Sylvicanthon attenboroughi, Cupello & Vaz-De, 2018

Cupello, Mario & Vaz-De, Fernando Z., 2018, A monographic revision of the Neotropical dung beetle genus Sylvicanthon Halffter & Martínez, 1977 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini), including a reappraisal of the taxonomic history of ‘ Canthon sensu lato’, European Journal of Taxonomy 467, pp. 1-205 : 136-140

Sylvicanthon attenboroughi sp. nov.

urn:lsid:zoobank.org:act:2CE9C82F-6618-4579-BA18-D49F3C8BCED7

Figs 11H , 15L , 18D , 20 , 34 , 35B , 38 C–D

Sylvicanthon cf. sp. nov. – Larsen 2004: 261.

Sylvicanthon sp. 1 – Silva et al. 2014: 348.

Sylvicanthon sp. – Pacheco et al. 2016: 143, 147, fig. P.

Etymology

A tribute to the great British naturalist and broadcaster Sir David Attenborough. In recognition of his profoundly influential work on the public understanding of natural history and evolutionary biology, which, for more than six decades, has been inspiring young people of successive generations (including the first author) to pursue a career as a biologist and the general public to know and preserve the beautiful world in which we live. We paraphrase him: “I did so because I know of no pleasure deeper than that which comes from contemplating the natural world and trying to understand it” ( Attenborough 2002). The specific name is a noun in the genitive case.

Material examined

Holotype

BRAZIL: ♂, Mato Grosso, Cláudia, Fazenda Iracema , 11º37′44″ S, 55º05′54″ W, (“BRASIL: Mato Grosso. Cláudia. / Faz. Iracema. 11º37′44″ S; 55º0 / 5′54″ W. Hum. dung. 20-II-2011. / M. F.Souza. ”), genital capsule extracted and glued to a triangular card ( CEMT).

Paratypes (63 ♂♂, 67 ♀♀)

