Sylvicanthon furvus ( Schmidt, 1920 )

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 : 153-159

Sylvicanthon furvus ( Schmidt, 1920)

Figs 6G , 11E , 13E , 15N , 16A , 19C , 20 , 40C , 41 , 43 A–B

Canthon furvus Schmidt, 1920: 130–131 , 133.

Canthon furvus – Schmidt 1922: 64, 75. — Balthasar 1939: 188; 1941: 341; 1951: 326. — Halffter &

Martínez 1977: 63. — Krajcik 2012: 63. Canthon furvum – Blackwelder 1944: 199. Glaphyrocanthon (Glaphyrocanthon) furvus – Pereira & Martínez 1956: 126, 129. — Martínez et al.

1964: 5–6, 8, 10, 14, 20–21. — Vulcano & Pereira 1964: 662. Sylvicanthon furvus – Halffter & Martínez 1977: 63. — Ratcliffe et al. 2015: 196.

Etymology

From the Latin ‘ furvus ’, meaning ‘black’ or ‘dark’, probable reference to the consistently dark colouration of this species.

Material examined

Lectotype (here designated)

PERU: Ƌ, Cuzco, Quispicanchi, Marcapata. Former type locality cited by Schmidt (1920): “ Peru, Bolivien ”,(“Typ.”,“Marcapata/Peru”,“Typus”,“ furvus / Type m. ”,“ furvus / A.Schm.”,“ Glaphyrocanthon / furvus / (Schm.) / P. Pereira det. 60 ”, “9663 / E92 +”, “NHRS-JLKB / 000021102”, “ LECTOTYPE ♂ / Canthon / furvus Schm. / des. F. Z. Vaz-de-Mello, 20 13 ”) ( NHRS) ( Fig. 40 Ca–b).

Paralectotypes (4 ♂♂, 5 ♀♀)

BOLIVIA: 1 ♂, (“9662 / E92 +”, “ PARALECTOTYPE / ♂ / Canthon / furvus Schm. / des. F. Z. Vazde-Mello, 20 13 ” “Bolivia”, “ furvus ”) ( NHRS) ( Fig. 40 Cf); 1 ♂, (“ PARALECTOTYPE / ♂ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “ Canthon / furvus / A. Schmidt ”, “Typus”, “Coll. C. Felsche / Kauf 20, 1918”, “Bolivia”) ( SMTD) ( Fig. 40 Cc); 1 ♂, (“ PARALECTOTYPE / ♂ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “Coll. C. Felsche / Kauf 20, 1918”, “Bolivia”) ( SMTD); 1 ♀, (“ PARALECTOTYPE / ♀ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “Coll. C. Felsche / Kauf 20, 1918”, “Bolivia”) ( SMTD); 1 ♀, (“ PARALECTOTYPE / ♀ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “Coll. C. Felsche / Kauf 20, 1918”, “Bolivia”) ( SMTD); 1 ♀, (“ PARALECTOTYPE / ♀ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “Coll. C. Felsche / Kauf 20, 1918”, “Bolivia”) ( SMTD).

PERU: 1 ♀, (“ furvus / Schmidt”, “Marcapata / Peru”, “9664 / E92 +”, “ PARALECTOTYPE / ♀ / Canthon / furvus Schm. / des. F. Z. Vaz-de-Mello, 20 13 ”) ( NHRS); 1 ♀, (“ PARALECTOTYPE / ♀ / Canthon / furvus Schm. / des. F. Z. Vaz-de-Mello, 20 13 ”, “ Glaphyrocanthon / furvus / (Schm.) / P. Pereira det. 60 ”, “Marcapata / Peru”, “9661 / E92 +”) ( NHRS) ( Fig. 40 Ce); 1 ♂, (“ PARALECTOTYPE / ♂ / Canthon / furvus / Schmidt / des. F. Z. Vaz-de-Mello, 20 14 ”, “ Canthon / furvus / A. Schmidt ”, “Peru”, “Gehr. W. Müller / Vermächt. 1909”) ( SMTD) ( Fig. 40 Cd).

