Sylvicanthon genieri, 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 : 80-85

Sylvicanthon genieri sp. nov.

urn:lsid:zoobank.org:act:AB21F887-15E7-40CD-BBA8-673CDB21ED37

Figs 6C , 15D , 17D , 20 , 24 , 27

Sylvicanthon sp. 2: Celi et al. 2004: 46.

Sylvicanthon candezei – Carvajal et al. 2011: 117, 316 (error).

Etymology

The specific name, a noun in the genitive case, is an homage to the Canadian entomologist François Génier, one of the leading specialists in Scarabaeinae and author of some great works on this group, in recognition of his immense help and kindness in loaning more than one thousand specimens of Sylvicanthon (including 131 S. genieri sp. nov.) from the CMNC to us. As far as we know, Génier was the first entomologist to recognize this species as new, having identified since 1998 several specimens in the CMNC collection as “S ylvicanthon sp. nov”.

Material examined

Holotype

ECUADOR: ♂, Tungurahua, 6 km east from Río Negro , 1500 m (“ECU: Tungurahua / 6 kmE RioNegro 1500m / 13–17.vii.76 S.Peck / for. car. tps. 40–50”), genital capsule removed and glued to a triangular label ( CMNC).

Paratypes (112 ♂♂, 89 ♀♀)

ECUADOR: Morona Santiago: 1 ♂, Untsuants, site 4, 1100 m, 20 Jan. 2002, J. Celi and M. Ortega leg. ( CMNC); 3 ♂♂, Untsuante, site 7, 900 m, 23 Jan. 2002, J. Celi and M. Ortega leg. ( CMNC). – Napo: 11 ♂♂, 7 ♀♀, “Km 7.3 Sarayacu-Loreto Road”, 1200 m, 11 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♀, same collecting data as for preceding ( MCNZ); 7 ♂♂, 3 ♀♀, “Km 7.3 Sarayacu-Loreto Road”, 1200 m, 14 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 3 ♂♂, 2 ♀♀, “Km 7.3 Sarayacu-Loreto Road”, 1200 m, 20 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♂, 1 ♀, “Km 11.1 Sarayacu-Loreto Road”, 1200 m, 20 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♀, same collecting data as for preceding ( MCNZ); 2 ♂♂, 2 ♀♀, “Km 25.4 Sarayacu-Loreto Road”, 950 m, 12 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 3 ♂♂, 3 ♀♀, “Km 25.4 Sarayacu-Loreto Road”, 950 m, 14 Jul. 1994, dung trap, F. Génier leg. ( CMNC). – Orellana: 1 ♀, Onkone Gare Camp, 00º39′10″ S, 76º26′00″ W, 220 m, 4–12 Oct. 1995, active collecting beneath the leaf litter, T.L. Erwin Ecuador Expedition 1995, G.E. Ball and D. Shpeley leg. ( CMNC). – Pastaza: 3 ♀♀, 4.3 km Rio Negro, 1200 m, 18 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♂, “09 km ESE Veracruz”, 22–24 Aug. 1975, dung trap, R. Webster leg. ( CMNC); 1 ♂, 1 ♀, Pastaza, “ 22 km SE Puyo”, 900 m, 16 Jul. 1976, dung trap, S. Peck leg. ( CMNC); 5 ♂♂, 2 ♀♀, Pastaza, Puyo, Llandia, “ 17 km N Puyo”, 19 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 2 ♂♂, Mera, 1 km E Mera, 1100 m, 13–17 Jul. 1976, dung trap, S. Peck leg. ( CMNC); 1 ♂, Pastaza, Puyo, Llandia, “ 17 km N Puyo”, 20 Jul. 1994, dung trap, F. Génier leg. ( CMNC). – Tungurahua: 1 ♂, 3 km W Río Negro, 1200 m, 18 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♂, 4.3 km E Río Negro, 1200 m, 18 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 1 ♂, 3 ♀♀, 4.3 km E Río Negro, 1200 m, 20 Jul. 1994, dung trap, F. Génier leg. ( CMNC); 3 ♂♂, 6 km E Río Negro, 1500 m, 13–17 Jul. 1976, dung trap, S. Peck leg. ( CMNC); 16 ♂♂ (1 dissected), 19 ♀♀, 8 km E Rio Negro, 10 km W Pastaza (= Shell), 1400 m, 13–17 Jul. 1976, dung trap, S. Peck leg. ( CMNC); 9 ♂♂, 6 ♀♀, Baños, “El Topo”, 01º23′41″ N, 78º22′52″ W, 1590 m, 23 Jan. 2011, human faeces, G. Maldonado leg. ( CEMT). – Zamora Chinchipe: leg. 2 ♂♂, Upper Río Comainas, Cordillera del Cóndor, 03º54′ S, 78º25′ W, 1150 m, Jul. 1994, human faeces, A. Forsyth ( MUSM).

