Streblopus punctatus ( Balthasar, 1938 )

Cupello, Mario, Ribeiro-Costa, Cibele S. & Vaz-De, Fernando Z., 2020, Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae), European Journal of Taxonomy 603, pp. 1-85 : 38-46

Streblopus punctatus ( Balthasar, 1938)

Figs 3–4 , 5 C–D, F, 6B, 7B, 8C–D, 12B, 13E–F, 14C–D, 15C–D, G–H, 16B–C, 17C–F, 18C–D, 19D–E, 20C–D, 21C, E, 23E–H, 25, 26B, 27B–D, 28

Streblopoides punctatus Balthasar, 1938: 216 .

Streblopoides punctatus ‒ Balthasar 1941: 346 ; 1951: 331. — Vulcano & Pereira 1964: 580–581. — Bezděk & Hájek 2011: 374 .

Streblopoides punctata ‒ Blackwelder 1944: 203 .

Streblopoides? punctatus ‒ Krajcik 2006: 163 .

Streblopus punctatus ‒ Halffter & Martínez 1966: 153–154 , 157–158, 162–165, figs 18–21. — Horgan 2005a: 131; 2005b: 609–610; 2006: 364; 2009: 3529, 3538. — Bezděk & Hájek 2011: 374. — Carvajal-López et al. 2011: 316. — Carvajal-López 2012: 195–197, 199, figs 1–2. — Krajcik 2012: 249. — Ratcliffe et al. 2015: 196. — Chamorro et al. 2018: 85, 98, fig. 8c; 2019: 234 ‒ 235, 313, fig. 50d. — Cupello & Vaz-de-Mello 2019: 168 ‒ 169.

Streblopus opatroides (error) ‒ Paulian 1939: 27 (in part: specimen from Oxapampa, Peru).

Material examined

Holotype

PERU? • ♀; dubious provenance [label indicates Peru, Piura, Huancabamba, 3000 m a.s.l., but this does not seem to be precise; see comments below] (“ punctatus / m.” [Balthasar’s handwriting], “ genotyp / stv. [?] punctatus / n. sp. / Dr. V. Balthasar det.” [Balthasar’s handwriting], “TYPUS”, “Huancabamba / N.Peru, 3000 m / H. Rolle.”, “ HOLOTYPE ♀ / Streblopoides punctatus / Balthasar, 1938 / Labelled by Cupello, 2019 ” [MC’s handwriting]); NMPC.

Other material (24 ♂♂, 16 ♀♀)

ECUADOR – Zamora-Chinchipe • 1 ♂, 1 ♀; Refugio de Vida Silvestre El Zarza, Campamento Las Peñas , “Parcela 5”; 1536 m a.s.l.; 9 Sep. 2012; William Chamorro leg.; pitfall baited with human faeces ;

CEMT • 1 ♀ (antennae, mouthparts, left posterior wing and genitalia dissected); Refúgio de Vida Silvestre El Zarza, Campamento Las Peñas (“C. Las Peñas”), “via a P4”; 1533 m a.s.l.; 11 Sep. 2011; William Chamorro leg.; pitfall baited with human faeces; CEMT • 1 ♂ (genitalia dissected), 1 ♀ (left posterior wing and genitalia dissected); Yantzaza; 1433 m a.s.l.; 11 Dec. 2010; T. Ghia leg.; pitfall baited with human faeces; CEMT.

