The genus Betiscoides Sjöstedt, 1924 was described by Sjöstedt (1924) and revised by Key (1937). Further descriptions of the morphological characters of Betiscoides with altered terminology followed (e.g. Dirsh 1965) and considering the new findings of this study, it is reasonable to provide a revised definition of the genus.

Type species: Betiscoides meridionalis Sjöstedt, 1924, type locality: South Africa. Body slender or thin, elongate, stick-like, smooth or hairy, with a prominent ventral line of hairs along the abdomen or at the end of tergits five to eight. Antennae thin or thick ensiform, segments strongly separated, finely and evenly rugose. Head in dorsal view conical. Frontal ridge compressed between antennae. Eye longitudinal or ovate and prominent. Pronotum cylindrical, with very weak or no median carina. Prosternal process trapezoid, T-shape, or lamelliformly compressed, laterally either flat or raised in the anterior or posterior part. Mesosternal interspace reduced, mesosternal lobes connected. Abdomen with fine longitudinal, dorsal median carina. Tympanum absent. Anterior and middle legs short; sometimes hairy, hind femur slender. External apical spine present. Tarsus shorter or half the length of the tibia. Arolium large, sometimes white. Male supra-anal plate triangular in dorsal view. Cerci very small, straight and conical. Subgenital plate elongate and conical. Female ovipositor valves slightly or strongly recurved at the tips; not toothed.

A phylogenetic reconstruction revealed that the sister taxon of the genus Betiscoides is Gymnidium Karsch, 1896 (Matenaar et al. 2016). The genus Betiscoides contains the following species: Betiscoides meridionalis Sjöstedt, 1924, B. parva Key, 1937, B. sjostedti Key, 1937, B. nova sp. nov. and B. muris sp. nov. While the description of B. meridionalis and B. sjostedti remains valid concerning the hairiness of the body, the body and especially the end of the abdomen of B. parva is fairly hairy (Fig. 6) and thus, its character description is hereby revised. In order to extend the definition for the three known nominal species considered by Key (1937), I provide the following additional morphological characters analyzed in the specimens collected at, or very close to, the type localities of the nominal species: ratio of the eye length to the length of the fastigium, degree of the male subgenital plate, volume of the body and, volume of the visible part of the eye. In addition, I provide measurements of the “female genital characters” for B. meridionalis.

Figure 6.

Overview of the nominal species of Betiscoides. A. B. sjostedti, HLMD-Cael-382, B21, DEL8; B. B. parva, HLMD-Cael-381, B103, DEL 9, and C. B. meridionalis HLMD-Cael-380, LB11, DEL 3. Specimens were collected at or close to the respective type localities. Scale bars: 1 mm.

Specimens of DEL 3 from Cape Point, Table Mountain NP, are hereby considered as Betiscoides meridionalis as they coincide with its overall morphological diagnostic characters. The large and slender body, the 24-jointed, ensiform, antennae, the very elongate head with the Fv nearly twice the length of the eye as well as the extremely elongate and acutely pointed subgenital plate, reason this allocation. The specimens occur in sympatry with the specimen from DEL 8.

Male. HLMD-Cael-380, LB11, Genbank Acc: KU206332, KU214602, 34°19'2.30"S, 18°25'12.66"E, 42 m above sea level, South Africa, Western Cape, Table Mountain National Park, Cape Point, Restio wetland, November 2013, L. Bröder leg.

Vol: 88.3974 mm³ or 8.83974e-08 m³; Vol. eye: 1002540337.66 µm³, ratio eye length/ fastigium length: 1.30, Subgenital plate degree: 31.6°

Female. HLMD-Cael-383, B24, Genbank Acc: MG243742.1, MG244116.1, MG243841.1, MG243940.1, MG244036.1, 33°57'32.18"S, 18°23'16.06"E, South Africa, Western Cape, Table Mountain National Park, Twelve Apostle, Restio wetland, March 2012, S. Wirtz leg.

Upper valve degree: 52°, upper valve distance = 182 µm, upper valve width = 201 µm, protrusion degree = 129°, Pr line 1 = 146 µm, Pr line 2 = 75 µm, Pr distance = 205 µm, Pr distance to tip of the dorsal valve = 280, 11 µm, Pr vertical distance = 78 µm, Pr straight-lined, apex acute.

