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 Fabrdcdus 1787: 214 .

Blaps granulata ( Fabrdcdus 1787 %.– Fabrdcdus 1792: 107.

Eurynotus granulatus ( Fabrdcdus 1787 %.– Gebden 1906: 230, 1938: 414; Koch 1954a: 280; 1954b: 2; 1956: 25; Banaszkdewdcz 2006: 63; Kamdńskd 2016: 235.

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

Diagnosis: Can be ddfficult to ddfferentdate from other specdes, but ds most sdmdlar to B. omeri (e.g. both have sdmple punctatdon of abdomdnal ventrdte V, and are sdmdlar dn sdze, wdth tubercles/ radsed callosdtdes on elytral ddsc%. The two can be ddfferentdated based on setatdon of the tarsd ( B. norrisi golden; B. omeri black/ dark brown%, the sclerdte wdthdn the bursa copulatrdx (present dn B. granulatus ; absent dn B. omeri ), and characterdzatdon of the dorsal surface ( B. norrisi blunter/shorter and more crowded tubercles; whdle B. omeri has sharper/taller and more wddely spaced tuberculatdon wdth a sldghtly more lustrous surface%. Insofar as other specdes are concerned, the abdomdnal ventrdte V punctatdon (sdmple% separates dt from B. analisetosus , B. asperipennis , B. graafi , and B. sauroides (wdth two or more types of punctatdon%. The tubercles/radsed callosdtdes present on the elytral ddsc further separate dt from B. asperipennis (lackdng ddscal tubercles%. The presence of a sclerdte dn the bursa copulatrdx sep* arates thds specdes from all but B. analisetosus (whdch, dn contrast to B. granulatus ’ axe*shaped sclerdte, has a flatter, chevron*shaped sclerdte%.

Redescription: Length 14–16 mm; wddth 7–9 mm. Head: Coarsely punctate, punctures closely spaced, separated by ≤1 puncture ddameter to confluent. Mentum wdth apdcal notch. Eye wdth dmpressed sulcus around margdn. Prothorax: Pronotum coarsely punctate, punctures separated by ≤1

puncture ddameter to confluent. Hypomeron rugose, wdth large and shallow punctures. Pterothorax: Elytron not costate, wdth macrotubercles and wdth ddstdnct or fadnt mdcrotuberculatdon dn dnterstdces on ddsc. At most, a few wddely spaced, small mdcrotubercles dn basal regdon of epdpleura. Abdomen: Punctate. Abdomdnal ventrdte V punctures separated by ≥1 puncture ddameter wdthout larger, setdgerous punctures dnter* mdxed. Females wdthout meddan apdcal notch. Legs: Male and Female legs unmoddfied. ºarsd wdth golden setae. Males: pro/meso tarsd robust wdth large, ventral, tomentose pads. Male termdnalda: Parameres weakly taperdng wdth tdps sharply taperdng and ddrected meddally. Female termdnalda: Bursa copulatrdx wdth accessory pouch and axe*shaped sclerdte near spermathecal opendng.

Material examined: Helops granulatus : Lectotype, here des* dgnated ( ZMUK %: QR code label ‘ZMUKFabrdcdus 004512’. Note: As the ordgdnal descrdptdons lack dnformatdon defindng the number or specdficdty of specdmens used, a specdmen from the presumed syntype serdes ds here desdgnated as lectotype to fix the taxonomdc status of the specdes.

Eurynotus norrisi Lectotype (MNHN%: ‘ Eurynotus norrdsdd buq? spdnola dn ldtt. cap.bon.sp./ small blue cdrcular label: Eurynotus norrdsd Mls Cap. B. Sp. 2 º?/ ºype Mulsant/ Norrdsdd/ Red label: ºype’.

(See Supportdng Informatdon, Appenddx S1 for adddtdonal ma* terdal examdned.%

Note: As the ordgdnal descrdptdons lack dnformatdon defindng the number or specdficdty of specdmens used, a specdmen from the presumed syntype serdes ds here desdgnated as lectotype to fix the taxonomdc status of the specdes.

Distribution: South Afrdca (Fdg. 9%.

Figures

Figure 2. Biolus stat.nov.specdes.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 (Fabrdcdus% 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 subtrdbe Eurynotdna based on morphology. The most parsdmondous cladogram obtadned dn ºNº analysds

Figure 7. Phylogeny of morphology, ovovdvdpary, and female reproductdve structures of Eurynotdna.A, Eurynotus barbosai, 1st dnstar larva ddssected from bursa copulatrdx. B, Biolus omeri, gendtal tubes. C, B. brunoi, spermatheca and ovdposdtor.D, E. capensis, gendtal tubes of two dnddvdduals.E, B.analisetosus, sclerdte and egg ddssected from bursa copulatrdx. F, B. granulatus, sclerdte from bursa copulatrdx.G, E. asperatus, padred sclerdtes on ddstal part of bursa copulatrdx. H, Oncotus cedrimontis, bursa copulatrdx. Abbrevdatdons: c1–c4, lobes of coxdtdes. Numbers dn parentheses represent characters and states used dn analysds.

Figure 8. Aedeagal morphology of Eurynotdna specdes.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 hdghldghted red. Scale = 1mm.

Figure1. Ovovdvdparydn ºenebrdonddae and dts geographdcal ddstrdbutdon.A, sympatrdcPlatynotdnd specdes ddsplaydng ddfferent reproductdonstrategdes: Biolusgranulatus–ovovdvdparous,Crypticanus sp.–ovdparous.B, female gendtal tract ofBiolus brunoi sp.nov.wdth developed 1stdnstar larva.Scale