Additional material (17 ♂♂, 8 ♀♀)

BOLIVIA: 1 ♂, no further data ( MNHN); 1 ♂, no further data [labelled as homeotype of “ Canthon candezei ” by P. Pereira, 1942] ( MZSP). – La Paz: 1 ♀, Larecaja, Guanay, 10 Nov. 2004, A.U. Peña leg. ( CEMT); 2 ♂♂, Murillo, Farinas ( MNHN); 2 ♂♂, Murillo, Zongo (“Songo”) ( MNHN); 1 ♂, same collecting data as for preceding ( MZSP); 2 ♂♂, 3 ♀♀, Nor Yungas, Coroico ( BMNH); 1 ♂, same collecting data as for preceding ( MZSP); 1 ♀, same collecting data as for preceding ( NMPC, ex. leg. Balthasar); 1 ♂, 1 ♀, Nor Yungas, Suapi ( MNHN); 1 ♂, “Yungas de La Paz”, 1000 m (“ 3,300 ft. ”) ( BMNH).

PERU: Cuzco: 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º33′54.41″ S, 73º05′36.85″ W, 26–29 Jan. 2010, C. Carranza and C. Rossi leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º33′51.29″ S, 73º05′37.01″ W, 1457 m, 18–21 Sep. 2010, pitfall, M. Alvarado and J. Peralta leg. ( MUSM); 1 ♂, La Convención, Echarate, Comunidad Campesina Santa Rosa, 12º34′14.23″ S, 73º05′41.87″ W, 1459 m, 14 Oct. 2009, C. Carranza and C. Rossi leg. ( MUSM); 1 ♂, 1 ♀, Quispicanchi, Marcapata ( NMPC, ex. coll. Balthasar). – Junín: 1 ♂, Chanchamayo, O. Schunke leg. ( MNHN).

No data: 1 ♀ [small green label] ( MNHN).

Description

COLOURATION. Very variable and iridescent. In general, head with greenish or bluish sheen. Pronotum usually purple; occasionally, with greenish sheen. Elytra purple or blue. Meso- and metafemora reddishbrown. Metaventrite with strong green shine. Pygidium of same colour as elytra.

HEAD. Tegument slightly shiny, with strong alveolar microsculpture obliterating dense surficial micropunctation; micropunctation almost imperceptible or even absent throughout outer edge of head. Clypeus with two apical teeth obtuse and only slightly separated from one another ( Fig. 6F ); with single transverse row of setae covering base of both teeth. Genae with weak denticle immediately behind clypeal-genal juncture. Posterior edge of head with fine incontinuous line between eyes, or completely unmargined.

THORAX. Pronotum with shiny tegument, and dense and well-marked micropunctation at centre; towards sides, micropunctation progressively less dense and well marked, sometimes absent on lateral edge; tegument among micropunctation completely smooth, microsculpture limited to very narrow strip of tegument on anterior edge of pronotum and anterolateral angles. Posterior edge with evident fine transverse line at centre (usually extending almost to third elytral striae). Hypomeral cavity with long yellowish setae at centre; external edge with weak tubercle. Metaventrite glabrous at centre; sides with few and sparse setae close to anterior margin of metacoxae; anterior region of metaventrite with distinct rivose microsculpture; centre and posterior region with dense micropunctation and very effaced, almost imperceptible microsculpture.

LEGS. Ventral surface of all femora and tibiae shiny. Profemora with tegument with sparse micropunctation and without microsculpture at anterior half and with strong rivose microsculpture at posterior half and on anterior margin. Protibiae narrow and with distinct expansion on internal edge; at apical third, external edge with three small acute teeth; the two most apical ones subequal in length and larger than basal ( Fig. 11E ). Mesofemora margined anteriorly only at basal half; unmargined portion of anterior edge with row of short setae; posterior margin absent; tegument with strong rivose microsculpture. Metafemora margined only anteriorly, posterior margin absent; apical third of anterior edge covered by row of setae; with strong coarse elongate punctation at base and with sparse micropunctation on rest of tegument, which is covered by rivose microsculpture ( Fig. 13E ). Metatarsomeres II and V subequal in length and longer than the others; metatarsomere IV shorter than others.