Specimens of DEL 9 from the 12 Apostles, Table Mountain NP, are hereby considered as Betiscoides parva as they coincide with its overall morphological diagnostic characters. The small body size, the thick ensiform shape of the antennae and the large, prominent and ovate eyes, as well as the relatively large feet (half the length of the hind tibiae), reason this allocation. In order to assess the hairiness, the specimens of DEL 9 were compared with images of the holotype (male, NHMUK Cape Peninsular, South Africa, December 1930), both showed fairly hairy integument. In order to assess the female genital characters, I analyzed images of the allotype (female, NHMUK, Cape Peninsular, South Africa, December 1930).

Male. HLMD-Cael-381, B103, Genbank Acc: MG243794.1, MG243893.1, MG243991.1, MG244078.1, MG244168.1, 33°57'32.18"S, 18°23'16.06"E, South Africa, Western Cape, Table Mountain National Park, Twelve Apostle, Restio wetland, March 2012, S. Wirtz leg.

Vol: 42.080 mm³ or 4.20806e-08 m³; Vol. eye: 973202045.68 µm³, ratio eye length/ fastigium length: 0.54, Subgenital plate degree: 51° rounded.

The specimen of DEL 8 from Cape Point, Table Mountain NP, is hereby considered as Betiscoides sjostedti as it coincides with its overall morphological diagnostic characters. The intermediate body size, the almost equal length of the antennae and head, the fastigium about 2/3 the length of the eye, as well as the comparatively short subgenital plate, reason this allocation. In order to assess the hairiness, the specimen of DEL 8 was compared with images of the holotype (male, NHMUK, Table Mountain, South Africa, December 1929–1930), both showed fairly smooth integument. This specimen occurs in sympatry with specimens from DEL 3. In order to assess the female genital characters, I analyzed images of the allotype (female, NHMUK, Table Mountain, South Africa, December 1929–1930).

Male. HLMD-Cael-382, B21, Genbank Acc: MG243739.1, MG243838.1, MG243937.1, MG244033.1, MG244113.1, 34°19'2.30"S, 18°25'12.66"E, 42 m above sea level, South Africa, Western Cape, Table Mountain National Park, Cape Point, Restio wetland, March 2012, S. Wirtz leg.

Vol: 58.9269 mm³ or 5.89269e-08 m³; Vol. eye: 773244697.52 µm³, ratio eye length/ fastigium length: 0.78, Subgenital plate degree: 40.8°. The three nominal species are shown in Fig. 6.

The type material is deposited in the invertebrate collection of Hessisches Landesmuseum Darmstadt (HLMD). Holotype. Male, pinned. HLMD-Cael-4HT, B168, Genbank Acc. PP417675, South Africa, Western Cape province, Jonaskop, 33°58'9.75"S, 19°30'17.99"E, 1553 m above sea level, Plot 07, Restio wetland, 13 Dec 2013, leg. L. Bröder, E. Heym & D.Matenaar, Paratypes. Allotype. Female, pinned. HLMD-Cael-18AT, B166, paratypes HLMD-Cael-5-19PT, all same data as holotype. In total, five female and ten male paratypes are designated.

The specific epithet is a Latin adjective (feminine form) meaning ‘extraordinary’ or ‘new’. It refers to the species’ prominent habitus and to the fact that it is a new species characterized with the help of novel techniques.

The described species is assigned to the genus Betiscoides Sjöstedt, 1924 due to its phylogenetic position and overall morphological characteristics. The species is defined by the elongate, stick-like medium to large sized body, ensiform antennae, which are triangular in cross-section; its conical head, which is in profile almost equal in length and height, large eyes, the trapezoid shape of the prosternal process with two kinds of hairs and little variation in general. The end of the abdomen has only few and short hairs. The arolium is very large. The subgenital plate is elongate and conical; the ovipositor is short with robust, recurved valves.

This new species differs morphologically from the described ones mostly through its robustness and higher body volume and the almost identical HL and HH. The body is less elongate than Betiscoides meridionalis. Genital valves are less acute compared to Betiscoides meridionalis. Furthermore, the new species represents a divergent evolutionary lineage (DEL 19) as stated in Matenaar et al. (2018) and is genetically distinct representing the closest relative of B. parva (p-distance: ~ 0.067). As Key (1937) stated for B. parva and B. sjostedti, also this species is far less elongate than Betiscoides meridionalis, and B. parva, B. nova and B. sjostedti seem morphologically more related to another than any of these to B. meridionalis. In general, the new species seems to be morphologically intermediate between B. parva and B. sjostedti, whereby the robustness is more similar, to B. sjostedti but regarding the general coloration of the body, the shape of the Fr, the large eyes and the angularity of the end of the abdomen, there is a resemblance with B. parva, although B. nova is smooth, whereas B. parva is hairy. Furthermore, B. nova lacks the big feet of B. parva (Figs 7, 8).

