Scale bar = 0.5 mm.

“Madeira” [mislabelled].

(BMNH). Holotype ♀ (Figure 24A): “Scutopterus// imbricatus, Woll” [HW] Scutopterus// imbricatus Woll// M.E.Bacchus det 1977// HOLOTYPE” [Latin name, describer & last 7 HW] “Holo-// type” [small, circular label, red margin] “Meladema// imbricata (Woll.)// M. Balke det. 1989” [Latin name, describer & 89 HW] (dry pinned, BMNH, Wollaston Collection).

Note that as discussed by Machado (1987), the type specimen must have been mislabelled, as this species is now known to be endemic to the Canary Islands.

(genotyped specimens). Spain, Canary Islands. 1 ♂ “1998 SPAIN Islas Canarias// La Gomera// El Cedro – stream in laurysilva// D. T. Bilton leg.” “M. IMBRICATA” [HW] “G4 Mel// below G3” [HW] “DNA voucher// NHM-IRM3A” (IBE); 1 ♂ “1998 SPAIN Islas Canarias// La Gomera// El Cedro – stream in laurysilva// D. T. Bilton leg.” “M. IMBRICATA” [HW] “G1 Mel” [HW] “DNA voucher// NHM-IRM4A” (IBE); 1 ♂ “1998 SPAIN Islas Canarias// La Gomera// El Cedro – stream in laurysilva// D. T. Bilton leg.” “DNA voucher// NHM-IRM4b” (IBE); 1 ♀ "15/i/2000 SPAIN Islas Canarias// La Gomera// El Cedro// D. T. Bilton leg." "DNA voucher// NHM-IRM15A" (CBP); 1 ♂ "15/i/2000 SPAIN Islas Canarias// La Gomera// El Cedro// D. T. Bilton leg." "DNA voucher// NHM-IRM15B" (CBP); 1 ♂ “April 1999 SPAIN// Islas Canarias La Palma// Bco. Hoyo Verde Caldera de// Taburiente D. T. Bilton leg.” “DNA voucher// NHM-IRM6A” (CBP); 1 ♀ “April 1999 SPAIN// Islas Canarias La Palma// Bco. Hoyo Verde Caldera de// Taburiente D. T. Bilton leg.” “DNA voucher// NHM-IRM6B” (CBP); 1 ♀ “April 1999 SPAIN// Islas Canarias La Palma// Bco. Hoyo Verde Caldera de// Taburiente D. T. Bilton leg.” “DNA voucher// NHM-IRM6C” (CBP); 1 ♂ "April 1998 SPAIN// Islas Canarias La Palma// Bco. del Rio upper reaches in// Laurisylva D. T. Bilton leg." "DNA voucher// NHM-IRM7A" (CBP); 1 ♂ “1998 SPAIN Islas Canarias// Tenerife// Barranco del Río 1,600m// D. T. Bilton leg.” “M. IMBRICATA” [HW] “T7// Mel.” [HW] “DNA voucher// NMH-IRM5A” (IBE); 1 ♂ “1998 SPAIN Islas Canarias// Tenerife// Barranco del Río 1,600m// D. T. Bilton leg.” “DNA voucher// NMH-IRM5D” (IBE); 1 ♂ “13/i/2000 SPAIN Islas Canarias// Tenerife// Barranco del Rio 1,600m// D. T. Bilton leg.” "DNA voucher// NHM-IRM17A" (CBP); 1 ♀ “13/i/2000 SPAIN Islas Canarias// Tenerife// Barranco del Rio 1,600m// D. T. Bilton leg.” "DNA voucher// NHM-IRM17B" (CBP). All with “Meladema imbricata// (Wollaston, 1871)// D T Bilton [or I Ribera] det. 2017”.

Additional material examined (non-genotyped specimens). Spain, Canary Islands. 1 ♂ “April 1998 SPAIN// Islas Canarias la Gomera// El Cedro stream in Garajonay// laurisylva D. T. Bilton leg.” (CBP); 1 ♀ “April 1998 SPAIN// Islas Canarias La Palma// Bco. del Río upper reaches in// Laurisylva D. T. Bilton leg.” (CBP); 1 ♂ “Islas Canarias: Tene-// rife, 9.-10.vi. 1989// Bco. del Río 1100m.// Balke & Hendrich leg.” “Meladema// imbricata// M. Balke det 2011” “M. Balke// BMNH(E) 2013-119” (BMNH); 1 ♂ “I. Canarias/ Tenerife// Bco. del Rio, 1400m// 9.-10.6.1989, Bach// Balke/ Hendrich, leg.” “Meladema// imbricata Woll.// HENDRICH det. 1995” [Latin name & describer HW] (CBP); 2 ♂♂, 2 ♀♀ “9.-10.6.89 Islas Canarias// Tenerife, Barranco// del Rio, ca. 1000m// Balke, Hendrich, Fery leg.” [HW] “Meladema// imbricata// Woll.// Fery det. 89” [HW] (CBF); 2 ♂♂, 1 ♀ “ESP. Tenerife// Bco. del Río 1600m// 2.xi.1991 AN Nilsson” “Meladema imbricata// (Wollaston, 1871)// Det. AN Nilsson 1991” (CBP); 1 ♂ “April 1998 SPAIN// Islas Canarias Tenerife// Bco. del Río Upper// Reaches D. T. Bilton leg” (CBP). All with “Meladema// imbricata (Wollaston, 1871)// D T Bilton [or I Ribera] det. 2017”.