PERU – Cuzco • 1 ♂; La Convención, Echarate, Comunidad Campesina Santa Rosa (“CC. Santa Rosa ”); 12°33′54.41″ S, 73°05′36.85″ W; 1747 m a.s.l.; 26–29 Jan. 2010; C. Carranza and C. Rossi leg.; MUSM . – Junín • 3 ♂♂; Chanchamayo, Chanchamayo, Nueva Italia , Santuario Nacional Pampa Hermosa ; 10°59′52.7″ S, 75°25′34.4″ W; 1737 m a.s.l.; 30 May 2011; D. Silva leg.; MUSM • 1 ♂ (genitalia dissected); same collection data as for preceding; ex W.D. Edmonds collection; TAMU • 2 ♂♂; Chanchamayo, Chanchamayo, Nueva Italia , Santuario Nacional Pampa Hermosa ; 10°59′51.8″ S, 75°25′35.9″ W; 1879 m a.s.l.; 30 May 2011; D. Silva leg.; MUSM • 1 ♂; same collection data as for preceding; ex W.D. Edmonds collection; TAMU • 1 ♀ (genitalia dissected); Chanchamayo, Chanchamayo, Nueva Italia , Santuario Nacional Pampa Hermosa ; 10°59′51.8″ S, 75°25′35.9″ W; 1940 m a.s.l.; 23–31 May [201?]; M. Alvarado leg.; Podocarpus forest; MUSM • 1 ♀; Chanchamayo, San Ramón ; 11°10′ S, 75°18′ W; 1500 m a.s.l.; 25 Aug. 2002; F.G. Horgan leg.; forest; pitfall with human faeces; MUSM • 1 ♂ (genitalia dissected); Satipo ; 600 m a.s.l.; 23 May–3 Jun. 2004; A. Santibañez leg.; CEMT • 1 ♂ (genitalia dissected), 1 ♀ (genitalia dissected); Satipo ; 1100 m a.s.l.; Dec. 2004; A. Santibañez leg.; CEMT • 1 ♀; Satipo ; 1100 m a.s.l.; Dec. 2004; A. Santibañez leg.; MNHN • 1 ♀ (genitalia dissected); Satipo, Río Venado ; Feb. 2011; CEMT • 2 ♂♂ (1 with genitalia dissected), 3 ♀♀ (1 with genitalia dissected); Satipo , “near [city of] Satipo”;> 1000 m a.s.l.; Dec. 2004; A. Santibañez leg.; yungas; trap with human faeces; ex W.D. Edmonds collection; TAMU . – Pasco • 2 ♂♂ (1 with genitalia dissected), 1 ♀; Oxapampa, Villa Rica, Bosque de Protección de San Matías-San Carlos ; 10°38′51″ S, 75°12′22″ W; 1556 m a.s.l.; 3–5 May 2012; L. Figueroa and V. Borda leg.; pitfall with dung; MUSM • 3 ♂♂, 3 ♀♀ (1 with genitalia dissected); Oxapampa, Villa Rica, Bosque de Protección de San Matías- San Carlos ; 10°38′44″ S, 75°12′37″ W; 1596 m a.s.l.; 6–10 Aug. 2012; P. Sanchez and E. Razuri leg.; MUSM • 1 ♂; Oxapampa, Villa Rica, Bosque de Protección de San Matías-San Carlos ; 10°36′27″ S, 75°12′18″ W; 1627 m a.s.l.; 3–5 May 2012; L. Figueroa and V. Borda leg.; pitfall with dung; MUSM • 4 ♂♂ (1 with genitalia dissected), 1 ♀; Oxapampa, Villa Rica, Bosque de Protección de San Matías-San Carlos ; 10°38′44″ S, 75°12′37″ W; 1702 m a.s.l.; 3–5 May 2012; L. Figueroa and V. Borda leg.; pitfall with dung; MUSM .

Erroneous record

PERU • 1 ♀; Lima; Sep. 1949; ex Martínez collection; CMNC .

Not studied (label information provided by François Génier (CMNC); 2 ♀♀)

PERU • 1 ♀; Huánuco, Leoncio Prado, La Divisoria (“Divisoria”); 1300 m a.s.l.; Aug. 1974; Bordon leg.; ex Martínez collection; CMNC .

Etymology

Derived from the Latin word for punctate ( Brown 1956), a probable reference to the fine umbilicate punctation seen on the pronotum of this species.

Redescription

MEASUREMENTS. Males (4 spec.): TL: AV 13.53, MX 14.4, MN 12.45. EW: AV 8.08, MX 8.64, MN 7.04. PW: AV 7.23, MX 8.06, MN 6.51. – Females (5 spec.): TL: AV 12.96, MX 13.95, MN 12.6. EW: AV 7.71, MX 8.0, MN 7.52. PW: AV 6.88, MX 7.13, MN 6.66.

COLOURATION. Antennal club usually with dark tegument and yellowish pubescence; some individuals with lighter tegument. Dorsum light brown; venter, including legs, dark brown to almost black. Some parts of body, especially elytra, pygidium and outer sides of metaventrite, sometimes with silky reddish sheen.

HEAD. Canthus short, with short penetration into eyes ( Fig. 5F ). Clypeal process a transverse carina, acuminate at apex ( Fig. 6C ). Antennae with 8 articles; first visible article of funicle little longer than combined length of second and third visible articles of funicle combined ( Fig. 7B ). Labrum with anterolateral regions not well projected and with external margin more or less straight; anterior labral margin not broad; epipharynx with short medial brush and median area almost entirely covered by dense, thick socketed setae ( Fig. 8 C–D).