ELYTRA. With only seven narrow visible striae: the first three or four striae well marked, finely carinulate, and widened at base; fifth to seventh striae progressively more effaced and interrupted; all striae lack carinulae before reaching apex of elytra, marked only by microsculpture or completely indistinct; humeral carina absent. Tegument of interstriae matte, with strong alveolar microsculpture throughout elytra surface; micropunctation totally oblitered by microsculpture and almost imperceptible.

ABDOMEN. Tegument of ventrites I–V with strong rivose microsculpture; ventrite VI smooth at centre and with weak rivose microsculpture on sides. Pygidium with shiny tegument; at centre, with diffuse microsculpture and dense micropunctation; on sides, with evident rivose microsculpture.

AEDEAGUS. Parameres longer than half-length of phallobase and with no noticeable asymmetry, both external faces flat. In lateral view, parameres with apices widely bifurcate, with inferior branch distinctly projected and divergent from superior branch; without ventral keel or notch ( Fig. 19C ).

SEXUAL DIMORPHISM. Males: Protibial spur broad and bifid, with external projection spiniform and not much longer than internal projection, which is bent and widened ( Fig. 15N ). Abdomen without lateral foveae. Ventrite VI strongly narrowed at centre, with posterior posterior edge emarginate; anterior edge covered by weak medial flange of posterior edge of ventrite V. Pygidium very long (length between 1.4 and 1.1 mm) and convex; apical margin much wider than lateral margins. Females: Protibial spur simple, spiniform. Abdomen with three pairs of lateral transverse foveae between ventrites I–II, II–III, and III–IV, respectively; foveae not margined by row of long setae ( Fig. 16A ). Ventrite VI very broad at centre, posterior edge straight, without emargination; anterior edge subtly covered by weak medial flange of posterior edge of ventrite V. Pygidium shorter (between 1.1 and 0.9 mm) and flat; apical margin of pygidium only slightly wider than lateral margin.

Measurements

Males (N = 11). TL: AV: 7 ± 0.53; MX: 7.9; MN: 6.2. EW: AV: 5.2 ± 0.29; MX: 5.8; MN: 4.7. PrL: AV: 2.4 ± 0.13; MX: 2.6; MN: 2.2. PrW: AV: 4.4 ± 0.21; MX: 4.8; MN: 4.1. PgL: AV: 1.2 ± 0.07; MX: 1.4; MN: 1.1. PgC: AV: 2.2 ± 0.12; MX: 2.4; MN: 2.

Females (N = 6). TL: AV: 7.4 ± 0.40; MX: 8.1; MN: 7.0. EW: AV: 5.4 ± 0.15; MX: 5.5; MN: 5.1. PrL: AV: 2.4 ± 0.12; MX: 2.5; MN: 2.2. PrW: AV: 4.6 ± 0.12; MX: 4.8; MN: 4.5. PgL: AV: 1.0 ± 0.09; MX: 1.1; MN: 0.9. PgW: AV: 2.3 ± 0.15; MX: 2.4; MN: 2.0.

Geographical distribution

Eastern slopes of the Andes in Peru and Bolivia.

Ecoregions

Peruvian Yungas, Bolivian Yungas. Eastern slopes of the Andes in Peru and Bolivia.

Collecting sites ( Fig. 41 )

PERU. Junín: Chanchamayo. Cuzco: La Convención (Echarate), Quispicanchi (Marcapata).

BOLIVIA. La Paz: Larecaja (Guanay), Murillo (Farinas, Zongo), Nor Yungas (Coroico, Suapi).