PERU: Amazonas: 5 ♂♂, 3 ♀♀, Comainas River, Jul. 1994, A. Forsyth leg. ( MUSM); 4 ♂♂, 1 ♀, Comainas River, 21 Jul. 1994, A. Forsyth leg. ( MUSM); 7 ♂♂, 12 ♀♀, Comainas River, 23 Jul. 1994, A. Forsyth leg. ( MUSM). – Cuzco: 1 ♀, La Convención, Echarate, Campamento Segakiato, 11º45′38.6″ S, 73º14′57.7″ W, 908 m, 2 Mar. 2011, M. Alvarado and E. Rázuri leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Otsanampiato, 12º39′32.18″ S, 73º09′18″ W, 1681 m, 22–24 Jan. 2010, C. Carranza and C. Rossi leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Otsanampiato, 12º39′41.03″ S, 73º09′30.98″ W, 1723 m, 15–17 Sep. 2010, pitfall, M. Alvarado and J. Peralta leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º33′54.41″ S, 73º05′36.85″ W, 1747 m, 26–29 Jan. 2010, C. Carranza and C. Rossi leg. ( MUSM); 1 ♀, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′13.56″ S, 73º05′12.13″ W, 1707 m, 18–21 Sep. 2010, M. Alvarado and J. Peralta leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′12.73″ S, 73º05′13.73″ W, 1709 m, 18–21 Sep. 2010, M. Alvarado and J. Peralta leg. ( MUSM); 1 ♂ (dissected), 2 ♀♀, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′13.18″ S, 73º05′50.23″ W, 1422 m, 14 Oct. 2009, light, C. Carranza and C. Rossi ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′10.12″ S, 73º05′13.85″ W, 1587 m, 18–21 Sep. 2010, M. Alvarado and J. Peralta leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′21.70″ S, 73º05′16.91″ W, 1685 m, 26–29 Jan. 2010, C. Carranza and C. Rossi leg. ( MUSM); 1 ♂, 2 ♀♀, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′10.72″ S, 73º05′15.17″ W, 1693 m, 18–21 Sep. 2010, pitfall, M. Alvarado and J. Peralta leg. ( MUSM). – Huánuco: 1 ♂, 3 ♀♀, Leoncio Prado, Rupa-Rupa, Tingo María, Universidad Nacional Agraria de la Selva (“Tingo María Universidad”), Jul. 1974 ( CMNC); 10 ♂♂ (1 dissected), 5 ♀♀, Leoncio Prado, Rupa-Rupa, Tingo María, Universidad Nacional Agraria de la Selva (“Tingo María Universidad”), Dec. 1974 ( CMNC). – Junín: 1 ♀, Chanchamayo, San Ramón, 10 May 2002, F.G. Horgan leg. ( MUSM); 2 ♀♀, Chanchamayo, San Ramón, Oct. 2002, pitfall with human faeces, F.G. Horgan leg. ( MUSM). – San Martín: 1 ♀, Rioja, Yuracyacu, 05º57′07″ S, 77º11′12″ W, 805 m, 6–7 Sep. 2008, C. Albujar leg. ( MUSM); 1 ♂ (dissected), Mariscal Cáceres, Juanjuí, Mar. 2010 ( CEMT).

Incorrect data: VENEZUELA: Bolívar: 1 ♂ (dissected), “ 150 km sur Caicara, R. Suapare”, 3–10 Apr. 1977, Bordón leg. ( CMNC).

Description

COLOURATION. Head, pronotum, elytra, metaventrite, and pygidium largely coppery; pygidium and, more rarely, centre of head occasionally with greenish reflections. Meso- and metafemora orangish.

HEAD. Tegument shiny, with strong alveolar microsculpture on frons, and more diffuse (yet clearly present) microsculpture on clypeus and genae; micropunctation fine and very subtle, sometimes almost imperceptible throughout most of the head, but often clearly seen on frons ( Fig. 6C ). Clypeus with two apical teeth obtuse and contiguous at base; with a single transverse row of setae covering the base of both teeth. Genae with a weak tooth immediately behind clypeal-genal juncture. Posterior edge of head unmargined between eyes.