BRAZIL: Acre: ♀, Manoel Urbano, Parque Estadual Chandless, 09º22′26″ S, 69º55′20″ W, 24 Jun. 2013, T.F. Brito leg. ( CEMT); 2 ♂♂ (1 dissected), Mâncio Lima, 07º28.584′ S, 72º54.110″ W, 5 Dec. 2012, H.M.B. Luiz & N.S.G.F. Adem leg. ( CEMT); 1 ♀, Senador Guiomard [“Rio Branco”], Fazenda Experimental Catuaba [“ Catuaba ”], 9 Apr. 1996, A. Bonaldo leg. ( MCNZ); 1 ♂, 4 ♀♀, Senador Guiomard [“Rio Branco”], Fazenda Experimental Catuaba, Feb. 1997 , F.Z. Vaz-de-Mello leg. ( CEMT). – Amazonas: 1 ♂, Benjamin Constant, Sep. 1962, Seabra leg. ( CMNC); 1 ♂, Borba, BR-319, km 220, 04º22′55″ S, 60º57′19″ W, 29 Nov. 2015, pitfall with human faeces, D. Pires leg. ( CEMT); ♀, Borba, BR-319, km 220, 04º24′06″ S, 60º55′22″ W, 30 Nov. 2015, pitfall with human faeces, D. Pires leg. ( INPA); 1 ♂ (dissected), Careiro, BR-319, KM 34, 03º21′46″ S, 59º51′10″ W, 9 Dec. 2015, pitfall with human faeces, D. Pires leg. ( INPA); 1 ♂ (dissected), Manaquiri, BR-319, km 100, 03º40′34″ S, 60º17′46″ W, 6 Dec. 2015, pitfall with human faeces, D. Pires leg. ( INPA). – Mato Grosso: 8 ♀♀, Alta Floresta, 09º56′52″ S, 56º03′02″ W, May 2008, pitfall with human faeces, E. Berenguer leg. ( CEMT); 3 ♂♂, 5 ♀♀, Alta Floresta, 09º53′40″ S, 56º16′35″ W, Jun. 2008, pitfall with human faeces, E. Berenguer leg. ( CEMT); 1 ♀, Aripuanã, 10º03′10″ S, 59º29′42″ W, 320 m, 26 Jan. 2012, H.A.B. Faria leg. ( CEMT); 1 ♀, Cláudia, Fazenda Iracema, 11º37′44″ S, 55º05′54″ W, 20 Feb. 2011, human faeces, M.F. Souza leg. ( CEMT); 2 ♀♀, Novo Mundo, Parque Estadual do Cristalino, 09º27′53″ S, 55º49′30″ W, May 2013, pitfall, V. Magalhães leg. ( CEMT); 1 ♀,Querência, Fazenda São Luiz, 12º39.85′ S, 52º22.18′ W, 50 m, Feb. 2009, flight interception trap, R. Andrade leg. ( CEMT). – Pará: 1 ♂ (dissected), 1 ♀, Itaituba, Uruá, 65 km SW of Itaituba on BR230 [“BR320”], 12–15 Oct. 1977, B.C. Ratcliffe leg. ( CMNC); 1 ♀, Santarém, Reserva Extrativista Tapajós-Arapiuns, 03º03′ S, 55º30′ W, 22 Dec. 2008, flight interceptation trap ( CEMT); 1 ♂ (dissected), 1 ♀, Santarém, Reserva Extrativista Tapajós-Arapiuns, 02º36′662″ S, 55º36′513″ W, 7 Jan. 2009, pitfall with human faeces, R. Andrade leg. ( CEMT). – Rondônia: 1 ♂, Itapuã do Oeste, Floresta Nacional [“FloNa”] do Jamari, 09º05′21″ S, 63º09′48″ W, 20 Feb. 2013, Mazama gouazoubira dung, J.F. Cerveira leg. ( CEMT); 1 ♂, 1 ♀, Itapuã do Oeste, Floresta Nacional [“FloNa”] do Jamari, 09º05′20″ S, 63º09′47″ W, 24 Feb. 2013, Mazama Nana dung, J.F. Cerveira leg. ( CEMT); 2 ♂♂, 2 ♀♀, Itapuã do Oeste, Floresta Nacional [“FloNa”] do Jamari, 09º05′19″ S, 63º09′42″ W, 26 Feb. 2013, Mazama Nana dung, J.F. Cerveira leg. ( CEMT); 21 ♂♂, 6 ♀♀, Guajará-Mirim, 10º44′53.56″ S, 65º17′31.1″ W, 14–16 Feb. 2010, pitfall with human faeces, Fabricio Coletti leg. ( CEMT); 1 ♂, Guajará-Mirim, Fazenda Agropecuária A.J.B., 10º37′47.59″ S, 64º59′52.58″ W, 180 m, 15 Jan. 2010, human faeces, F. Coletti leg. ( CEMT); 1 ♀, Guajará-Mirim, Fazenda Benezi, 10º44′53.56″ S, 65º17′31.10″ W, 170 m, 16 Feb. 2010, human faeces, F. Coletti leg. ( CEMT); 1 ♂, Ji-Paraná (“V. Rondonia (378 km S de P. Velho)”), 25 Jan.–9 Feb. 1961, Pereira and Machado leg. ( MZSP); 1 ♂, Pimenta Bueno, 11º43′04.43″ S, 61º28′21.45″ W, 249 m, 9–11 Dec. 2015, pitfall with human faeces, D.C. Castro leg. ( CEMT); 1 ♂, Porto Velho, Bom Jesus, Rio das Garças, 08º49′48″ S, 63º46′45″ W, 20 Apr. 2017, flight interception trap, D.C. Santos and K.K.G. Silva leg. ( CEMT); 1 ♂, Porto Velho, Bom Jesus, Rio das Garças, 08º49′48″ S, 63º46′45″ W, 27 Jul. 2017, flight interception trap, D.C. Santos and K.K.G. Silva leg. ( CEMT); 1 ♂, Porto Velho, Bom Jesus, Rio das Garças, 08º49′47″ S, 63º46′51″ W, 29 Nov. 2017, flight interception trap, D.C. Santos and K.K.G. Silva leg. ( CEMT); 1 ♂ (dissected), Porto Velho, ESEC Cuniã, 08º04′11.82″ S, 63º28′34.64″ W, 15 Aug. 2012, pitfall with human faeces, M.A.P.A. Silveira leg. ( CEMT); 1 ♂, 1 ♀, Porto Velho, ESEC Cuniã, 08º04′11.82″ S, 63º28′34.64″ W, 83 m, 10–12 Nov. 2013, pitfall with human faeces, M.A.P.A. Silveira leg. ( CEMT); 1 ♂, 4 ♀♀, Porto Velho, Nova Mutum-Paraná, 09º26′24″ S, 60º13′19″ W, 27–28 Feb. 2010, human faeces, L.R. Silva and R.L.R. Silva leg. ( CEMT); 1 ♂, Porto Velho, Nova Mutum-Paraná [“Mutum”], 09º35′46″ S, 65º02′27″ W, Jan. 2012, R.V. Nunes leg. ( CEMT); 1 ♀, Porto Velho, Nova Mutum-Paraná [“Mutum”], 09º35′46″ S, 65º02′27″ W, 250 m, Sep. 2012, pitfall with human faeces, R.V. Nunes leg. ( CEMT); 1 ♀, Vilhena, Nov. 1986, O. Roppa and P. Magno leg. ( CEMT); 7 ♂♂ (1 dissected), 7 ♀♀, Same collection data as for preceding ( MNRJ); 1 ♂, 2 ♀♀, Same collection data as for preceding ( BMNH).