Figure 7.

Lateral views of female allo- and paratypes of B. nova sp. nov. A. HLMD-Cael-8PT, B162; B. HLMD-Cael-16PT; C. HLMD-Cael-12PT; D. HLMD-Cael-14PT, B165; E. HLMD-Cael-18AT, B166. HLMD-Cael-18AT is the allotype.

Figure 8.

Lateral views of the paratypes of B. nova sp. nov. A. HLMD-Cael-7PT; B. HLMD-Cael-15PT, B167; C. HLMD-Cael-9PT; D. HLMD-Cael-6PT; E. HLMD-Cael-10PT, B163; F. HLMD-Cael-19PT; G. HLMD-Cael-11PT; H. HLMD-Cael-17PT; I. HLMD-Cael-5PT; J. HLMD-Cael-13PT.

Body robust, of medium size (Figs 9A, 10A). BL 25.81 mm, Integument finely rugose, shiny, smooth. Body of type specimen curved to the right in dorsal view, starting at the 10^th tergit. Antennae 22-jointed, ~ 4.5 mm, ensiform, slightly flattened above and slightly tapering, in length longer than the head, reaching the pronotum, evenly punctured, grooves darker than the rest, fine hairs, end of each segment lined in lighter ground color. HL 3.82 mm, HDL 6.13 mm, HH 3.77 mm, head from above elongate, conical, from above app. 1.5 as long as its width at the occiput. Fv 1.6 mm, 0.78 the length of an eye, angular, the margins well raised but obtuse-angled, margins of contrasting color, forming a thin, dark brown line, narrowing from the anterior margin of the eye forwards, protruding in front of the eyes culminating at the tip of the vertex as a protruding dot. Median carinula faintly discernible on the fastigium. Head in profile very slightly concave above, face slightly incurved, angle acute (app. 57.9°). Apex of fastigium in profile slightly raised; frons oblique. Fr between the antennae in profile somewhat projecting, thin, lamelliform, lateral carinae below the antennae very narrow, shallowly sulcate, in basal part obliterated. Fc straight, distinct throughout. Eyes ovate, about twice as long as their maximum width, their surface strongly convex, both margins somewhat curved, the upper more so than the lower, lower margin slightly raised, as of callus. Ratio of eye length and fastigium length 0.60. PL 3 mm. Pronotum cylindrical, with weak median and indistinct lateral carinae; no sulcus crossing dorsum. Hind margin of metazona very slightly concave, anterior margin straight; sides of pronotum with the lower margin almost straight, sloping down- and forwards in the anterior part, posterior margin practically straight, anterior lower angle slightly more than 90°, rounded; posterior lower angle 90°, rounded. Pp trapezoid, about twice as long as broad, its margins and angles rounded, the anterior end little broader than the posterior, finely haired and rugose (anterior width: 299.27 µm, posterior width: 263.23 µm, length ~ 492 µm), see Fig. 11A. Mesosternal interspace reduced/ closed with mesosternal lobes connected. Metasternal interspace like a pigs’ nose. Anterior and middle legs strongly shortened. Carinae of anterior and middle femora with fine and sparse hairs, anterior and middle tibiae and tarsi hairy, HFL 9.47 mm, hind legs reaching beyond the end of abdomen. Hind femora robust, about five times as long as their maximum width, upper carina of femur finely toothed as of small frequent tubercles, hind tibiae with 8 outer and 10 inner spines. External apical spine of hind tibia present. Hind tarsus shorter than half of the length of the tibia. Arolium very large and of white color. Supra-anal plate elongate, 1.33 mm, acutely angular. Cercus short, conical. Supra-anal plate with the basal part about one third the length of the apical part, the longitudinal depression distinct; the apical part of the plate shaped like an equilateral triangle with the base curved and the sides straight; apical angle widely rounded; no depression on the apical angle. Subgenital plate acute and elongate (angle app. 50°), but apex rounded, comparatively sharply pointed in dorsal view, lower margin convex, upper margin straight. Outer genitalia only sparsely haired with the subgenital plate featuring a fine line of hairs along the fold. Genital apparatus about the length of 2.5 tergits. SGP: 2.33 mm, three tergits: 3.35 mm ratio: 0.6955. The total body volume is 1.33029e-07 m³ (133.029 mm³). The volume of the visible part of the eye is 1160104343.13 µm³, the longitudinal height being 540.54 µm, and the length 2339.22 µm. The cross-sectional height is 657.29 µm and the width itself is 1372.3 µm. General coloration deep -brown. Antennae, fore and middle legs and knees are slightly paler than the brown primary color of the body. The upper part of the head, thorax and abdomen, within the callus line, is of green color. A longitudinal shallow stripe extends from the front of the head backwards along the central ridge of the body. The second and third tergit show brown markings between the callus lines; the posterior part of each tergit is of brown color, like a thin line, starting at the 5^th tergit. The lateral stripe backwards from the base of the eye is beige-white. It starts at the lower hind corner of the eye (as a somewhat raised callus ridge) and stops right behind the middle leg. Eyes brown, anterior femora reddish-brown, middle femora green, tibia and tarsi dark brown, hind femora of the same green as the dorsal part, knees and upper and lower external carina of the hind femur pale brown. Hind tibia brown and very dark in the lower side, spines and spurs black-tipped.