THORAX. Pronotum with shiny tegument and dense micropunctation at centre, denser and with more marked punctures than head (in some specimens, however, punctation can be weaker); towards the sides, micropunctation progressivily less dense and well marked, being completely absent on the lateral margin; tegument with strong alveolar microsculpture throughout its surface. Posterior edge with a fine transverse line at centre (usually extending little beyond the second elytral stria). Hypomeral cavity with variable density of setae, from almost glabrous to with long yellowish setae covering the entire interior of the hypomeral cavity; external margin simple, without any trace of a tubercle. Metaventrite entirely glabrous; anterior region with tegument with distinct alveolar microsculpture; centre and posterior region with dense alveolar microsculpture obliterating weak micropunctation, or micropunctation completely absent.

LEGS. Profemora with tegument with strong alveolar microsculpture on their anterior half and with strong rivose microsculpture on their posterior half. Protibiae narrow and straight on their internal edge; at their apical seventh, external edge with two small acute teeth – the apical one slightly longer than the second. Mesofemora margined anteriorly only at their basal half; unmargined portion of anterior edge with row of very short setae; tegument with strong alveolar micosculpture. Metafemora margined only anteriorly, posterior margin absent; apical third of anterior edge covered by row of setae; tegument with strong alveolar microsculpture obliterating micropunctation, which is almost imperceptible; base without coarse elongate punctation. Metatarsomeres II and V subequal in length and longer than the others; metatarsomere IV shorter than the others.

ELYTRA. With only eight visible striae: first four or five striae clearly marked, finely carinulate, and without basal widening; fifth to seventh striae progressivily more effaced and interrupted, the seventh occasionally vestigial; eighth stria vestigial, present only as a weak mark parallel to the apical half of epipleura; all striae lack their carinulae at the apex of elytra, where they are marked only by

microsculpture or are completely indistinct; humeral carina absent. Tegument of interstriae with strong alveolar microsculpture obliterating micropunctation, which is almost imperceptible.

ABDOMEN. Ventrite VI smooth at centre and with weak rivose microsculpture on the sides. Lateral foveae absent in both sexes. Pygidium with tegument with strong alveolar microsculpture and effaced, frequently imperceptible micropunctation; of equivalent length in both sexes.

AEDEAGUS. Parameres about half as long as phallobase, and slightly asymmetrical: left paramere laterally flat or concave, right paramere convex (difference more easialy seen from ventral view). In lateral view, parameres with strong ventral keel and with short notch close to base ( Fig. 17D ).

SEXUAL DIMORPHISM. Males: Protibial spur broad and bifid, with external projection spiniform, straight, and long, and internal projection bent and widened ( Fig. 15D ). Ventrite VI strongly narrowed at middle; ventrite V usually without medial flange on its posterior edge or only slightly indicated over anterior edge of ventrite VI. Females: Protibial spur spiniform. Ventrite VI broad at middle and with anterior edge only slightly covered by weak flange of posterior edge of ventrite V.

Measurements

Males (N = 16). TL: AV: 7.0 ± 0.36; MX: 7.4; MN: 6.5. EL: AV: 4.9 ± 0.39; MX: 5.7; MN: 4.3. PL: AV: 2.1 ± 0.17; MX: 2.5; MN: 1.9. PW: AV: 4.1 ± 0.26; MX: 4.5; MN: 3.7. PgL: AV: 1.4 ± 0.09; MX: 1.5; MN: 1.3. PgW: AV: 2.1 ± 0.13; MX: 2.3; MN: 1.9.

Females (N = 21). TL: AV: 7.2 ± 0.50; MX: 8.2; MN: 6.6. EW: AV: 4.9 ± 0.23; MX: 5.3; MN: 4.6. PL: AV: 2.1 ± 0.14; MX: 2.3; MN: 1.9. PW: AV: 4.1 ± 0.19; MX: 4.5; MN: 3.9. PgL: AV: 1.4 ± 0.08; MX: 1.5; MN: 1.3. PgW: AV: 2.1 ± 0.11; MX: 2.3; MN: 1.9.

Geographical distribution

Cloud forests of western Amazonia and slopes of the Andes in Ecuador and Peru.

Ecoregions

Napo Moist Forests , Cordillera Oriental Montane Forest, Marañón Dry Forest, Ucayali Moist Forests, Peruvian Yungas.