PERU: Cuzco: 1 ♀, La Convención, Echarate, Comunidad Kitaparay, 12º12′47.24″ S, 72º49′16.12″ W, 479 m, 13 Nov. 2009, C. Espinoza and E. Rázuri leg. ( MUSM). – Junín: 1 ♂, Satipo, Mashira, Río Tambo, 11º25′20.25″ S, 73º27′16.14″ W, 672 m, 2 Feb. 2010, J. Grados leg. ( MUSM). – Madre De Díos: 1 ♂, Manu, Huepetuhe (“Huaypetue”), 12º59′27.07″ S, 70º32′06.74″ W, 15–17 Nov. 2010, J. Costa and M. Vilchez leg. ( MUSM); 2 ♂♂ (1 dissected), 1 ♀, Manu, Manu, Salvación (“near Salvación”), 13º50′37″ S, 71º19′57″ W, 650 m, Nov. 1999, pitfall with human faeces, T. Larsen leg. ( CMNC); 1 ♂, Manu, Pakitza Biological Station, 28 Sep. 1987, B. Trocha leg. ( MUSM); 2 ♀♀, Manu, Pantiacolla Lodge, 8 km NW of El Mirador Trail, Alto Madre de Dios River, 12º38′30″ S, 71º16′41″ W, 800 m, 23–26 Oct. 2000, flight interception trap, R. Brooks leg. ( CMNC); 1 ♂ (dissected), 1 ♀, Manu, Parque Nacional del Manu (“Manu National Park”), 15–30 Aug. 1986, A. Forsyth leg. ( CMNC); 1 ♀, Manu, Parque Nacional del Manu, Estación Biológica Cocha Cashu, 11º53′45″ S, 71º24′24″ W, 350 m, 17–19 Oct. 2000, flight interception trap, R. Brooks leg. ( CMNC); 1 ♀, Manu, Reserva Comunal Amarakaeri, 12º59′51.87″ S, 70º50′26.05″ W, 864 m, 2–4 Nov. 2010, J. Costa and M. Vilchez leg. ( MUSM); 2 ♀♀, Manu, Reserva Comunal Amarakaeri, 12º59′51.87″ S, 70º50′26.05″ W, 864 m, 10–14 Nov. 2010, J. Costa and M. Vilchez leg. ( MUSM); 1 ♂ (dissected), Río Amiguillos, 12º22′25.4″ S, 70º22′13.2″ W, 260 m, May 2000, human faeces, T. Larsen leg. ( MUSM); 1 ♂, 2 ♀♀, Tambopata, Puerto Madonado, Sudadero, 12º21′19″ S, 69º01′48″ W, 221 m, 26–27 Mar. 2009, L. Figueroa leg. ( MUSM).

Description

COLOURATION. Entire body with very dark tonalities. Head and pronotum dark purple. Elytra dark green or dark blue; when green, striae dark blue and slightly contrasting with the rest of tegument. Metaventrite black. Meso- and metafemora orangish-brown, reddish-brown, or dark brown, with base distinctly darker than at least apical two-thirds. Pygidium usually coppery; occasionally, with greenish reflections.

HEAD. Tegument little shiny, with strong alveolar microsculpture; micropunctation evident on posterior region of clypeus and especially on frons. Clypeus with two apical teeth obtuse and only slightly separated; with single transverse row of short setae covering base of both teeth. Genae with strong tooth immediately behind clypeal-genal juncture. Posterior edge of head completely unmargined.

THORAX. Pronotum with slightly shiny and lustrous tegument, with very fine alveolar microsculpture (sometimes effaced at centre), and with dense, clearly marked micropunctation at centre, progressively weaker towards the sides. Posterior edge with fine transverse line at centre (usually extending only up to second elytral stria; sometimes effaced and almost indistinct). Hypomeral cavity entirely glabrous or at most with very few short setae at centre; long setae, if present, restricted to posterior region; external margin with weak tubercle. Metaventrite entirely glabrous; tegument with strong rivose microsculpture on anterior region and weaker adjacent to internal margin of mesocoxae; at centre, alveolar microsculpture very fine and progressively more diffuse and undifferentiated towards posterior region; micropunctation very fine, but usually distinct.