Intraspecific variation and taxonomic discussion

The furvus subgroup represents the assemblage with the most complex taxonomy in Sylvicanthon . One of the reasons for this is the small number of specimens found in collections: while we could examine 2060 specimens of the species of the bridarollii subgroup, the other subgroup with an intricate taxonomy, we had in front of us only 53 specimens of the three rarest species of the furvus subgroup (i.e., excluding S. obscurus ). At least in part, this is a consequence of the small number of collections made in the species’ occurrence area (the slopes of the Andes in Bolivia and Peru for S. furvus , southern Amazonia for S. monnei sp. nov. and western Amazonia for S. mayri sp. nov.). Nevertheless, it is probable that these species should have natural low population densities, since even modern collections made with pitfall traps baited with dung did not yield a large number of individuals (e.g., specimens of S. monnei sp. nov. collected in the northern region of Mato Grosso and the holotype of S. mayri sp. nov. in Rio Branco, Acre). A second reason for such a complex taxonomy in the furvus subgroup is the tenuous morphological differences found between S. mayri sp. nov., S. furvus , and S. monnei sp. nov. (distinctions between S. obscurus and these three species were discussed anteriorly; see more details on Table 5).

The differences between S. furvus , S. mayri sp. nov. and S. monnei sp. nov. may be classified into two groups: those referring to the microsculpture and those referring to the shape of the parameres. Regarding the former, we observe that S. furvus possesses a much stronger microsculpture than S. mayri sp. nov. and S. monnei sp. nov. over the body. The head of the first species is covered by a dense, strongly impressed alveolar microsculpture, which makes the tegument more opaque, while the other two species have the head largely microsculptured, but this microsculpture is much weaker and, in some areas of the head (especially on frons), it can be totally absent, which makes the tegument much brighter. The meso- and metafemora of S. furvus are entirely covered by a strong rivose microsculpture and have micropunctation almost imperceptible, while these structures are smooth in almost the entirety of their tegument (except for the anterior region of the apical area, which has a rivose microsculpture) and show an evident micropunctation in both S. mayri sp. nov. and S. monnei sp. nov. 22 (the profemora also have rivose microsculpture more strongly marked in S. furvus than in the other two species, but this difference is less noticeable). The pygidium of S. furvus has microsculpture ranging from having a clear alveolation to a completely diffuse one, while the microsculpture in both S. mayri sp. nov. and S. monnei sp. nov. is absent in almost the entire tegument and may be present only at the base (and, in this latter case, it is more evident in S. monnei sp. nov. than in S. mayri sp. nov.). Finally, on the elytra rest the major differences between the three species: in S. furvus , the entire elytral tegument is covered by a strongly-marked alveolar microsculpture so that the tegument has a very weak, diffuse silky shine. In S. mayri sp. nov., the tegument presents a very diffuse microsculpture, without an evident alveolar pattern, and, consequently, is more intense bright. Lastly, in S. monnei sp. nov., the microsculpture is completely absent on the elytra, which gives the tegument a smooth appearance with a strong, lustrous shine, and micropunctation is present.

The differences in the shape of the parameres are determinant for the separation of both S. furvus and S. mayri sp. nov. from S. monnei sp. nov. The paramere apex of the first two species, which do not show any differences from each other, has a strong bifurcation, with two widely divergent branches, being the inferior branch well developed and with a strong excavation posteriorly to it ( Fig. 44 A–B). In S. monnei sp. nov., in contrast, the apical bifurcation is much weaker, and the inferior branch is short and straight ( Fig. 44 C–D).

Distinctions related to the coarse punctation at the base of the metafemora and to the female abdominal foveae approach S. furvus to S. monnei sp. nov. on the one side, and separate them from S. mayri sp. nov. on the other. The coarse punctures of metafemora are usually longer and more well impressed in the first two species ( Fig. 13C, E ), while they are finer and much weaker in S. mayri sp. nov. ( Fig. 13D ). Nonetheless, there is a variation in the intensity on which these punctures are marked in different individuals of a same species (and also in S. foveiventris and S. bridarollii , the other two species in which these punctures are present). It is important to bear in mind, however, that it is only possible to see this distinction at the populational level, not at the individual one. The real difference should rest, therefore, when we see populations as a whole (in average, punctures are more profound in S. monnei sp. nov. and S. furvus than in S. mayri sp. nov.). When we have larger series of S. furvus , S. monnei sp. nov. and, especially, S. mayri sp. nov., we will be able to test more firmly whether this observation is correct or not. As for the abdominal foveae, they are as profound and clearly marked in S. monnei sp. nov. as in S. furvus ( Fig. 16A ), while they are very shallow (particularly the one between ventrites III–IV) in the only two known females of S. mayri sp. nov. ( Fig. 16B ).