Collecting sites ( Fig. 24 )

ECUADOR. Orellana: Onkone Gare Camp. Napo. Tungurahua: Baños de Agua Santa, Río Negro. Pastaza: Pastaza (Puyo: Llandia), Mera. Morona Santiago: Untsuante. Zamora Chinchipe: Cordillera del Cóndor.

PERU. Amazonas. San Martín: Rioja (Yuracyacu), Mariscal Cáceres (Juanjuí). Huánuco: Leoncio Prado (Rupa-Rupa: Tingo María). Junín: Chanchamayo (San Ramón). Cuzco: La Convención (Echarate).

Intraspecific variation and taxonomic discussion

It is interesting to note that many specimens of S. genieri sp. nov. studied for this work bear labels identifying them as S. candezei . It is not surprising that both species have been confused for such a long time, since they, together with S. foveiventris , share several characteristics not found in any other Sylvicanthon . The most obvious of them is the presence of only two protibial teeth ( Fig. 11J ), while all the other species have three, but we can list also protibiae without internal expansion (absent also in the aequinoctialis subgroup and in the majority of the species of the bridarollii group) and elytral striae very fine and progressively less marked from the elytral suture. Furthermore, among Sylvicanthon , it is only in S. genieri sp. nov. and S. candezei that the hypomeral tubercle is completely absent, fact that leads these two species to key out at the same couplet in all the identification keys published so far ( Schmidt 1922; Balthasar 1939; Pereira & Martínez 1956; Martínez et al. 1964; Vulcano & Pereira 1967).

Even so, also based on identification labels, it is possible to see that with the increasing accumulation of material collected during the 1990s and 2000s, some entomologists were able to recognize the true identity of S. genieri sp. nov. as a distinct species. From 1998 on, François Génier identified some specimens of this species as ‘ Sylvicanthon sp. nov.’, whereas Trond Tarsen, in 2012, determined others as ‘ Sylvicanthon sp. ≠ candezei ’. In fact, even at first sight, S. genieri sp. nov. and S. candezei are easily differentiated by colouration: the former species has a bright coppery colouration (occasionally, with greenish reflexions on head) ( Fig. 27A ), while S. candezei is entirely dark green ( Fig. 25A ). The degree of punctation and microsculpture throughout the body also varies between the two species: in S. candezei , punctures at the centre of pronotum are strong and easily seen, little obliterated by microsculpture, while punctures are very weak in S. genieri sp. nov. and usually almost completely obliterated by the strong microsculpture at the centre of the pronotum. On the elytra and pygidium, the microsculpture is always very strong and punctures are weak or even absent in S. genieri sp. nov., whereas, although not as well marked as at the centre of pronotum, they are still completely visible in S. candezei .

A difference is also seen on the ventrites: in males of S. candezei , ventrite five shows a weak medial flange over the anterior margin of ventrite six ( Fig. 14D ), while in females this flange is much stronger, giving a false impression of a medial narrowing to ventrite six (characteristic usually associated to males in Sylvicanthon ) ( Fig. 14E ). In S. genieri sp. nov., on the other hand, males usually lack any trace of a flange on ventrite five or, at most, it is very weak, almost imperceptible, while in females this flange is not very strong either, being similar to the one present in males of S. candezei , which does not obscure the true broad, typically female nature of ventrite six. Lastly, the clearest morphological difference between these two species lies on the shape of the parameres: in S. candezei , they have a deep ventral notch, dividing the parameres into two regions (the anterior one subrectangular) ( Fig. 17C ), and lack a ventral keel, whereas S. genieri sp. nov. presents a much shorter notch (which does not divide the parameres into two distinct parts) and has a strong ventral keel ( Fig. 17D ).

From S. foveiventris , the third species of the candezei subgroup, S. genieri sp. nov. is distinct in presenting a strong alveolar microsculpture on the pronotum, elytra and pygidium, whilst the former species does not possess any sign of microsculpture, with tegument marked only by micropunctation. The absence of paired foveae on the female ventrites (present in three pairs in S. foveiventris , Figs 14B , 16D ), the paramere shape (with a ventral keel and a notch in S. genieri sp. nov., Fig. 17D ; and simple in S. foveiventris , Fig. 17B ), and colouration also distinguish both species. On the other hand, S. genieri sp. nov. and S. foveiventris are similar by the presence of a weak (rather than strong) medial flange of ventrite five of females (although this structure is even less developed in S. foveiventris , Fig. 14B , than in S. genieri sp. nov.).