(based on all material examined).Size: Holotype TL = 22.13 mm; EL = 15.79 mm; MW = 10.22 mm. Other material examined TL = 18.05–21.38 mm; EL = 13.57–15.62 mm; MW = 8.83–9.98 mm.

Colour. Dorsum (Figure 3C), dark reddish brown to yellow. Labrum yellowish; clypeus yellowish except central 1/4 red to blackish, connected to dark pigmentation on frons. Frons with transverse pale strip anterolaterally, adjacent to pale parts of clypeus, otherwise dark reddish brown. Medial, paired interocular patches on frons yellow; strongly transverse apicolaterally, almost reaching channel around interior margin of compound eye. Pronotum dark reddish brown on disc; narrowly reddish along anterior margin; lateral margins broadly yellowish to pale red. Elytra yellowish brown, with black irrorations; pattern much more clearly visible when lifted (Figure 4C). Legs yellowish brown to black; posterior tibiae and tarsi darkest. Antennae and maxillary and labial palpi yellowish to reddish. Venter reddish brown; prementum and posterior genae yellow; mentum and submentum reddish. Pronotal hypomeron and shoulder, outer portion of elytral epipleurs and apex of metacoxal process yellowish.

Head. Labrum shining, with medium to fine, sparse punctures. Reticulation absent anteriorly, clearly evident in posterior half, here fine and composed of small, isodiametric to slightly transverse meshes. Clypeus weakly shining, with medium to fine, sparse punctures and traces of very fine, shallow, close punctures. Frons weakly shining, entire surface with coarse, open reticulation, becoming stronger and more evident posteriorly. Meshes transverse to isodiametric apically and medially, strongly elongate posteriorly and onto vertex. Paired epicranial foveae on anterior frons, one immediately behind the other. Anterior foveae transverse, posterior foveae elongate oval. Foveae all strongly reticulate; anterior and posterior foveae linked by reticulated channel. Internal and posterior borders of compound eyes distinct, raised relative to level of adjacent cuticle. Lateral margins bordered by distinct narrow channel; deeper anteriorly than posteriorly and continuing behind posterior margin of eye onto vertex. Channel with dense punctures, bearing long, stiff, yellow recumbent to decumbent setae.

Pronotum. Posterior margin weakly sinuate laterally (Figure 3C). Surface somewhat shining, strongly rugose. Reticulation meshes large, open, flat and with varying sizes and orientations. Transverse irregular row of medium punctures bearing long, yellow recumbent to decumbent setae 1/5 behind anterior pronotal margin; interrupted briefly in centre, obscured by reticulation inside lateral margins but continuing inside lateral third of posterior margin. Reticulation weak and obsolete anterior to transverse row, surface here clearly doubly punctate, with very fine, dense and medium, sparse to very sparse punctures. Scattered medium punctures visible elsewhere, amongst meshes of reticulation. Centre of disc with traces elongate, narrow, slit-like fovea, typically obscured by reticulation, but traceable as an elongate reticulation channel. Lateral margins slightly raised, shining, without rugose sculpture and with fine, scattered punctures.

Elytra. Shining, with short, transverse, usually straight or weakly curved crescentic striolae of varying sizes and density (Figures 5C, 25D, F). Striolae shallow and moderate on shoulder and anterior disc, widely separated; becoming closer, larger and more curved posteriorly. Posterior third of elytra with an almost scaly appearance (Figure 3C); striolae here almost touching each other laterally. Surface between crescentic striolae doubly punctate and reticulate (Figure 5C); with very fine, close and medium, sparse to very sparse punctures (the latter bearing short, peg-like setae). Reticulation fine, somewhat obsolete, meshes isodiametric; more evident in posterior half, sometimes obscuring very fine punctation. Puncture rows well-marked, continuous almost to elytral apices; punctures shallower posteriorly than anteriorly.