Along with the morphological differences listed so far, these three species are totally allopatric, without any known overlapping zone between their distributions ( Fig. 41 ). Sylvicanthon furvus is restricted to the cloud forests and the yungas on the eastern slopes of the Andes, in northern Bolivia and southern Peru. Sylvicanthon monnei sp. nov. is widely distributed in southern Amazonia, in the Brazilian states of Mato Grosso and Pará. Finally, S. mayri sp. nov. was recorded from four points very distant from one another in western Amazonia, in the Peruvian department of Madre de Díos, in the Brazilian states of Acre and Amazonas and in the Colombian state of Meta.

Comments

A male S. furvus collected in Bolivia and now housed at the MZSP is labelled “HOMEOTIPO” and “ Canthon / candezei / Har. / P. Pereira det. 942 ”. Since the term homeotype refers to an ordinary specimen compared to a type specimen ( Evenhuis 2008), it is curious to note that none of the known S. candezei syntypes bears any label by Padre Francisco Pereira, so indicating he did not have access to them. Besides, S. candezei and S. furvus are very distinct in terms of morphology and, therefore, it is improbable that such an experienced entomologist as Pe. Pereira could have confused them. On the other hand, he had access to the type specimens of S. furvus in 1960 (as stated by Pereira & Martínez 1960), and, hence, it might be possible that the homeotype label refers in fact to this latter name and has nothing to do with the 1942 identification label.

Natural history

Almost nothing is known on the biology of S. furvus . Most of the specimens examined, having likely been collected in the 19 th and early 20 th centuries, do not have bionomic data on their labels. The only information on the collecting methods comes from the label of a male collected in a pitfall trap (baited with human faeces?) in La Convención (Cuzco, Peru) in 2010. The only four specimens with data on the collecting month report January, September, October and November. Lastly, concerning the altitudinal range, specimens of S. furvus were found mostly between 1350 and 1850 m, being 450 m the lowest record (the female collected in Guanay, Bolivia). The type locality of S. furvus, Marcapata (Quispicanchi, Cuzco), is located around 3100 m a.s.l., which, if correct, would represent the highest altitudinal record for an American Deltochilini .

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. 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. 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.

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. 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).

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. 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. 40. Type material of three species of Sylvicanthon described by Schmidt (1920). A. S. secures (Schmidt, 1920) comb. nov. a. Lectotype. b. Lectotype’s labels. B. S. obscurus (Schmidt, 1920). a. Lectotype. b. Lectotype’s labels. c. Paralectotype 1. d. Paralectotype 3. e. Paralectotype 9. C. S. furvus (Schmidt, 1920). a. Lectotype. b. Lectotype’s labels. c. Paralectotype 4. d. Paralectotype 6. e. Paralectotype 3. f. Paralectotype 1.

Fig. 41. Distribution of Sylvicanthon securus (Schmidt, 1920) comb. nov. and the four species of the furvus subgroup.

Fig. 43. The three Amazonian species of the furvus subgroup. A, D. Sylvicanthon furvus (Schmidt, 1920). A. Dorsal view. D. Ventral view. B, E. S. monnei sp. nov. B. Dorsal view. E. Ventral view. C, F. S. mayri sp. nov. C. Dorsal view. F. Ventral view.

Fig. 44. Differences on the parameres of Sylvicanthon mayri sp. nov. and S. monnei sp. nov. (grayish zones represent original membraneous areas). A–B. S. mayri sp. nov. C–D. S. monnei sp. nov. Note that both branches of the apical bifurcation of the parameres of S. mayri sp. nov. are much more divergent than those of S. monnei sp. nov., which makes the internal angle between them more open in the first species (~110º) than in the second (~78º). In the same way, as the inferior branch is much more projected in S. mayri sp. nov. than in S. monnei sp. nov., the angle between the posterior region of that branch and the rest of the paramere is more open in S. monnei sp. nov. (~147º) than in S. mayri sp. nov. (~137º), species that seems to have a strong excavation at this point of the paramere (indicated by red arrow in A).