Limited morphological variation was observed in S. genieri sp. nov., none of it related to distribution. The few variable characteristics are related to the micropunctation of the pronotum and the elytra (which can be clearly present or almost entirely effaced and imperceptible; on the elytra, in particular, sometimes it is completely absent), the pilosity of the hypomeral cavity (which can be entirely glabrous or present several yellowish setae all over its surface), and the colouration (which can vary from a very dark coppery with little sheen to light reddish with greenish reflections on the head, with the majority of the specimens being in an intermediate position between these two extremes, with a bright coppery colouration without any trace of greenish reflexions). The colouration of the meso- and metafemur also varies, with one end of the variation spectrum being light yellow and the other dark brown. Teneral specimens are entirely yellow.

Comments

Judging from the cited locality, specimens identified as S. candezei by Carvajal et al. (2011) from Ecuador are probably S. genieri sp. nov.. In addition, having studied some specimens of the species that Celi et al. (2004) called “S ylvicanthon sp. 2”, it was possible to see that this species is actually S. genieri sp. nov. (see discussion about this and other identifications by Celi et al. on the comments under S. edmondsi sp. nov.).

Natural history

Based on the label information, it is possible to say that S. genieri sp. nov. is mainly collected with pitfall traps baited with human faeces. A male was collected in La Convención (Cuzcu, Peru) with a light trap and a female was found beneath the leaf litter in Onkone Gare Camp (Orellana, Ecuador). Living on the Amazon slopes of the Andes, S. genieri sp. nov. is in the candezei group the species which occurs at the highest elevation, having been recorded mostly in altitudes ranging from 805 to 1685 m; just one specimen was caught at 200 m in the Onkone Gare Camp. This species was collected throughout the year (January, March, April, May, July, August, September and October), but July (125) and January (22) concentrated the record of the vast majority of the 202 specimens studied.

Figures

Fig. 6. Head, dorsal view. A. Sylvicanthon enkerlini (Martínez et al., 1964) comb. nov. B. S. candezei (Harold, 1869). C. S. genieri sp. nov. D. S. proseni (Martínez, 1949) stat. et comb. nov. E. S. seag sp. nov. F. S. securus (Schmidt, 1920) comb. nov. G. S. furvus (Schmidt, 1920).

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. 17. Aedeagus morphological diversity in Sylvicanthon. A. S. enkerlini (Martínez et al., 1964) comb. nov. B. S. foveiventris (Schmidt, 1920). C. S. candezei (Harold, 1869) (arrows points to the ventral notch). D. S. genieri sp. nov. (idem). E. S. aequinoctialis (Harold, 1868). F. S. proseni (Martínez, 1949) stat. et comb. nov.

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. 24. Distribution of the three species of the candezei subgroup.

Fig. 27. Sylvicanthon genieri sp. nov. A. Dorsal view. B. Ventral view.

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. 25. Sylvicanthon candezei (Harold, 1869). A. Dorsal view. B.Ventral view.

Fig. 14. A. Ventral view of the anterior region of the pterothorax of S. proseni (Martínez, 1949) stat. et comb. nov. Abbreviations: MsEs = mesoventrite; SEs = meso-metaventrite suture; MsC = mesocoxa; RAMtEs = anterior region of metaventrite. B–E. Sexual dimorphism in the abdominal ventrites of Sylvicanthon Halffter & Martínez, 1977. B. S. foveiventris (Schmidt, 1920), ♀. Note the three lateral foveae between ventrites I–II, II–III, and III–IV and the broad ventrite VI at the middle, without emargination on its posterior edge and without its anterior edge covered by a medial flange of ventrite V. C. S. proseni stat. et comb. nov., ♀, ventrite VI broad. D. S. candezei (Harold, 1869), ♂. Note how the ventrite VI is narrow at the middle due to the emargination on its porterior edge and to the medial flage of the posterior edge of the ventrite V. E. S. candezei (Harold, 1869), ♀. Despite its posterior edge not being emarginate, its anterior edge is covered by a well-developed medial flange of ventrite V, which gives a narrowed aspect to the centre of ventrite VI.

Fig. 16. Abdominal lateral foveae of females (arrows). A. Sylvicanthon furvus (Schmidt, 1920). B. S. mayri sp. nov. C. S. obscurus (Schmidt, 1920). D. S. foveiventris (Schmidt, 1920).