Figure 9.

Holotypes of Betiscoides nova sp. nov. and Betiscoides muris sp. nov. in lateral view. A. Shows B. nova, HLMD-Cael-4HT, B168; B. Shows B. muris, HLMD-Cael-364HT, B156.

Figure 10.

Holotypes of Betiscoides nova sp. nov. and Betiscoides muris sp. nov. in dorsal view. A. Shows B. nova, HLMD-Cael-4HT, B168; B. Shows B. muris, HLMD-Cael-364HT, B156.

Figure 11.

Prosternal proceses, ×200. A. Holotype of B. nova sp. nov., HLMD-Cael-4HT, B168; B. Shows the Pp of HLMD-Cael-10PT, B163 (B. nova) in ventral topview.

Larger than the male (Fig. 7E). BL 30 mm, HL 5 mm, HDL 7 mm, HH 3.79 mm, Head in profile straight, angle acute (app. 57.5°). Antennae 22-jointed, 5.5 mm, ensiform, finely punctured. Fr strongly protruding, lateral carinae of frontal ridge distinct beneath the sulcus, almost until clepeus. Fastigium 0.6 the length of an eye. PL 3.7 mm, sides of pronotum with lower margin straight; anterior margin sloping forward, only very slightly incurved, posterior margin slightly incurved, sloping forward in the upper part. Anterior lower angle more than 90°, posterior lower angle 90°, both rounded. Pp trapezoid, slightly resembling the number 8, rugose and finely haired with two different kinds of hair, margins rounded, the anterior part slightly broader than the posterior (anterior width: 297.32 µm posterior width: 212.37 µm, length: ~ 562 µm; measured in tilted angle (see Fig. 12). HFL 10.5 mm, hind tibiae with 10 outer and 12 inner spines, black tipped. Genital valves recurved, rounded in the apical part, hairs inside. Supra-anal plate 1.4 mm in total, subgenital plate acute. Genital valves spread. Substrate on the lower valves. Degree of protrusion acute (124°), Pr line 1 slightly longer (174 µm) than Pr line 2 (109 µm), Pr distance is 253,15 µm, Pr area is of 13755.38 µm² and protrusion vertical distance to tip is 180.92 µm. Protrusion to tip of valve distance is 311.02 µm. Upper valve degree acute of 163°, short upper valve distance: 239 µm, upper valve width 311 µm (see Fig. 13A). General coloration brown, in dorsal view head greenish brown, pronotum and abdomen colored in mixture of orange-brown, greenish brown, end of abdomen dark brown. Ventral ridge of the body finely but densely haired, head and hind femur of yellow-green color, knee and beginning of hind tibia pale brown, apically slightly darker, arolium large and white; black claws. The volume of the visible part of the eye is 1729519566.68 µm³. With the longitudinal height being 642.85 µm, and the length 2737.3 µm. The cross-sectional height is 646.17 µm and the width itself is 1412.78 µm.

Figure 12.

Image of the prosternal process of the allotype of Betiscoides nova sp. nov., HLMD-Cael-18AT, B166, ×200, captured in ventral view, with a tilt angle of 60°.

Figure 13.

Ovipositor valves, lateral view. A. Allotype of B. nova sp. nov. HLMD-Cael-18AT, B166; B. Allotype of B. muris sp. nov. HLMD-Cael-368AT, B155.