Venter. Prementum shining, tumid in centre, with fine and medium, sparse punctures. Mentum shining; central projection with shallow median emargination. Lateral lobes with very fine, close punctures, scattered, whitish recumbent to decumbent setae and longitudinal wrinkles. Submentum shining, with transverse wrinkles. Central 1/4 with medium, sparse punctures bearing long, white-yellowish, erect setae. Gula shining, with sparse, shallow, transverse wrinkles; patch of medium-coarse punctures posterolaterally. Genae shining, strongly reticulate; meshes transverse anteriorly and posteriorly, almost isodiametric in centre. Prosternum shining, with weak, low irregular transverse ridges laterally. Arched in centre and with fine, moderate to close punctures laterally, bearing long, white-yellowish, recumbent to erect setae; punctures and setae extending in an irregular row onto process, just below arch. Process lanceolate, arched; apex acuminately rounded. Centre of prosternum and process with double punctation of very fine, close to very close and medium, sparse to moderate punctures. Pronotal hypomeron shining, impunctate. Elytral epipleurs shining, with fine wrinkles; irregular puncture row close to internal margin, from centre to close to apex, punctures bearing fine, whitish, erect setae. Metaventrite shining, central portion with reticulation reduced to sparse, transverse scratches and very fine, close and fine to medium, sparse punctures. Metaventral process strongly reticulate, with transverse to elongate, rugose meshes and traces of fine, sparse punctures; small central area with reticulation of very small meshes. Metaventral process relatively broad; apex acuminate and upturned slightly anterolaterally. Metacoxal lines not reaching anterior border of metacoxae, disappearing approx 1/10 from margin. Internal laminae of metacoxae shining, sculpture as on centre of metaventrite. Metacoxal lobes sculptured as internal laminae, strongly rounded, with irregular, elongate field of medium to coarse punctures close to lateral margins, bearing fine, white, recumbent to erect setae. External laminae of metacoxae shining, smooth close to process, but with strong reticulation elsewhere; reticulation meshes very elongate posteromedially, to transverse anteriorly. Abdominal ventrites shining. Ventrites 3–5 with cluster of golden, erect setae anteromedially. Ventrite 1 with elongate reticulation throughout. Ventrite 2 with similar reticulation; absent close to centre. Ventrite 3 with elongate reticulation laterally, becoming transverse close to smooth central 1/5. Reticulation of ventrites 4 and 5 restricted to lateral third and with superimposed elongate furrows. Ventrites 2–5 doubly punctate; very fine, moderate and fine, sparse to very sparse punctures; punctures most evident in areas without reticulation. Ventrites 3–5 with transverse irregular row of long, yellowish, recumbent to decumbent setae laterally. Ventrite 6 (Figure 11C) with very fine, moderate punctures and medium to coarse, sparse to moderate punctures; punctures coarser close to apex. Elongate, semicircular wrinkles and channels apicolaterally and centrally; apicolateral sculpture extending basally around central portion of ventrite. Some punctures in channels bearing elongate, whitish, erect setae.

Figure 11. 

Meladema species males, abdominal ventrite 6 (DNA voucher codes where applicable). AM. coricaea, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., Corsica, Cap Corse (NHM-IRM12F) C M. imbricata, La Gomera, El Cedro (NHM-IRM3A) D M. lanio, Madeira, Ribeira dos Cedros (NHM-IRM8A). Scale bar = 1 mm.

Figure 12. 

Meladema species males, fore (A–D) and mesotarsi (E–H), ventral view (DNA voucher codes where applicable). A, E M. coriacea Spain, Cáceres, nr. Plasencia B, F M. lepidoptera sp. n. holotype, Corsica, Cap Corse (NHM-IRM12E) C, G M. imbricata, La Gomera, El Cedro (NMH-IRM3A) D, H M. lanio, Madeira, Rabacal. Scale bar = 1 mm.

Figure 13. 

Meladema species males, fore tarsal claws, lateral view (DNA voucher codes where applicable). A M. coriacea Spain, Cáceres, nr. Plasencia B M. lepidoptera sp. n. holotype, Corsica, Cap Corse (NHM-IRM12E) C M. imbricata, La Gomera, El Cedro (NMH-IRM3A) D M. lanio, Madeira, Rabacal. Scale bar = 1 mm.

Male. Foretarsi (Figure 12C) with articulo-setal counts as follows (base to apex): row 1, 8; row 2, 10; row 3, 10; row 4, 8. Curved, golden setae bordering articulo-setal field dense, particularly basally. Foretarsal claws (Figures 12C, 13C) elongate, curved. Mesotarsi (Figure 12G) with articulo-setal counts as follows (base to apex): row 1, 8; row 2, 9–10; row 3, 8–9; row 4, 7 (2 clusters, 4 on inner side, 3 on outer side, situated laterally). Curved, golden setae bordering articulo-setal field relatively dense, especially basally. Mesotarsal claws (Figure 12G) elongate, curved. Abdominal ventrite 6 (Figure 11C) with apex rounded, with well-marked median emargination. Median lobe asymmetrical (Figure 14C), sinuation strong, approximately 1/4–1/3 from apex; ventral margin of apical portion weakly concave in lateral view. Parameres (Figure 14C) with inner margin almost right-angled at base; outer and inner margins undulated slightly.

