Biolus granulatus (Fabricius) Lumen, Ryan & Kamdńskd, Marcdn J., 2024

Lumen, Ryan & Kamdńskd, Marcdn J., 2024, evdsdon of the darkldng beetle genus Eurynotus (Blaptdnae: Platynotdnd % and new records of ovovdvdpary dn ºenebrdonddae, Zoological Journal of the Linnean Society 201 (201), pp. 358-386 : 378

6. Biolus granulatus (Fabricius) comb. nov.

( Figs 1A , 2F, I , 5A , 7F , 8F )

Helops granulatus Fabricius 1787: 214 .

Blaps granulata ( Fabricius 1787).– Fabricius 1792: 107.

Eurynotus granulatus ( Fabricius 1787).– Gebien 1906: 230, 1938: 414; Koch 1954a: 280; 1954b: 2; 1956: 25; Banaszkiewicz 2006: 63; Kamiński 2016: 235.

= Eurynotus norrisi Mulsant and Rey 1854a: 27 syn. nov. – Mulsant and Rey 1854b: 171; Lacordaire 1859: 239; Gebien 1938: 414; Koch 1954a: 277; 1954b: 2; Kamiński 2016: 235.

Diagnosis: Can be difficult to differentiate from other species, but is most similar to B. omeri (e.g. both have simple punctation of abdominal ventrite V, and are similar in size, with tubercles/ raised callosities on elytral disc). The two can be differentiated based on setation of the tarsi ( B. norrisi golden; B. omeri black/ dark brown), the sclerite within the bursa copulatrix (present in B. granulatus ; absent in B. omeri ), and characterization of the dorsal surface ( B. norrisi blunter/shorter and more crowded tubercles; while B. omeri has sharper/taller and more widely spaced tuberculation with a slightly more lustrous surface). Insofar as other species are concerned, the abdominal ventrite V punctation (simple) separates it from B. analisetosus , B. asperipennis , B. graafi , and B. sauroides (with two or more types of punctation). The tubercles/raised callosities present on the elytral disc further separate it from B. asperipennis (lacking discal tubercles). The presence of a sclerite in the bursa copulatrix sep - arates this species from all but B. analisetosus (which, in contrast to B. granulatus ’ axe -shaped sclerite, has a flatter, chevron -shaped sclerite).

Redescription: Length 14–16 mm; width 7–9 mm. Head: Coarsely punctate, punctures closely spaced, separated by ≤1 puncture diameter to confluent. Mentum with apical notch. Eye with impressed sulcus around margin. Prothorax: Pronotum coarsely punctate, punctures separated by ≤1

puncture diameter to confluent. Hypomeron rugose, with large and shallow punctures. Pterothorax: Elytron not costate, with macrotubercles and with distinct or faint microtuberculation in interstices on disc. At most, a few widely spaced, small microtubercles in basal region of epipleura. Abdomen: Punctate. Abdominal ventrite V punctures separated by ≥1 puncture diameter without larger, setigerous punctures inter - mixed. Females without median apical notch. Legs: Male and Female legs unmodified. Tarsi with golden setae. Males: pro/meso tarsi robust with large, ventral, tomentose pads. Male terminalia: Parameres weakly tapering with tips sharply tapering and directed medially. Female terminalia: Bursa copulatrix with accessory pouch and axe -shaped sclerite near spermathecal opening.

Material examined: Helops granulatus : Lectotype, here des - ignated ( ZMUK): QR code label ‘ZMUKFabricius 004512’. Note: As the original descriptions lack information defining the number or specificity of specimens used, a specimen from the presumed syntype series is here designated as lectotype to fix the taxonomic status of the species.

Eurynotus norrisi Lectotype (MNHN): ‘ Eurynotus norrisii buq? spinola in litt. cap.bon.sp./ small blue circular label: Eurynotus norrisi Mls Cap. B. Sp. 2 T?/ Type Mulsant/ Norrisii/ Red label: Type’.

(See Supporting Information, Appendix S1 for additional ma - terial examined.)

Note: As the original descriptions lack information defining the number or specificity of specimens used, a specimen from the presumed syntype series is here designated as lectotype to fix the taxonomic status of the species.

Distribution: South Africa (Fig. 9).

Figures

Figure 2. Biolus stat.nov.species.A, Biolus analisetosus (Koch) comb. nov. Holotype; B, B.asperipennis (Mulsant and Rey) stat.res., comb.nov. Lectotype; C, B. brunoi sp. nov. Holotype; D, B. denticosta (Mulsant and Rey) comb. nov. Lectotype; E, B. graafi (Koch) comb. nov. Holotype; F,B. granulatus (Fabricius) comb. nov. Lectotype; G, B. omeri (Koch) comb.nov.Holotype; H, B. sauroides (Koch) comb. nov. Holotype; I, Eurynotus norrisi Mulsant and Rey syn. nov. of B. granulatus, Lectotype.

Figure 5. Phylogeny of the subtribe Eurynotina based on morphology. The most parsimonious cladogram obtained in TNT analysis

Figure 7. Phylogeny of morphology, ovovivipary, and female reproductive structures of Eurynotina.A, Eurynotus barbosai, 1st instar larva dissected from bursa copulatrix. B, Biolus omeri, genital tubes. C, B. brunoi, spermatheca and ovipositor.D, E. capensis, genital tubes of two individuals.E, B.analisetosus, sclerite and egg dissected from bursa copulatrix. F, B. granulatus, sclerite from bursa copulatrix.G, E. asperatus, paired sclerites on distal part of bursa copulatrix. H, Oncotus cedrimontis, bursa copulatrix. Abbreviations: c1–c4, lobes of coxities. Numbers in parentheses represent characters and states used in analysis.

Figure 8. Aedeagal morphology of Eurynotina species.A, Biolus analisetosus; B, B.asperipennis; C, B. brunoi sp. nov.; D, B. denticosta; E, B. graafi; F, B. granulatus; G, B. omeri; H, B. sauroides; I, Eurynotus asperatus; J, E.barbosai; K, E. capensis; L, E. graniderma; M, Capidium tuberculatum; N, Oncotus cedrimontis; O, Heteropsectropus amaroides; P, Schyzoschelus africanus. Parameres highlighted red. Scale = 1mm.

Figure1. Ovoviviparyin Tenebrionidae and its geographical distribution.A, sympatricPlatynotini species displaying different reproductionstrategies: Biolusgranulatus–ovoviviparous,Crypticanus sp.–oviparous.B, female genital tract ofBiolus brunoi sp.nov.with developed 1stinstar larva.Scale