Specimens show little differences in the shape of the head in profile, varying from described morphology of the holotype to a slightly more convex form. By contrast, the face differs concerning the distinction of the lateral carinae below the sulcus and the shape of the sulcus itself. In some specimens, the lateral carinae are distinct throughout, also beneath the sulcus. The sulcus is shaped as a mustache sometimes. Median Carina/ Central ridge of the body: Whereas the median carina is present in all specimens, the central ridge of the body is clearly distinct in males on the abdomen, starting on the third segment. This pattern is truly sex specific. Females show a distinct median carina throughout, but in three females (HLMD Cael-18AT, B166; HLMD-Cael-14PT, B165; and HLMD-Cael-16PT) the median carina culminates in a black spot at the verge of the fastigium of vertex. The prosternal process was described as laminate or lamelliform compressed by Key (1937) and Dirsh (1965). In contrast to the other known Betiscoides species, the Pp of B. nova can be described as trapezoid with vague variability towards a square shape. In 2D some specimens seem to show a proximal concave Pp (see Fig. 11B). Detailed 3D image analysis revealed a straight but rugose progression of the surface of the Pp (see Fig. 14). The curve progression showed a gradient of ~ 54 µm (HLMD-Cael-10PT, B163, see Fig. 14). Supra-anal plate and subgenital plate more elongate than in the holotype. Variation in spines on the hind tibia: In male, the hind tibiae had 8–9 outer and 10–11 inner spines. In females the hind tibia showed 9–10 outer spines and 11–12 inner spines. General coloration in males dark-brown. Antennae, fore and middle legs are of the same brown primary color as the body. The upper part of the head, thorax and abdomen, within the callus line, is of light brown or green color. When of green color, sometimes the hind margin of each segment of the abdomen is of light brown or orange color. Eyes brown. Hind femora pale, especially when light brown within the callus line; or green, both inside and outside, except the carina, which is sometime black-tipped, herring bone pattern distinct. Knees light brown. Hind tibiae brown and very dark in the lower side, spines and spurs black-tipped. Female: General primary coloration similar to, but slightly less dark than, that of the male. No variation toward green or light brown color within the callus line. Hind femora, both outside and inside, paler than the rest of the body, varying in one female towards yellow-green, herring bone pattern distinct, hind tibiae darker apically than basally. Detailed measurements are provided in Table 1.

Figure 14.

3D image profile analysis of the prosternal process of specimen HLMD-Cael-10PT, B163 (B. nova sp. nov.). A. Shows the 3D image turned to profile view and the two points set to measure the distance and height progression curve of the selected part; B. Shows the depth image composition of the 3D image and the two points set to measure the distance; C. Shows the profile analysis results, including the length and height and the curve progression.

The longitudinal height cannot be safely derived from the measurements since most of specimens eyes have been subject to certain degree of decay, which caused the highest part of the eyes to collapse at the highest point. Specimens HLMD-Cael- 6, 8, 9, 13, and 15PT were excluded from the cross sectional height analysis since the eyes had collapsed.

Described from 11 males and five females from Jonaskop, Western Cape, South Africa, 2013. The species occur on top of Jonaskop in restio-dominated wetland. Abundant in patches with water covering the bare ground between the single restio plants. As the three formally recognized species of Betiscoides are considered endangered (Hochkirch 2012a, 2012b, 2012c), I propose this status as well for B. nova because itis only known from a single locality and its habitat is in decline.

The type material is deposited in the invertebrate collection of Hessisches Landesmuseum Darmstadt (HLMD). Holotype. Male, pinned, HLMD-Cael-364HT, B156, (Genbank Acc. PP411598, PP417668), South Africa, Western Cape province, Groot Winterhoek Wilderness Area, 32°59'21.52"S, 19°3'26.84"E, 991 m above sea level, Plot 134, Restio wetland, 18 April 2016, leg. D. Matenaar.