Female. As male, except for simple fore and mesotarsi and differently shaped abdominal ventrite 6 (with bluntly pointed apex - Figure 15C).

Variation. The size and density of the crescentic striolae on the elytra differs somewhat between individuals and localities (e.g. Figure 25D, F). At least some of this variation may be due to hybridization with M. coriacea (see below), making the extent to which this is truly intraspecific unclear. On La Palma, however, an island with no known populations of M. coriacea, and no genetic evidence of hybridization, the crescentic striolae are relatively very large and dense in some females (Figure 25F), approaching the situation seen in some females of M. lanio (see below).

Morphologically somewhat intermediate between M. coriacea/M. lepidoptera sp. n. and M. lanio. From M. coriacea and M. lepidoptera sp. n. M. imbricata can be distinguished on its different dorsal colouration, particularly the strongly mottled elytra, with much smaller, sparser crescentic striolae, as well as the less strongly sinuate posterior pronotal margin, details of the male genitalia (median lobe with sinuation further away from apex, with concave ventral margin in lateral view) and the last abdominal ventrites of both sexes. The habitus of M. imbricata is also typically more elongate than either of the above species (Figure 3). There are also additional minor differences in dorsal and ventral sculpture, as described above. With the exception of some females (see below), M. imbricata can be separated from M. lanio on the presence of crescentic striolae on the elytra. The male genitalia of the two species are also different, the sinuation of the median lobe of M. lanio occurring further from the apex than M. imbricata (see Figure 14C, D). M. imbricata also differs from all individuals of M. lanio in its less elongate habitus (Figure 3) and the much stronger sculpture of the metacoxae and abdominal ventrites.

Figure 14. 

Meladema species male genitalia. Median lobe, lateral and ventral view; paramere (DNA voucher codes where applicable). A M. coriacea Spain, Cáceres, nr. Plasencia B M. lepidoptera sp. n. holotype, Corsica, Cap Corse (NHM-IRM12E) C M. imbricata, La Gomera, El Cedro (NMH-IRM3A) D M. lanio, Madeira, Rabacal. Silhouette indicates orientation of median lobe for imaging in ventral view. Scale bar = 1 mm.

Figure 15. 

Meladema species females, abdominal ventrite 6 (DNA voucher codes where applicable). AM. coricaea, Spain, Murcia, Fte. Caputa B M. lepidoptera sp. n., Corsica, Cap Corse (NHM-IRM12C) C M. imbricata, La Palma, Bco. Hoyo Verde D M. lanio, Madeira, Ribeira dos Cedros (NHM-IRM9A). Scale bar = 1 mm.

Figure 16. 

Meladema coriacea, Var, France, La-Londe-les-Maures, female reproductive tract and genitalia (DNA voucher codes). A reproductive tract anatomy (NHM-IRM11A) B gonocoxae and gonocoxosternite (left gonocoxosternite removed) (NHM-IRM11B) C gonocoxae with laterotergites expanded (NHM-IRM11B). Scale bars = 1 mm.

Figure 17. 

Meladema coriacea, Sahara, elytral sculpture; shoulder and middle left and right, respectively. A male, Chad, Tibesti, Koudou B female, Chad, Tibesti, Bassin de Gorrom C male, Algeria, Hoggar, Aguelmanne D female, Algeria, Hoggar, Aguelmanne. Scale bar = 0.5 mm.

Endemic to the western Canary Islands (Figure 23), being erroneously reported from Madeira in the original description, as discussed by Machado (1987). We have only seen material from a single locality on Tenerife (upper reaches of Barranco del Río), one on La Gomera (El Cedro, in the laurel forest of Garajonay National Park), and two streams on La Palma (Barranco del Río and Barranco Hoyo Verde, situated on opposite sides of the Caldera de Taburiente). Lüderitz et al. (2010) report the species from one additional locality each on La Gomera and Tenerife, although the Tenerife locality (Barranco del Infierno, 500 m) has been visited by DTB in the early 2000s, when it contained only M. coriacea, as reported by Malmqvist et al. (1995). Additionally, neither of these records are mentioned by Lüderitz et al. (2016), casting some doubt on both of them. The number of permanent stream systems on the Canary Islands has declined seriously in recent decades, as a result of unsustainable water use. Streams have been variously diverted, piped, dammed, and negatively affected by abstraction directly from aquifers (Malmqvist et al. 1993, 1995, Kelly et al. 2002, Lüderitz et al. 2010, 2016). As a consequence, M. imbricata, which appears to be restricted to permanent reaches at relatively high altitude, particularly in forested regions, is very rare, being listed as Critically Endangered (1Ac) in the IUCN Red List (Foster 1996a). The species is also potentially threatened by hybridization with M. coriacea (see Ribera et al. 2003 and below), a process which may be further facilitated by ongoing climate change favouring the expansion of this more eurytopic species. Lüderitz et al. (2016) suggest that M. imbricata may have disappeared from the El Cedro stream on La Gomera recently, apparently being replaced by M. coriacea between 2006 and 2013. It is not clear, however, whether the same stream reaches were sampled on these two occasions. Work establishing the current status of this species in the Canary Islands is clearly a conservation priority.