THORAX. Pronotum with sparse umbilicate punctation at centre (approximately at area between eyes) ( Fig. 12B ); towards sides and posterior region, punctation progressively denser, but punctures never confluent; tegument between punctures with diffuse microsculpture. Prosternum, mesoventrite, mesanepisterna, metanepisterna and metaventrite entirely covered by sparse umbilicate punctation, particularly sparse and with smaller punctures on anterior lobe and posterior region of metaventrite; tegument between punctures lustrous, with extremely diffuse microsculpture ( Fig. 13 E–F).

LEGS. Trochantofemoral articulation not produced into spur in either sex. Mesocoxae with few sparse umbilicate punctures and tegument lustrous, covered by diffuse microsculpture. Mesofemora always glabrous on posterior edge of both sexes ( Fig. 17 C–F; sometimes with a few isolated setae in males); ventral surface covered by small and sparse umbilicate punctures at base and centre, and larger and denser punctures near apex and posterior edge; tegument between punctures lustrous and with extremely diffuse microsculpture, except near posterior edge, which is sexually dimorphic (see below). Mesotibiae with internal edge with a few erect setae, never forming well-defined longitudinal rows in either sex ( Fig. 18 C–D). Metafemora with sexually dimorphic brush of long setae on posterior edge (see below) ( Fig. 19 D–E). Metatibiae with ventral surface without umbilicate punctures (Ecuadorian specimens) or with at least some umbilicate punctures along entire tegument (Peruvian specimens); internal edge smooth, without row of sharp teeth at base in both sexes ( Fig. 20 C–D).

ELYTRA. Striae very fine and sometimes discontinuous, especially those on pseudepipleura and stria VII on disc. Tegument covered by small umbilicate punctures, each puncture glabrous or with short central seta; tegument between punctures with diffuse microsculpture (alveoli sometimes better delimited towards apices). Lateral carina (pseudepipleural carina) smooth, not interrupted by row of umbilicate punctures ( Fig. 21C ). Apices of elytra strongly deflected downwards; distal calluses strongly projecting ( Fig. 21E ).

TERMINALIA. Aedeagus: Right paramere with pronounced, broadly rounded apical projection with clear external spine ( Fig. 23 E–H);in dorsal view, parameres with external edges more or less straight ( Fig. 23E ); in lateral view, ventral side straight, with no week keel nor inclination ( Fig. 23 F–G). Internal sac with axial and subaxial sclerites distinct in shape: subaxial sclerite curved apically and surrounding axial sclerite, which is simply curved and shorter than subaxial sclerite ( Fig. 25D ). – Genital sclerite: Medial sclerotized plate (MSP) well sclerotized and with lateral arches more or less straight, not concave; posterior arms of MSP well developed and separated from one another by a deep ‘U-shaped’ emargination ( Fig. 26B ). – Spermatheca: Entirely slender and with a pair of apical hooks ( Fig. 27 B–C).

SEXUAL DIMORPHISM. Male: Clypeus slightly emarginated at apex ( Fig. 5C ). Metaventrite with excavation on posterior region slightly wider and deeper in males than in females ( Fig. 13E ). Profemora with anterior margin raised into a single spur ( Fig. 14C ). Protibiae as described above for genus; protibial spur very short, bent inwardly, acuminate ( Figs 15C, G , 16C ). Mesofemora near posterior edge with tegument covered by large umbilicate punctures surrounded by dense alveolar microsculpture ( Fig. 17C, E ). Metafemora with brush of long setae covering basal two-thirds of posterior edge; posterior edge not crenulate and with small spur bent inwards rising just after trochanter ( Fig. 19D ). – Female: Clypeus with apical emargination absent or only slightly marked ( Fig. 5D ). Metaventrite with excavation on posterior region slightly narrower and shallower in females than in males ( Fig. 13F ). Profemora with anterior margin simple, not raised into spur ( Fig. 14D ). Protibiae with two apical teeth subequal in length, contiguous at base and apart from basal tooth ( Fig. 15D, H ); all three lateral teeth (especially two apical ones) more or less directed forwards ( Fig. 15D, H ); protibial spur bent inwards apically, never bifurcated ( Fig. 15D, H ). Mesofemora near posterior edge with tegument covered by dense umbilicate punctation and tegument between punctures almost smooth, with very diffuse microsculpture (as rest of tegument of mesofemora) ( Fig. 17D, F ). Metafemora with brush of sparse, long setae covering approximately basal two-thirds of posterior edge; posterior edge crenulate along brush of setae and devoid of any spur ( Fig. 19E ).