Paratypes. Allotype. Female, pinned. HLMD-Cael-368AT, B155, South Africa, Western Cape, Groot Winterhoek Wilderness Area, 32°59'53.06"S, 19°4'12.40"E, 929 m above sea level, Plot 133, Restio wetland, 18 April 2016, D. Matenaar. Paratypes. HLMD-Cael-362PT, B153; HLMD-Cael-366PT, South Africa, Western Cape, Groot Winterhoek Wilderness Area 32°59'35.34"S, 19°3'32.90"E, 969 m above sea level, Plot 65, Restio wetland, 18 April 2016, D. Matenaar. HLMD-Cael-363PT; HLMD-Cael-367PT, B154, South Africa, Western Cape, Groot Winterhoek Wilderness Area, 32°58'54.76"S, 19°3'18.37"E, 1001 m above sea level Plot: 66, Restio wetland, 18 April 2016, D. Matenaar. HLMD-Cael-365PT South Africa, Western Cape, Groot Winterhoek Wilderness Area, 32°59'53.06"S, 19°4'12.40"E, 929 m above sea level, Plot 133, Restio wetland, 18 April 2016, D. Matenaar. HLMD-Cael-360PT, HLMD-Cael-361PT; HLMD-Cael-369PT, South Africa, Western Cape, Groot Winterhoek Wilderness Area, 134: 32°59'21.52"S, 19°3'26.84"E, 991 m above sea level, Plot 134, Restio wetland, 18 April 2016, D. Matenaar. HLMD-Cael-379PT, B169, South Africa, Western Cape, Groot Winterhoek Wilderness Area, Restio wetland 33°0'11.81"S, 19°4'20.91"E, 908 m above sea level, Plot 95, Restio wetland, 06 March 2020, D. Matenaar. In total, five female and five male paratypes are designated.

The specific epithet is a Latin noun, meaning mouse. It refers to the species’ very slender and delicate habitus.

The described species is assigned to the genus Betiscoides Sjöstedt, 1924 due to phylogenetic and overall morphological characteristics. The very slender, elongate, stick-like and medium sized body defines the species. The antennae are ensiform and triangular in cross-section. The head is acutely conical elongated and the end of the abdomen shows dense and long hairs. The arolium is large. The subgenital plate is elongate and acutely conical; the ovipositor is short with well-curved valves and sharp tips.

This new species differs morphologically from the described ones mostly by its overall delicate habitus, the comparatively densly and longhaired end of the abdomen of the males. The body is smaller and more delicate than Betiscoides meridionalis. The head is more elongate than in B. nova and B. parva. The genital valves are strongly acute, sharply pointed, upper valve width narrower than in Betiscoides meridionalis. Just as B. nova, the new species represents a divergent evolutionary lineage as stated in Matenaar et al. (2018) being most closely related to B. parva (p-distance: 0.0734). As Key (1937) stated for B. parva and B. sjostedti, also this species is far less elongate than Betiscoides meridionalis, and B. parva, B. nova and B. sjostedti as well as B. muris are morphologically more similar to each other than any of these to B. meridionalis. Morphologically B. muris relates the most to B. sjostedti although B. sjostedti lacks the hairy end of the abdomen, which instead can be found even stronger in B. parva (Fig. 15). The genetic distance of B. muris to B. meridionalis is closer than to B. nova or B. sjostedti. The frontal ridge of the new species is more prominent and protruding than in B. nova; Fig. 16.

Figure 15.

Images of the end of the abdomen of specimens A. HLMD-Cael-364HT, B156 (Betiscoides. muris sp. nov.); B. HLMD-Cael-4HT, B168 (B. nova sp. nov.); C. B. parva (HLMD-Cael-381, B103) and D. B. sjostedti (HLMD-Cael-382, B21).

Figure 16.

Lateral views of female and male type specimens B. muris sp. nov. A–D. HLMD-Cael-360-363PT; E. HLMD-Cael-364HT, B156; F–H. HLMD-Cael-365-367PT; I. HLMD-Cael-368AT, B155; J. HLMD-Cael-369PT.