“Maderae aquis”

(BMNH). Lectotype ♀ (herein designated, Figure 24B): “Dytiscus lanio// Fab. Entom. p. 231. n. 8.” [Latin name & 231. n. 8. HW] “Dytiscus lanio// Fabricius, 1775// LECTOTYPE// DT Bilton & I Ribera des. 2017” (dry pinned, BMNH, Banks Collection). Supposed syntypes comprise the above specimen and one other female, located in the Zoological Museum, University of Copenhagen (Zimsen, 1964; A. Solodovnikov, pers. comm.). Sree Selvantharan and Aslak Hansen have kindly communicated a photo of the Copenhagen specimen labelled as lanio (both on its pin, and above it in the cabinet). This is in fact a species of Cybister.

(genotyped specimens). Portugal, Madeira: 1 ♂ “vi/1998 PORTUGAL Madeira// Ribeira dos Cedros// L. C. Kelly leg.” “M3 Hand// Coleoptera” [HW] “Meladema// lanio” [HW] “DNA voucher// NHM-IRM8A” (IBE); 1 ♀ “vi/1998 PORTUGAL Madeira// Levada das Faias// L. C. Kelly leg.” “M2 handsearch// Coleoptera” [HW] “Meladema// lanio” [HW] “DNA voucher// NHM-IRM9A” (IBE); 1 ♀ “vi/1998 PORTUGAL Madeira// Levada das Faias// L. C. Kelly leg.” “DNA voucher// NHM-IRM9B” (IBE); 1 ♂ “Madeira, Canhas// lado sur de Paul da Serra// 1398 m 32°44'36"N 17°05'56"W// charca sobre arroyo de pietras// 19.7.2011 A. Rudoy leg.” “DNA Voucher// IBE-DV298” (IBE). All with “Meladema// lanio (Fabricius, 1775)// D T Bilton [or I Ribera] det. 2017”

(non-genotyped specimens). Portugal, Madeira: 1 ♀ “pouzo” [HW] “var. squamata” [HW] “passage à la m.// imbricatum Woll.” [HW] (ISNB); 1 ♀ “Madeira// 90.32.” [HW] (BMNH); 1 ♀ “Madeira// 90.32.” [HW] “Colymbetes// lanio// Fabr.” [HW] (BMNH); 1 ♀ “Madeira” reverse “48// 99” [small, circular label, HW] “Colymbetes// lanio Fabr.// Madeira” [HW] (BMNH); 1 ♀ “Madeira” reverse “48// 99” [small, circular label, HW] (BMNH); 1 ♂ “Sharp Coll.// 1905-313.” “Dytiscus lanio Fab.// (Scutopterus Woll. Meladema D. S.)// Madeira T. V. Wollaston” [HW] (BMNH); 2 ♂♂, 2 ♀♀ “Madeira// in Fauvel” [HW] “Coll. Odier.// B. M. 1921-288” (BMNH); 3 ♂♂, 3 ♀♀ “MADEIRA// Ribeira da St.// Luzia// 24.VIII.1964// E. W. Classey” [HW] “Brit. Mus.// 1968-48.” [8-48 HW] (BMNH); 3 ♂♂, 3 ♀♀ “MADEIRA:// Ribeira de St. Martinho// 13.VIII.1966// E. W. Classey” [HW] “Brit. Mus.// 1966-446” [6-446 HW] (BMNH); 6 ♂♂, 4 ♀♀ “MADEIRA:// Ribeiro de St.// Martinho// 13.VIII.1966// E. W. Classey” [HW] “Brit. Mus.// 1968-48.” [8-48 HW] (BMNH); 1 ♂, 2 ♀♀ “MADEIRA:// Ribeira da Abilheira// 19/VIII/1966// E. W. Classey” [HW] (BMNH); 2 ♂♂ “23/iii/1995 MADEIRA// Ribera on 103 below// Poiso & Monte ca.// 900m D. T. Bilton leg.” (CBP); 2 ♂♂, 1 ♀ “March 2001 MADEIRA// Levada @ Rabacal// D. T. Bilton leg.” (CBP); 1 ♂ “Madeira 1.-4.8.86// 1kn nördl. Poiso// Quellarn d. R. Frio// leg. Balke// Hendrich” “COLL.// HENDRICH// BERLIN” “Meladema// lanio F.” [HW] (CBP); 3 ♂♂, 3 ♀♀ “Madeira 8 Anchadas da Cruz// 932 m 32°48'35"N 17°17'47"W// charca de arroyo// 4.07.15 A. Rudoy leg.” (IBE); 1 ♂ “Madeira, Canhas// lado sur de Paul da Serra// 1398 m 32°44'36"N 17°05'56"W// charca sobre arroyo de pietras// 19.7.2011 A. Rudoy leg.” (IBE); 1 ♂ “lanio Fab.” [HW] (BMNH); 1 ♂ “40// 4 2// 2156” [small, circular label, HW] (BMNH); 1 ♂ “40// 4 2// 2157” [small, circular label, HW] “C. Lanio Fabr.” (BMNH); 2 ♂♂, 2 ♀♀ without labels (BMNH, Wollaston Collection); 1 ♂ “Meladema// lanio// Fauvel 1897// Madère” [HW] “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN); 1 ♂ “MUSÉUM PARIS// Coll. P. de Peyerimhoff// 1950” (MNHN). All with “Meladema lanio// (Fabricius, 1775)// D T Bilton [or I Ribera] det. 2017”.