Distribution

Sub-Andean areas in the Peruvian and Ecuadorian Yungas ( Fig. 4 ).

Ecoregions. Eastern Cordillera Real Montane Forest and Peruvian Yungas.

Collection sites. Ecuador – Zamora Chinchipe: Cordillera del Cóndor (Sector Colibrí), Refugio de Vida Silvestre ‘El Zarza’, Yantzaza. Peru – Huánuco: Leoncio Prado (La Divisoria); Pasco: Oxapampa (Villa Rica: Bosque de Protección de San Matías-San Carlos); Junín: Chanchamayo (Nueva Italia: Santuario Nacional Pampa Hermosa; San Ramón), Satipo (Satipo); Cuzco: La Convención (Echarate).

Intraspecific variation

Until fairly recently, only two specimens of S. punctatus were known to entomologists, the holotype female housed in the NMPC and another female deposited in the MNHN, both collected in Peru. Over the past two decades, however, with the growing interest for dung beetle studies among South American scholars and the subsequent increase in the number of collections in regions hitherto unexplored in terms of their scarabaeine fauna, that number has risen significantly. For this paper, we were able to examine 43 specimens (24 males and 19 females, including the holotype) deposited in five museums, but we know that some additional individuals are housed in at least two other collections in Ecuador (the Escuela Politécnica Nacional, Quito, and the Museo de Zoología de la Universidad Técnica Particular de Loja, Loja; Carvajal-López 2012; Chamorro et al. 2019). While still not a huge sample, these specimens reveal that S. punctatus shows some noticeable morphological variation, one of them being clearly geographical.

Firstly, two individuals, a Peruvian male and an Ecuadorian female, possess an antennal club with the tegument considerably lighter than on the other specimens examined, therefore departing from the typical dark tegument of S. punctatus and approaching the pattern observed in S. opatroides , whose antennal club has a yellowish tegument. A more significant variation, however, concerns the metatibial punctation: while all Ecuadorian individuals examined show the ventral surface of the metatibiae to be devoid of umbilicate punctures, all the Peruvian specimens studied by us, including the holotype, have at least some sparse umbilicate punctures along their entire ventral surface. However, since all the Peruvian individuals we had the chance to see came from much further south in Peru than its border with Ecuador, we cannot tell exactly how one form replaces the other over the distribution of S. punctatus , i.e., whether the transition between them is smooth or abrupt. Unfortunately, Carvajal-López (2012), who redescribed S. punctatus based on a female collected in the Cordillera del Cóndor, on the border between Ecuador and Peru, did not comment on what the tegument of the metatibiae of his specimen was like. As mentioned above for the colour variation observed in S. opatroides , future collections will give us more detail on how this variation occurs in nature and whether intermediate populations show intergradation between the two forms seen here. As for many other characters showing geographical variation in dung beetle species, nothing is known about the functional role of the umbilicate punctures of the metatibiae in this species, although it is reasonable to suppose they perform some kind of tactile or chemical function. Why this should vary geographically, however, remains totally unknown.

A final variation observed among the specimens is clearly non-genetic: two Peruvian males lack the protibial spur, but it is evident that this condition is due to wear, because one of them has a very short indication of a right spur, which is clearly reduced due to abrasion. As will be discussed on the following pages, in S. opatroides the protibial spur is never expressed in males, being restricted to females in that species.

Remarks

The information on the presence of S. punctatus in Cordillera del Cóndor (Zamora Chinchipe, Ecuador) is based on Carvajal-López (2012), who collected a female at 1446 m a.s.l. in that mountain range using a pitfall trap. Conversely, we question two other geographical records present in the literature. First, the type locality mentioned by Balthasar (1938) and indeed stated on the holotype’s label – namely, some place at about 3000 m a.s.l. in the province of Huancabamba (Piura, Peru) – is probably incorrect, since no other specimens are know to have been collected either above ~ 1950 m a.s.l. or in the Páramo ecosystem. Likewise, the record in Halffter & Martínez (1966) of S. punctatus from the outskirts of Lima (“arredores de Lima ”) is also most probably erroneous, since no other individuals are known from the west side of the Andes nor from the Sechura Desert ecoregion. Luis Figueroa (pers. comm. to MC, 23 Sep. 2018), the Coleoptera curator at the MUSM and specialist in Peruvian dung beetles, agrees that both records are inaccurate.