Body of medium length, slender, delicate (Figs 9B, 10B). BL 22.5 mm. Integument finely rugose, shiny golden. Antennae 22-jointed, ~ 6 mm, finely ensiform, slightly flattened above and slightly tapering, finely pointed and shiny, evenly punctured, in length longer than the head, reaching the pronotum. Left antenna between the 12^th and 13^th and on the 15^th segment marked with shiny orange fluid-like spot. Head elongate, conical, from above almost 4 times as long as its width at the occiput and twice the diagonal length than the height, HL 3.8 mm, HDL 5.52 mm, HH 2.59 mm. Fv 1.4 mm, 0.7 the length of an eye, angular, the margins well raised but obtuse-angled, narrowing from the anterior margin of the eye forwards; callus line above the eyes culminating into the margins of Fv, (straight), strongly protruding in front of the eyes; median carinula faintly discernible on the fastigium. Head in profile convex above, face slightly incurved; angle acute (app. 49.4°). Apex of fastigium in profile slightly raised culminating dot-like at the very apex; frons oblique. Fr between the antennae in profile somewhat projecting, thin, lamelliform, lateral carinae distinct, below the antennae very narrow, shallowly sulcate, crossing the sulcus and only close to the clepeus at the basal part obliterated. Fc straight, distinct throughout. Eyes oval, 1 ¾ as long as their maximum width, their surface strongly convex, both margins somewhat curved, the upper more so than the lower, upper and lower margin slightly raised, as of callus. Ratio of eye length to fastigium length 0.61. PL 2.6 mm, lateral ventral length 1.78 mm, pronotum cylindrical, with weak median and indistinct lateral carinae; no sulcus crossing dorsum. Hind margin of metazona very slightly concave, anterior margin straight; sides of pronotum with the lower margin straight, anterior lower angle slightly more than 90°, rounded; posterior lower angle 90°, rounded. Pp destroyed through pinning. Mesosternal interspace reduced with mesosternal lobes connected. Anterior and middle legs strongly shortened. Tibia and tarsi haired, tarsi with black claws. Hind legs reaching beyond the end of abdomen. HFL 8.1 mm, hind femora very slender, about 5.5 times as long as their maximum width; outer apex of knees acute; hind tibiae with 10 outer and 13 inner spines. External apical spine of hind tibia present. Hind tarsus shorter than half of the length of the tibia. Arolium extremely large, its margin of beige color. Male supra-anal plate elongate, acutely angular. Cerci short, conical. Subgenital plate strongly elongate, 2.85 mm, acutely conical, 35.8°. End of abdomen with dense but fine, comparatively long hairs, see Fig. 15A. Supra-anal plate with the basal part about 0.65 the length of the apical part, the longitudinal depression distinct; the apical part of the plate shaped like an equilateral triangle with the base curved and the sides straight; apical angle widely rounded; no depression on the apical angle. Subgenital plate about twice the length of the supra-anal plate; comparatively sharply pointed; lower margin convex, upper margin straight; apex acute. Genital apparatus about 3.5 the length of one tergit. SGP: 2.85 mm, last three tergits: 2.32 mm ratio: 1.22. The total body volume is 5.38305e-08 m³. The volume of the visible part of the eye is 587729028.76 µm³. The longitudinal height being 493.28 µm, and the length 2137.79 µm. The cross-sectional height is 483.82 µm and the width itself is 1213.70 µm. General coloration dark-brown. Antennae, fore and middle legs and knees are slightly paler than the brown primary color of the body. The upper part of the head, thorax and abdomen, within the callus line, is beige. The Fv is dark brown almost black. A longitudinal stripe extends from the front of the head backwards along the central ridge of the body. The lateral stripe backwards from the base of the eye is beige-pink. It starts at the lower hind corner of the eye (as a somewhat raised callus ridge) and reaches the end of the middle leg. Eyes brown, hind femora of brown color processing into yellow beneath, knees, upper and lower external carina of the hind femur yellow. Hind tibia orange and brown in the lower side, spines and spurs black-tipped.

Larger than the male (Fig. 16I). BL 26.5 mm, antennae 22-jointed, ~ 5 mm. fastigium about as long as an eye. The anterior and posterior margin of the eye are somewhat raised, as of torus. HL 4.3 mm, HDL 5.89 mm, HH 2.92, head in profile acute-angled (50.9°). PL 3 mm, sides of pronotum with lower margin straight; anterior margin sloping forward, only very slightly incurved, posterior margin very little incurved, sloping forward in the upper part. Anterior lower angle more than 90°, posterior lower angle 90°, both rounded. Pp T-shape, proximal concave, 673.43 µm in length, anterior part 233.54 µm width and 52.87 µm higher than the posterior part (of 122.29 µm width, see Figs 17, 18), fine and thin hairs on the broader anterior part and at the sides only. HFL 9 mm, hind tibiae with 10 outer spines, 11 inner spines. Protrusion of lower genital valves crossed by two diagonal sulci. Pr lines 1 and 2 slightly curved, bending inwards, apex of Pr thereby rounded although degree of protrusion strongly acute (79°), Pr line 1 slightly longer (155 µm) than Pr line 2 (91 µm), Pr distance is 199 µm, Pr area is of 12455.69 µm² and protrusion vertical distance is 129.66 µm, protrusion to tip of valve distance (horizontal distance) is 330.13 µm (see Fig. 13B). Tips of the valves black and strongly acute. End of upper valve slender and acute pointed, of 134°, upper valve distance 173 µm, upper valve width is 173 µm. General coloration similar to, but slightly less dark, than that of the male. Eyes and hind femora, both outside and inside, paler than the rest of the body; outer side of femur with black stripe and fishbone pattern; hind tibiae darker than the femur but without gradient in color. The volume of the visible part of the eye is 1356324476.51 µm³. With longitudinal height being 760.13 µm, and the length 2131.68 µm. The cross-sectional height is 696.96 µm and the width itself is 1175.43 µm.

