Research Article |
Corresponding author: Renzo Perissinotto ( renzo.perissinotto@mandela.ac.za ) Academic editor: Andrey Frolov
© 2019 Renzo Perissinotto, Petr Šípek.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Perissinotto R, Šípek P (2019) New species of Xiphoscelis Burmeister, 1842 (Coleoptera, Scarabaeidae, Cetoniinae) from arid regions of South Africa and Namibia. ZooKeys 879: 57-89. https://doi.org/10.3897/zookeys.879.37721
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Two new species of the southern African genus Xiphoscelis Burmeister, 1842 are recognised and described, X. braunsi sp. nov. from the Eastern and Western Cape Karoo (South Africa) and X. namibica sp. nov. from the Huns Mountains of southern Namibia and adjacent ranges in South Africa. These were previously overlooked and grouped together with X. schuckardi Burmeister, 1842, but further material and more in-depth analyses have now revealed their clear separation on the basis of key diagnostic features, including clypeal structure, metatibial spur development and aedeagal shape. The densely and coarsely costate elytral structure and the black to brown colour of these species are symplesiomorphies shared with a number of the most primitive genera among the African Cetoniinae. However, these characters also reflect the convergent adaptation to hot and arid conditions they share with several other species occurring in this region. Phylogenetic relationships of the genus with other Cetoniinae are explored using the larval characters highlighted in the description of the 3rd instar larva of X. braunsi sp. nov. The extraordinary hypertrophy observed in the male metatibial spur of species in this genus, and particularly in X. schuckardi, appears to represent a defence mechanism against potential predators on the ground, apart from playing a role during mating.
Afrotropical region, fruit chafers, identification key, immature stages, life cycle, Succulent Karoo, taxonomy, Xiphoscelidini
The genus Xiphoscelis has been recognised since its first description as characterised by unique features, such as a narrow mesometasternal process, enlarged metafemur, low subhumeral elytral arch, round pronotum and regularly costate elytra. To accommodate these, apparently plesiomorphic features, a supra-generic grouping was proposed with the aim of clustering together a variety of genera sharing the key characters.
However, later
As already pointed out by
To complicate matters further, recent molecular analyses indicate that the phylogeny of Cetoniinae in general needs a substantial revision, as large incongruences with the traditional concepts are emerging (
There are currently three species described within the genus Xiphoscelis, two of which (X. lenxuba Perissinotto, Villet & Stobbia, 2003 and X. sneeubergensis Perissinotto, Villet & Stobbia, 2003) were only recently separated from the type and only species previously recognised, X. schuckardi (
Holotype specimens of both Xiphoscelis schuckardi Burmeister, 1842 (♂, 17 mm total length, “Pr. b. sp. Sch.”) and Xiphoscelis gariepena Schaum, 1849 (nec Gory & Percheron) (♀, 15.8 mm total length, “Afr. Austr”) were studied in detail through high-resolution photographs kindly provided by Karla Schneider (
Other specimens were obtained through field collections during the period 1995–2018 (R Perissinotto & L Clennell legit), or from museum and private collections (as per list provided below). Fresh specimens were either caught in flight using standard nets after rainfall events, excavated from underground or obtained after rearing third instar larvae collected in the wild under laboratory controlled-conditions. In the laboratory, larvae were kept in plastic containers of 1–5 L capacity, containing the natural soil and detrital material found in situ. Water was sprayed at the soil surface at regular intervals of about 1–2 weeks until pupation.
Data on distribution, period of adult activity and other biological information for all the species of the genus Xiphoscelis were also obtained from
As in previous works, the description of adult morphological characters follows the terminology of
The identity of the larvae was confirmed by both rearing specimens to adulthood and by molecular match (COI) with adult specimens. Specimens for DNA extraction were stored in 96% ethanol immediately after capture. Genomic DNA from thoracic leg muscle tissue (both larvae and adults) was extracted non-destructively using a Qiagen Blood and Tissue Kit, following standard protocols. Voucher specimens are deposited at the NMCR. Partial sequences of the mitochondrial protein coding gene cytochrome oxidase subunit 1 (Cox1) were used in the study. For a detailed description of the laboratory protocol see
Larval material was examined with an Olympus SZ9 and a Nikon SMZ 745 stereomicroscope, under which measurements were taken with an ocular grid. Habitus photographs were taken using a Canon EOS 70D camera fitted with Canon EF–S 60 mm f/2.8 Macro USM lens or Canon MP–E 65 mm f/2.8 1–5× macro lens. Microscopic slides were photographed using a Canon EOS 70D camera mounted on an Olympus SZ9 stereomicroscope. Partially-focused images of specimens were combined using Zerene Stacker (Zerene Systems LLC, Richland, USA). Structures examined using scanning electron microscopy (JEOL, Model 6380, Tokyo) were cleaned in 10% lactic acid for 24 h and submerged into a Sonorex ultrasonic bath (Bandelin electronics, Berlin) for 30 s, dried in a heating chamber, or using critical point drying, and mounted on aluminium plates. All pictures were digitally enhanced using Adobe Photoshop CC.