Ecology

Like its congeneric S. opatroides , almost nothing is known about the biology of S. punctatus . The specimens studied here were collected in both forest fragments and continuous forests at altitudes between 600 and 1940 m, and the holotype is labelled as having been collected at 3000 m (but see comments above about the accuracy of this record). The species is attracted to pitfall traps baited with human and pig excrement ( Horgan 2005a, 2005 b, 2006, 2009; Carvajal-López 2012; Chamorro et al. 2019; data from specimen labels), and adults, like those of S. opatroides , seem to be active throughout the year, having been recorded in the period “May–June” and all individual months from August to February ( Carvajal-López 2012; Chamorro et al. 2019; data from specimen labels). Horgan’s classification of S. punctatus as a roller ( Horgan 2005a) was probably based on inference from its morphology, not on actual observations of the food relocation behaviour of this species. Finally, as discussed above for S. opatroides , the wide eyes and dull colouration of this species lead us to hypothesize that flight activity in S. punctatus is nocturnal.

Figures

Fig. 3. Streblopus punctatus (Balthasar, 1938). A‒B. Holotype, ♀. A. Dorsal view. B. Attached labels. C‒D. Ordinary specimens, dorsal view. C. ♂. D. ♀.

Fig. 4. Distribution of Streblopus van Lansberge, 1874. Note how the two species are widely separated by the vast South American Dry Diagonal stretching between Amazonia and the Atlantic Forest. During several different times over the Neogene, forest corridors connected the two biomes and the populations of Streblopus could move from one to the other. When the forest retreated, the distribution of the genus became relict.

Fig. 5. Head. A‒D. Dorsal view. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Note how the emargination of the apex of the clypeus varies between the species and sexes, being greater in males of S. opatroides than in males of S. punctatus and females of both sexes. E‒F. Lateral view. E. S. opatroides. F. S. punctatus. Red arrow indicates the canthus. Observe that the canthus is much longer in S. opatroides than in S. punctatus.

Fig. 6. Head. A‒B. Clypeal process. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). C. Ventral view of head, S. opatroides.

Fig. 7. Antennae. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note the differences between the two species concerning the colour of the apical flagellomeres and the number and shape of the articles. In S. punctatus, the suppression of arthrogenesis between ancestral articles III and IV has resulted in a reduction of the total number of antennomeres from 9 to 8.

Fig. 8. Labrum. A‒B. Streblopus opatroides van Lansberge, 1874. A. Aboral surface (dorsal view). B. Oral surface (ventral view; epipharynx). C‒D. S. punctatus (Balthasar, 1938). C. Aboral surface. D. Oral surface. Note the many differences between the two species, particularly the much denser and shorter setation of the median area in S. punctatus, the much longer median brush of S. opatroides and the dissimilarities in the shape of the anterior epypharingeal margin. The black arrow shows the anterior margin of the labrum, while the red arrow indicates the anterolateral region. Abbreviations: afr = apical fringe of labrum; ampr = antero-median process of labral suspensorium; lf = lateral files; mbr = medial brush of labrum; pmpr = postero-median process of labral suspensorium; ppm = proplegmatium; ssma = socketed setae of median area; topr = tormal process of labral suspensorium.

Fig. 12. Pronotum. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note the differences in the umbilicate punctation and colour between the species.

Fig. 13. Venter. A‒B. Streblopus opatroides van Lansberge, 1874. A. Hypomeron. B. Mesoventrite and anterior region of metaventrite. C‒F. Metaventrite and abdomen. C‒D. S. opatroides. C. ♂. D. ♀. E‒F. S. punctatus (Balthasar, 1938). E. ♂. F. ♀. Note in particular the distinction between the species and sexes in the shape of the concavity of the posterior region of the metaventrite.

Fig. 17. Mesofemora. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. Note the sexual dimorphism in the presence of a long row of setae on the posterior edge of the male mesofemur. C‒F. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. E‒F. Detail of posterior region of metafemora. E. ♂. F. ♀. Note the strong alveolar microsculpture present near the posterior edge in males and the high concentration of umbilicate punctures in the same region in females.

Fig. 18. Mesotibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Observe that females of S. opatroides differ from those of S. punctatus by having a deep emargination at the apical edge of the mesotibiae (red arrow shows emargination).