LEGS. Ventral surface of all femora and tibiae shiny. Profemora with tegument with strong rivose microsculpture and without micropunctation. Protibiae narrow and with internal edge straight and simple, without expansion; at apical third, with three small acute teeth on external edge, two most apical ones of subequal length and larger than basal ( Fig. 11H ). Mesofemora margined anteriorly only at basal half; unmargined portion of anterior edge with row of very short setae; posterior margin absent; tegument with effaced rivose microsculpture. Metafemora margined only anteriorly, posterior margin absent; apical half of anterior edge covered by row of setae; tegument covered by effaced rivose microsculpture and, in a few specimens, with traces of coarse elongate punctation at base. Metatarsomeres II and V subequal in length and longer than the others; metatarsomere IV shorter than the others.

ELYTRA. With at most nine very narrow visible striae: in general, first five or six striae well marked, very finely carinulate and slightly widened at base; remaining striae progressively more effaced; eighth and ninth striae seen only in specimens with very well-marked striae and, in these cases, always very subtle; all striae lack carinulae before reaching apex of elytra, where completely indistinct; humeral carina absent. Tegument of interstriae with diffuse shine; at centre, with alveolar microsculpture ranging from clearly marked, in most specimens, to diffuse; at apex, always with strong alveolar microsculpture; micropunctation in general clearly visible at 20 × magnification.

ABDOMEN. Tegument of ventrites I–V with strong rivose microsculpture at centre; ventrite VI with very diffuse rivose microsculpture and very subtle micropunctation; both sexes without lateral foveae. Pygidium with bright tegument covered by alveolar microsculpture; micropunctation subtle, but always evident among microsculpture.

AEDEAGUS. Parameres at least half as long as phallobase and asymmetrical: external face of right paramere flat, external face of left paramere concave, strongly excavated. In lateral view, parameres with strong ventral keel ( Fig. 18D ).

SEXUAL DIMORPHISM. Males: Protibial spur narrow and bifid at apex, with spiniform projections, external projection longer than internal one ( Fig. 15M ). Ventrite VI with posterior margin strongly narrowed at centre; anterior edge covered very slightly by weak medial flange of ventrite V. Females: Protibial spur simple, spiniform. Ventrite VI very broad at centre; anterior edge slightly covered by medial flange of posterior edge of ventrite V.

Measurements

Males (N = 13). TL: AV: 8.5 ± 0.54; MX: 9.6; MN: 7.5. EW: AV: 6.1 ± 0.35; MX: 6.6; MN: 5.5. PrL: ME: 2.5 ± 0.14; MX: 2.9; MN: 2.3. PrW: AV: 5.1 ± 0.29; MX: 5.5; MN: 4.6. PgL: AV: 1.6 ± 0.11; MX: 1.7; MN: 1.3. PgW: AV: 2.6 ± 0.18; MX: 2.9; MN: 2.3.

Females (N = 13). TL: AV: 8.4 ± 0.55; MX: 9.4; MN: 7.2. EW: AV: 6.1 ± 0.29; MX: 6.8; MN: 4.9. PrL: AV: 2.5 ± 0.14; MX: 2.8; MN: 2.4. PrW: AV: 5.2 ± 0.32; MX: 5.8; MN: 4.6. PgL: AV: 1.7 ± 0.15; MX: 1.9; MN: 1.4. PgW: AV: 2.6 ± 0.20; MX: 2.9; MN: 2.3.

Geographical distribution

Southern Amazonia in Brazil and Peru.

Ecoregions

Southwest Amazon Moist Forests, Peruvian Yungas, Madeira-Tapajós Moist Forests, Mato Grosso Tropical Dry Forests.

Collecting sites ( Fig. 34 )

BRAZIL. Acre: Mâncio Lima, Manoel Urbano (Parque Estadual Chandless), Rio Branco. Amazonas: Benjamin Constant, Borba, Careiro, Manaquiri. Pará: Itaituba (Uruá), Santarém (Reserva Extrativista Tapajós-Arapiuns) . Rondônia: Guajará-Mirim, Itapuã do Oeste (Floresta Nacional do Jamari), Ji- Paraná, Pimenta Bueno, Porto Velho, Vilhena. Mato Grosso: Alta Floresta, Aripuanã, Cláudia, Novo Mundo (Parque Estadual do Cristalino), Querência.