Figure 18. 

Meladema species, Italy, male elytral sculpture; shoulder and middle left and right, respectively (DNA voucher codes, where applicable). A M. lepidoptera sp. n., Liguria, Levante B intermediate specimen, Campania, S. Michele (IBE-AN694) C intermediate specimen, Apulia, Vieste D M. coriacea, Lazio, Grotta di Pastena. Scale bar = 0.5 mm.

Figure 19. 

Meladema coriacea, Italy, Sicily elytral sculpture; shoulder and middle left and right, respectively (DNA voucher codes, where applicable). A male, Bosco Ficuzzo (IBE-AN691) B female, Monte Maganoce. Scale bar = 0.5 mm.

(based on all material examined).Size: Lectotype TL = 20.30 mm; EL = 16.30 mm; MW = 10.18 mm. Other material examined TL = 17.40–21.12 mm; EL = 12.67–15.36 mm; MW = 8.70–10.50 mm.

Colour. Dorsum (Figure 3D), dark reddish brown to yellow. Labrum yellowish, red anterolaterally in some specimens; clypeus yellowish except central ca. 1/6 red to blackish, connected to dark pigmentation on frons. Frons with transverse pale strip anterolaterally, adjacent to pale parts of clypeus, otherwise dark reddish brown. Medial, paired interocular patches on frons yellow; strongly transverse apicolaterally, almost reaching channel around interior margin of compound eye; confluent in centre in some specimens. Pronotum dark reddish brown on disc; narrowly reddish along anterior margin; lateral margins broadly yellowish. Elytra yellowish brown to greenish brown, with black irrorations; pattern visible without being lifted, but more evident when done so (Figures 3D, 4D). Legs yellowish brown to black; posterior tibiae and tarsi darkest. Antennae and maxillary and labial palpi yellowish to reddish. Venter reddish brown; prementum yellow; posterior genae, mentum and submentum reddish. Pronotal hypomeron and shoulder and outer portion of elytral epipleurs yellowish.

Head. Labrum shining, with medium to fine, sparse punctures. Reticulation absent anteriorly, clearly evident in posterior half, here fine and composed of small, isodiametric to transverse meshes. Clypeus weakly shining, with medium to fine, sparse punctures and traces of very fine, shallow, close punctures. Frons weakly shining, entire surface with, open reticulation; weak in front of interocular patches, becoming stronger and more evident posteriorly. Meshes transverse to isodiametric apically and medially, strongly elongate posteriorly and onto vertex. Paired epicranial foveae on anterior frons, one immediately behind the other. Anterior foveae transverse, posterior foveae elongate oval. Foveae reticulate; anterior and posterior foveae linked by reticulated channel. Internal and posterior borders of compound eyes distinct, raised relative to level of adjacent cuticle. Lateral margins bordered by distinct narrow channel; deeper anteriorly than posteriorly and continuing behind posterior margin of eye onto vertex. Channel with dense punctures, bearing long, stiff, yellow recumbent to decumbent setae.