Fig. 19. Posterior legs. A. Metacoxa of Streblopus opatroides van Lansberge, 1874. Arrow shows the lateral spur present in both sexes of both species. Note also that the spur covers the epipleura and possibly maintains the elytra pressed against the body, thus avoiding loss of water or the entry of sediment beneath the elytra in a similar way to that hypothesized for the metanepisternal tab of the Phanaeini and other tunneller groups (Edmonds 1972; Cupello & Vaz-de-Mello 2016). B‒E. Metafemora and metatrochanters. B‒C. S. opatroides. B. ♂. C. ♀. D‒E. S. punctatus (Balthasar, 1938). D. ♂. E. ♀. Note the strong sexual dimorphism and the many differences between the species.

Fig. 20. Posterior legs. A‒D. Metatibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Note the strong sexual dimorphism and the many differences between the species. E‒F. Metatarsi and apex of metatibia of S. opatroides. E. ♂. F. ♀. Observe how much shorter the male metatibial spur is in comparison to that of the female.

Fig. 21. Wings. A‒E. Elytra. A. Lateral view of elytra of Streblopus opatroides van Lansberge, 1874. Note the very wide pseudepiplera and narrow epipleura. B‒C. Pseudepipleural carina (indicated by red arrows). B. S. opatroides. C. S. punctatus (Balthasar, 1938). Observe that the pseudepipleural carina in B is completely interrupted by umbilicate punctures, whereas the pseudepipleural carina in C is almost completely smooth, with very tiny and isolated punctures. D‒E. Lateral view of apex of elytra. D. S. opatroides. E. S. punctatus. Note how the distal calluses (shown by blue arrows) of S. punctatus are much more strongly projecting than those of S. opatroides. F. Right hind wing of Streblopus opatroides. Red arrow shows the notch of the anal fold. Abbreviations: AA = anal anterior; AP = anal posterior; Cu = cubitus; J = jugal; MP = media posterior; MPa = accessory MP; r4 = cross-vein r4 (cross-vein connecting veins RA3+4 and RP2); RA = radius anterior; RP = radius posterior; ScA = subcosta anterior.

Fig. 23. Genital capsule. A‒D. Streblopus opatroides van Lansberge, 1874. A. Dorsal view. B‒C. Lateral view from slightly different angles (red arrow shows the ventral surface and its keel). D. Ventral view. E‒H. S. punctatus. (Balthasar, 1938). E. Dorsal view. F‒G. Lateral view from slightly different angles (red arrow indicates ventral surface). H. Ventral view. Observe the differences in the shape of the parameres, in particular how the apical hook is much more prominent in S. punctatus.

Fig. 25. Internal sac of Streblopus punctatus (Balthasar, 1938). A‒B. General aspect before dissection. C. Raspule. D. Apical and subapical sclerites. E. Superior right peripheral sclerite. Arrow indicates a triangular area of a more sclerotized tegument that is probably homologous to a similar triangular area on the SRP of S. opatroides van Lansberge, 1874. Abbreviations: A = axial sclerite; Rp = raspule; SA = subaxial sclerite; SRP = superior right peripheral sclerite; T = temones.

Fig. 26. Genital segment. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note that the medial sclerotized plate is larger and more sclerotized in S. punctatus than in S. opatroides; other differences refer to their lateral arches (indicated by blue arrows), which are concave in S. opatroides and more or less straight in S. punctatus, and the posterior arms of the MSP (indicated by red arrows), which are well developed and separated by a deep ʻU-shapeʼ in S. punctatus and only slightly indicated in S. opatroides. Abbreviations: LSP = lateral sclerotized plate; MSP = medial sclerotized plate.

Fig. 27. Female genitalia. A‒C. Spermatheca. A. Streblopus opatroides van Lansberge, 1874. B‒C. S. punctatus (Balthasar, 1938) (slightly different views). Observe that the spermatheca of the latter species is more slender and has a unique pair of apical hooks (red arrow). D. General view of the entire female genitalia of S. punctatus.

Fig. 14. Profemora. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Note the differences between the species and sexes in relation to the overall shape of the profemora and the presence of spurs on the anterior edge in males.

Fig. 15. Protibiae. A‒D. Dorsal view. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. E‒H. Ventral view. E‒F. S. opatroides. E. ♂. F. ♀. G‒H. S. punctatus. G. ♂. H. ♀. Notice the many differences between the species and sexes, in particular the presence of a protibial spur in males of S. punctatus (and its absence in S. opatroides) and the different shapes of the spurs and of the lateral teeth in females of the two species.

Fig. 16. Protibiae. A‒B. Lateral view. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note the how the protibia of S. opatroides is much more curved than that of S. punctatus. C. Frontal view of the protibia of S. punctatus. Note the presence of a protibial spur.