PERU. Junín: Satipo (Mashira). Cuzco: La Convención (Echarate). Madre de Díos: Manu (Huepetuhe; Manu: Salvación; Pakitza Biological Station; Pantiacolla Lodge; Parque Nacional del Manu; Reserva Comunal Amarakaeri), Tambopata (Puerto Madonado: Sudadero).

Intraspecific variation and taxonomic discussion

The distribution of S. attenboroughi sp. nov. is very interesting because, based on the data we have on hand, it seems to be clearly delimited by two major rivers in the Amazon Basin: the Amazon, which constrains dispersal towards forests farther north, and the Tapajós , the eastern limit of the distribution of S. attenboroughi sp. nov. in the centre of the Amazon Basin (i.e., excluding the populations in southern Amazonia in the state of Mato Grosso). Beyond those rivers (i.e., to the left banks of the Amazon and to the right banks of the Tapajós ) inhabits S. seag sp. nov., a species very similar to S. attenboroughi sp. nov. and its possible sister species. Southern populations of S. seag sp. nov. (i.e., those closer to the range of S. attenboroughi sp. nov.) show colouration and pygidial tegument very similar to (if not totally indistinguishable of) those of S. attenboroughi sp. nov., but both species are easily separated from one another by the shape of the parameres, which are much simpler in S. attenboroughi sp. nov. ( Fig. 18D ) than the very complex shape seen in S. seag sp. nov. ( Fig. 19A ). Other differences that do not vary geographically between these two species are the shape of the male protibial spur and the shape of the anterior margin of female ventrite VI ( Table 4; see also section on S. seag sp. nov. for a detailed discussion on the geographical variation seen in that species).

Sylvicanthon bridarollii is the only representative of the bridarollii subgroup to be found in sympatry with S. attenboroughi sp. nov. Both species are differentiated by the tegument of the head, pronotum, metaventrite and pygidium, the presence or absence of setae on the hypomeral cavity ( Fig. 35 ), the shape of protibiae ( Fig. 11 F–H), the presence or absence of a coarse punctation at the base of the metafemora ( Fig. 13 F–H) and, finally, by the shape of the parameres ( Fig. 18A, D ) ( Table 4). From S. edmondsi sp. nov., the fourth species of the bridarollii subgroup, S. attenboroughi sp. nov. is different in colouration ( Fig. 38 ), size, head tegument and hypomeral cavity; the parameres, which are usually very distinctive among the species of Sylvicanthon , do not show any remarkable differences between S. attenboroughi sp. nov. and S. edmondsi sp. nov. ( Fig. 18 C–D) ( Table 4).

Two intraspecifically-variable characters seen in S. attenboroughi sp. nov. call attention: the shape of the protibiae and the presence of coarse punctation at the base of the metafemora. In the great majority of the individuals of this species, the internal margin of the protibiae is straight ( Fig. 11H ) and the metafemora do not show any trace of coarse punctation at their base ( Fig. 13F ), in the same way as in S. seag sp. nov. and S. edmondsi sp. nov. However, some very few specimens from Rondônia ( Brazil) have the internal margin of the protibiae with a weak indication of expansion (similar to specimens of S. bridarollii from Ecuador and Colombia) and weak traces of a coarse punctation at the base of the metafemora (in the same way as in the populations of S. bridarollii from northern Peru). Future phylogenetic analyses will unveil whether those features are homologous to the states seen in S. bridarollii or whether they are homoplastics; if they are homologous, it will be interesting to know whether they constitute a synapomorphy between these two species or a symplesiomorphy in the subgroup that was completely lost independently in the majority of the S. attenboroughi sp. nov., the northern populations of S. bridarollii and by S. edmondsi sp. nov. and S. seag sp. nov..

Natural history

Very little information on the biology of S. attenboroughi sp. nov. is available on specimen labels. Adult activity seems to be restricted to the spring and summer months, with records from all months between September and March. Specimens were collected using pitfall traps baited with human faeces and flight interception traps. In relation to the altitudinal amplitude, S. attenboroughi sp. nov. seems to occur between 170 and 864 m a.s.l.