Pronotum. Posterior margin weakly sinuate laterally (Figure 3D). Surface somewhat shining, strongly rugose. Reticulation meshes large, open, flat and with varying sizes and orientations. Transverse irregular row of medium punctures bearing long, yellow recumbent to decumbent setae 1/5 behind anterior pronotal margin; interrupted briefly in centre, obscured by reticulation inside lateral margins but continuing inside lateral third of posterior margin. Reticulation weak and obsolete anterior to transverse row, surface here clearly double punctate, with very fine, dense and medium, sparse to very sparse punctures. Scattered medium punctures visible elsewhere, amongst meshes of microreticulation. Centre of disc with elongate, narrow, slit-like fovea, partly obscured by reticulation in some specimens, but then traceable as an elongate reticulation channel. Lateral margins slightly raised, shining, without rugose sculpture and with fine, scattered punctures.

Elytra. Shining, without crescentic striolae (except in some females – see below). Surface doubly punctate and reticulate (Figure 5D); with very fine, close and fine to medium, sparse punctures (the latter bearing short, peg-like setae). Reticulation fine, somewhat obsolete, meshes isodiametric; sometimes obscuring very fine punctation. Surface weakly tuberculate towards sides and apex, particularly in posterior third (see Figure 3D). Puncture rows well-marked, continuous almost to elytral apices; punctures shallower posteriorly than anteriorly.

Figure 20. 

Meladema species, variation in male median lobes, lateral view. A–N M. coriacea; O–U M. lepidoptera sp. n. (DNA voucher codes, where applicable). A Spain, Huesca, Bernués (IBE-DV292) B Spain, Girona, Port Bou (IBE-DV-293) C Spain, Tarragona, Esblada D Spain, Cáceres, nr. Plasencia E Spain, Murcia, Fte. Caputa (NHM-IRM13) F Spain, Córdoba, Baena (NHM-IRM14C) G Tenerife, Chamorga (MNCN-AI1095) H Morocco, Oued Massa (NHM-IRM2A) I Morocco, Tazekka (NHM-IRM1C) J Morocco, Sidi-Ibrahim (NHM-IR7) K Algeria, Aïn Damous (MNCN-HI6) L Italy, Sicily, Bosco Ficuzzo (IBE-AN691) M Greece, Poros, Kampos N Greece, Ios O–P Corsica, Francardo Q holotype, Corsica, Cap Corse (NHM-IRM12E) R Italy, Montecristo (IBE-DV289) S Sardinia, Villagrande Strìsaili (IBE-RA5) T Italy, Elba, Pomonte (IBE-AN692) U Italy, Tuscany, S. Luce (IBE-AN693). Scale bar = 1 mm.

Figure 21. 

Meladema lepidoptera sp. n. elytral sculpture (males); shoulder and middle left and right, respectively (DNA voucher codes). A holotype male, Corsica, Cap Corse (NHM-IRM12E) B Italy, Montecristo (IBE-DV289) C Italy, Elba, Pomonte (IBE-AN692) D Italy, Tuscany, S. Luce (IBE-AN693). Scale bar = 0.5 mm.

Figure 22. 

Meladema lepidoptera sp. n., Sardinia, elytral sculpture SEMs; shoulder and middle left and right, respectively (DNA voucher codes). A male, Villagrande Strìsaili (IBE-RA5) B female, Tortoli (IBE-RA18).

Venter. Prementum shining, tumid in centre, with fine and medium, sparse punctures. Mentum shining; central projection with shallow median emargination. Lateral lobes with medium, sparse punctures, and scattered, whitish recumbent to decumbent setae. Submentum shining, with transverse wrinkles centrally, and elongate wrinkles laterally. Central 1/4 with medium, sparse punctures bearing long, white-yellowish, erect setae. Gula shining, with sparse, shallow, transverse wrinkles; patch of medium-coarse punctures posterolaterally. Genae shining, with obsolete, open, elongate reticulation. Prosternum shining, with weak, low, irregular transverse ridges laterally. Arched in centre and with fine, moderate to close punctures laterally, bearing long, white-yellowish, recumbent to erect setae; punctures and setae extending in an irregular row onto process, just below arch. Process lanceolate, arched; apex acuminately rounded. Centre of prosternum and process with double punctation of very fine, close to very close and medium, sparse to moderate punctures. Pronotal hypomeron shining, impunctate. Elytral epipleurs shining, with fine wrinkles; irregular puncture row close to internal margin, from centre to close to apex, punctures bearing fine, whitish, erect setae. Metaventrite shining, central portion with reticulation reduced to sparse, transverse scratches and very fine, close and fine to medium, sparse punctures. Metaventral process strongly reticulate, with transverse to elongate, rugose meshes and traces of fine, sparse punctures. Metaventral process relatively broad; apex acuminate and upturned slightly anterolaterally. Metacoxal lines not reaching anterior border of metacoxae, disappearing approx 1/10 from margin. Internal laminae of metacoxae shining, sculpture as on centre of metaventrite. Metacoxal lobes sculptured as internal laminae, strongly rounded, with irregular, elongate field of medium to coarse punctures close to lateral margins, bearing fine, white, recumbent to erect setae. External laminae of metacoxae shining, smooth close to process, and with obsolete reticulation elsewhere; without distinct meshes, wrinkled, elongate around anterior and posterior margins; doubly ounctate, with very fine, close and fine, very sparse punctures. Abdominal ventrites shining. Ventrites 3–5 with cluster of golden, erect setae anteromedially. Ventrite 1 with weak reticulation of elongate scratches throughout. Ventrite 2 with similar reticulation; absent close to centre. Ventrite 3 and 4 with scratches restricted to lateral 1/3. Ventrites 2–5 doubly punctate; very fine, moderate and fine, sparse to very sparse punctures; punctures most evident in areas without reticulation. Ventrites 3–5 with transverse irregular row of long, yellowish, recumbent to decumbent setae laterally. Ventrite 6 (Figure 11D) with very fine, moderate punctures and medium to coarse, sparse to moderate punctures; punctures coarser close to apex. Elongate, semicircular wrinkles and channels apicolaterally and centrally; apicolateral sculpture extending basally around central portion of ventrite. Some punctures in channels bearing elongate, whitish, erect setae.