Figure 17.

High resolution DOF images of the prosternal process of Betiscoides muris sp. nov. clustering in DEL 22, Groot Winterhoek, revealing four different shapes of Pp. A. Image of HLMD-Cael-379PT, B169; B. Image of HLMD-Cael-366PT; C. Image of allotpye HLMD-Cael-368AT, B155; D. Image of HLMD-Cael-367PT, B154.

Figure 18.

3D-image of the prosternal process of the allotype of Betiscoides muris sp. nov. HLMD-Cael-368AT, B155. The output of the 3D profile image analysis is shown, including the curve progression from the anterior to the posterior part in profile, and total length and height.

The antennae are 22–23-jointed and the segments are covered with fine, black dots and shiny appearance in all specimens. Posterior margins of the antennae segments of one male specimen are matt beige, while the rest is of the usual common shiny brown color. Specimens show little differences in the shape of the head in profile, varying from described morphology of the holotype to a slightly more concave form. Similar to B. nova, the face varies concerning the degree of distinction of the lateral carinae below the sulcus and the shape of the sulcus itself. In some specimens, the lateral carinae are distinct throughout, also beneath the sulcus. The sulcus is shaped as a mustache sometimes. This intraspecific variation is not sex-specific. Variation in spines on the hind tibia: In male, the hind tibiae had 9–10 outer and 10–14 (!) inner spines. In females, the hind tibia showed 9–10 outer spines and 11–13 inner spines. In contrast to the other known species of Betiscoides, there seems to be a intraspecific variability in the shape of the prosternal process, in general the Pp is about twice as long as broad, its margins and angles rounded; the anterior end little broader than the posterior. However, four different shapes can be distinguished within the B. muris species, making the Pp itself a variable character within this species (see Fig. 17): a) thin lamelliform (one thin line from above), b) trapezoid, c) T-Shape, d) thick lamelliform. Furthermore, the trapezoid and thick lamelliform Pps are often tuberculate and hairy, whereas T-shape and thin lamelliform ones are rather smooth and not very rugose. The 3D image analysis revealed variations in the elevation of the prosternal process. In some specimens, the anterior part is higher than the posterior and vice versa (see Figs 18, 19). The Pp of specimen HLMD-Cael-379PT, B169 is 541 µm long and the posterior part is 167.91 µm higher than the anterior; the Pp of specimen HLMD-Cael-367PT, B154 is 614.53 µm long and the posterior part is 148.39 µm higher than the anterior. General coloration in males dark-brown. Antennae, fore and middle legs are of the same brown primary color as the body. The upper part of the head, thorax and abdomen, within the callus line, is of light brown, orange or yellow color. The lateral stripe backwards from the base of the eye is beige. It starts at the lower hind corner of the eye (as a somewhat raised callus ridge) and stops right behind the middle leg. In one male, the stripe seems to extend to the femur as a yellow line. Eyes brown, hind femora pale, especially when light brown within the callus line, or with blueish line. In life, sometimes of green color (Fig. 20). Knees light brown. Hind tibiae brown, spines and spurs black-tipped. Female: General primary coloration similar to that of the male. No variation toward green or light brown color within the callus line. Hind femora, both outside and inside, paler than the rest of the body, varying in one female towards blue, femoral stripe often present, herring bone pattern distinct hind tibiae brown. Detailed measurements are provided in Table 1.

Figure 19.

3D image of the prosternal process of B. muris sp. nov. specimen HLMD-Cael-379 PT, B169. The output of the 3D profile image analysis is shown, including the curve progression from the anterior to the posterior part in profile, and total length and height.

Figure 20.

Male individual of Betiscoides muris sp. nov. from Groot Winterhoek Wilderness Area, South Africa in life; image taken on 06 March 2020, 32°58'54.76"S, 19°3'18.37"E.

Specimen HLMD-Cael-379PT, B169 is not included in the subgenital plate degree measurement due to damage.

Described from six males and five females from Groot Winterhoek, Western Cape, South Africa, collected in 2016 and 2020. The species occurs on the plateau of the Groot Winterhoek Nature reserve in restio-dominated wetland. It is abundant in patches with water covering the bare ground between the single restio plants. As the three formally recognized species of Betiscoides are considered “endangered” (Hochkirch 2012a, 2012b, 2012c), I propose this status as well for B. muris as this species is only known to occur at a single locality and its habitat is in decline.