Figures

Fig. 11. Morphological diversity in the protibiae of Sylvicanthon Halffter & Martínez, 1977. A. S. enkerlini (Martínez et al., 1964) comb. nov. B. S. aequinoctialis (Harold, 1869) comb. nov. C. S. securus (Schmidt, 1920) comb. nov. D. S. obscurus (Schmidt, 1920). E. S. furvus (Schmidt, 1920). F. S. bridarollii (Martínez, 1949) (southern populations). G. S. bridarollii (northern populations). H. S. attenboroughi sp. nov. I. S. edmondsi sp. nov. J. S. candezei (Harold, 1869). Note the variation related to the presence and degree of development of an expansion on the internal edge of protibiae and to the number, size and position of the external teeth.

Fig. 15. Morphological variation and sexual dimorphism in the protibial spur of Sylvicanthon. A. S. enkerlini (Martínez et al., 1964) comb. nov., ♂. B. S. foveiventris (Schmidt, 1920), ♂. C. S. candezei (Harold, 1869), ♂. D. S. genieri sp. nov., ♂. E. S. aequinoctialis (Harold, 1868), ♂. F. S. aequinoctialis comb. nov., ♀. G. S. proseni (Martínez, 1949) stat. et comb. nov., ♂. H. S. proseni stat. et comb. nov., ♀. I. S. bridarollii (Martínez, 1949), ♂. J. S. seag sp. nov., ♂. K. S. edmondsi sp. nov.,♂. L. S. attenboroughi sp. nov., ♂. M. S. obscurus (Schmidt, 1920), ♂. N. S. furvus (Schmidt, 1920), ♂. O. S. securus (Schmidt, 1920) comb. nov., ♂.

Fig. 18. Aedeagus morphological diversity in the bridarollii subgroup.A. S. bridarollii (Martínez, 1949). B. S. seag sp. nov. (arrow points to the ventral notch). C. S. edmondsi sp. nov. (idem). D. S. attenboroughi sp. nov. (idem).

Fig. 20. Combined distribution of the species of Sylvicanthon in the Neotropical region. Note that the genus occupies the four great areas of tropical humid forest of the American continent: Central American forests, the Choco of the South American northwest, the Amazon Rainforest and the Atlantic Forest. Following the hypothesis of Martínez et al. (1964) that the Amazon Basin was the ancestral area of the Sylvicanthon of the candezei group, three lineages have independently dispersed from there (arrows): that of the ancestors of S. aequinoctialis (Harold, 1868), which, from the Choco, invaded Central America at the pace of the tropical forests’ gradual northwards advance, occurred during the Plio-Pleistocene, and those of the ancestors of respectively S. obscurus (Schmidt, 1920) and S. foveiventris (Schmidt, 1920), which reached the Atlantic Forest through the corridors of humid forests that bridged that biome to the Amazon forest during the wettest periods of the Neogene (see a more detailed discussion in the text).

Fig. 34. Distribution of the four species of the bridarollii subgroup.

Fig. 35. Variation on the pilosity at the centre of the hypomeral cavity among members of the bridarollii subgroup. A. Sylvicanthon seag sp. nov. B. S. attenboroughi sp. nov. C–D. S. bridarollii (Martínez, 1949). Note the first two species have the hypomeral cavity entirely glabrous or with setae limited to its periphery (centre always glabrous), whereas S. bridarollii possesses long and dense setae throughout the tegument of the hypomeral cavity.

Fig. 38. A–B. Sylvicanthon edmondsi sp. nov. A. Dorsal view. B. Ventral view. C–D. Sylvicanthon attenboroughi sp. nov. C. Dorsal view. D. Ventral view.

Fig. 19. Aedeagus morphological diversity in Sylvicanthon. A. S. securus (Schmidt, 1920) comb. nov. B. S. obscurus (Schmidt, 1920). C. S. furvus (Schmidt, 1920). D. S. monnei sp. nov. E. S. mayri sp. nov.

Fig. 13. Morphological variation in the metafemora of Sylvicanthon Halffter & Martínez, 1977. A. S. foveiventris (Schmidt, 1920). Note the presence of coarse elongate punctures at the base of the metafemur (arrow) and the tegument without microsculpture. B. S. obscurus (Schmidt, 1920). Observe the tegument entirely covered by coarse non-elongate punctures. C. S. monnei sp. nov. D. S. mayri sp. nov. E. S. furvus (Schmidt, 1920). F. S. edmondsi sp. nov. G. S. bridarollii (Martínez, 1949) (southern populations). H. S. bridarollii (Martínez, 1949) (northern populations). Note the differences in colouration and in the presence of coarse punctation at the base of metafemur in the two ends of the clinal variation observed in S. bridarollii.