Male. Foretarsi (Figure 12D) with articulo-setal counts as follows (base to apex): row 1, 8–9; row 2, 10; row 3, 10; row 4, 8. Number of setae in rows may differ from right to left tarsus in same beetle. Curved, golden setae bordering articulo-setal field relatively sparse, particularly laterally. Foretarsal claws (Figures 12D, 13D) elongate, curved. Mesotarsi (Figure 12H) with articulo-setal counts as follows (base to apex): row 1, 7–8; row 2, 8; row 3, 8; row 4, 6 (2 clusters of 3, situated laterally). Curved, golden setae bordering articulo-setal field relatively sparse, especially laterally. Mesotarsal claws (Figure 12H) elongate, curved. Abdominal ventrite 6 (Figure 11D) with apex rounded, with well-marked median emargination. Median lobe (Figure 14D) asymmetrical, sinuation strong, approximately 1/3–0.35 from apex; ventral margin of apical portion relatively straight in lateral view. Parameres (Figure 14D) with inner margin almost right-angled at base, with distinct small projection; outer margin undulated slightly; inner straight.

Female. As male, except for simple fore and mesotarsi, and differently shaped abdominal ventrite 6 (with bluntly pointed apex - Figure 15D).

Variation. Males and females generally have identical sculpture on the elytra. Two females studied, one from Ribeira da St. Luzia (Figure 26B, BMNH) and one from “pouzo” (ISNB) have distinct crescentic striolae, as have been observed in female M. imbricata from La Palma (see above). Females sculptured in this manner appear to be relatively rare, the only other occurrences of this form we can find being mentioned by Aubé (1838), who had seen a female with this sculpture (see also Sharp 1882) and Falkenström (1938). Such females may be mistaken for M. imbricata, as was the case with the ISNB animal listed above.

Figure 23. 

Meladema distribution, material examined. Symbols with black border show locations of genotyped specimens. Symbol colours as follows: M. coriacea – red; M. lepidoptera sp. n. – blue; M. imbricata – green; M. lanio – yellow. Bicoloured symbols, hybrid or morphologically intermediate individuals.

Figure 24. 

Meladema type labels and specimens in situ. A M. imbricata, holotype, BMNH, Wollaston Collection B M. lanio lectotpe, BMNH, Banks Collection.

Figure 25. 

Meladema species, Canary Islands, elytral shoulder sculpture SEMs A–E males F female (DNA voucher codes where applicable). A–B M. coriacea x imbricata hybrid, Tenerife, Bco. del Río 600 m (NHM-IRM16A) C M. coriacea, Tenerife, Bco. de Masca (NHM-IRM19A) D M. imbricata, La Gomera, El Cedro (NHM-IRM3A) E M. imbricata x coriacea hybrid, Tenerife, Bco. del Río 1,600 m (NHM-IRM5B) F M. imbricata, La Palma, Bco. Hoyo Verde.

Closest to M. imbricata; for diagnostic characters see under that species.

Restricted to the main island of Madeira (Figure 23), where it is relatively widely distributed in permanent streams, particularly in remnant laurel forests in the mountains. Can also be found in man-made levada systems. Much more abundant on Madeira than M. imbricata on the Canaries, but still listed as Vulnerable (B1 + 2b) on the IUCN Red List (Foster 1996b), reflecting its very small global range.