The terminology for larval description follows
To compare the observed larval morphological characters of X. braunsi sp. nov. with those of other cetoniines from various clades, a basic phylogenetic analysis based on a matrix of 77 larval morphological characters of 13 taxa was performed, as modified from
Specimen repositories are abbreviated as follows:
BMPC Jonathan Ball and Andre Marais Private Collection, Cape Town, South Africa
NMCR Národní Museum, Prague. Czech Republic
SRPC Sébastien Rojkoff Private Collection, Sourcieux les Mines, France
TGPC Thierry Garnier Private Collection, Montpellier, France
UKCR Univerzita Karlova, Katedra Zoologie, Prague, Czech Republic
This species differs from X. schuckardi by its matte, black to brown dorsal colouration (black and shiny in X. schuckardi) and the elytral costae which are weakly elevated and poorly visible, rather than prominent as in X. schuckardi (Figs
Finally, the parameres of the two species are also different at the level of the inner apical end of the dorsal lobes, which is finely pointed in X. braunsi, but rather blunt in X. schuckardi (Figs
(Figs
Body.
Black to dark brown, completely matte except for small worn ridges on elytral umbones (Fig.
Head. Black to dark brown, with round sculpture on frons, becoming irregularly shaped on vertex; ultrafine rugosity across entire surface; long, erect black setae restricted to eye canthus and antennal pedicel; clypeus markedly bilobate and deeply concave, with anterior margin sharply elevated and lateral margins perfectly rounded both posteriorly and anteriorly; antennal clubs and flagellum black to dark brown, of normal cetoniine length; pedicel dark brown, becoming lighter towards base.
Pronotum.
Dark brown and matte, becoming blackish at margins; smoothly rounded at all margins except antero-lateral, which exhibit sharp angles leading to medio-apical, weakly elevated transversal protuberance; posterior margin forming straight line in front of scutellum; small, scattered round to horse-shoe punctures throughout surface, becoming denser towards lateral and anterior margins; thick, black, medium to long setae regularly distributed across entire surface and emerging at centre of punctures, becoming denser and longer at lateral margins (Fig.
Scutellum.
Black on sides to dark brown on disc; isoscelic triangular with sharply pointed apex and deep but narrow lateral grooves; with regularly spaced horse-shoe punctures restricted to basal and baso-lateral margins; dense line of short, light setae along basal margin, just below posterior pronotal margin (Fig.
Elytron.
Black to dark brown, not covering entire abdomen and leaving external projection of sternites and pygidium partly exposed; subhumeral arch very low and postero-apical declivity extremely steep and abrupt; with costae 1–4 poorly elevated in basal two-thirds, becoming virtually flat in apical third; 5th costa and umbones raised, with latter showing shiny area on top; rest of surface matte and regularly sculptured with geminate striae along intercostal spaces 1–4, becoming round to horse-shoe beyond 5th costa; with sparse short, but thick and erect black setae across whole surface, except umbones; apex without spinal projection and weakly rounded (Fig.
Pygidium.
Black at base gradually becoming dark brown towards apex; remarkably narrow and triangular in shape, with basal and baso-lateral margins sharply upturned; with dense, rugose sculpture on basal third, becoming scattered and round towards apex; with moderate central bulge and shallow, symmetric baso-lateral depressions; without pubescence on general surface but with lining of black, long setae along entire apical margin (Fig.
Legs.
Tarsal segments consistently black and elongate, but tibiae reddish-brown with black tips and of normal cetoniine thickness and length; protibia tridentate, with proximal tooth drastically reduced; mesotibia bearing mid outer spine with four thick setae on its surface, two apical spines and two spurs of small to medium size; metatibia with extreme hypertrophic inner spine and spurs, inner spine 2–3 times as thick and 1.5 times as long as spurs; (Fig.
Ventral surface. Shiny, reddish-brown to black; with thin and long dark setae on prosternum, coxae and all femoral margins, becoming shorter and more sparse on other surfaces, particularly abdominal sternites; all setae emerging at centre of small and round sculptures; mesometasternal process extremely small, not protruding forward or upwards and partly covered by coxal bases, reddish-brown to black, with scattered round punctures and thin setae on surface; abdominal sternites initially flat, but forming concavity at middle particularly in area of sternites 5–7.
Aedeagus.
Parameres with dorsal lobes tapering gradually and smoothly towards apex, forming short spinal apical inner end, covering completely ventral lobes in dorsal view (Fig.
Derivatio nominis. This species is dedicated to the memory of Hans Brauns (1857–1929), renowned physician and entomologist, who during the early 20th century lived and collected extensively in the Willowmore District of the Eastern Cape Province. Most of the early specimens of the new species described here formed part of his collection.
The only reliable and consistent external feature of sexual dimorphism in this species lies in the development of the metatibial internal apical spine and spurs, which are far less hypertrophic in the female, compared to the male. Also, like in most cetoniines, the protibiae and protarsi are appreciably shorter in the female than in the male, and the abdominal sternites of the male are usually concave in the central area while those of the female tend to be flat or slightly convex.
This species appears to be restricted to the Eastern and Western Cape provinces of South Africa. Apart from the long series collected at Willowmore by Brauns in the early part of the 20th century, specimens have recently been found in mountainous areas of the western part of the Eastern Cape and in the interior regions of the Western Cape, at altitudes > 500 m but not exceeding 1000 m asl. Thus, the species appears to follow the geographic range of the Cape Fold Belt, where it inhabits the lower slopes of its mountain ranges (Fig.
Both literature and specimen data records report regular occurrences of association of this species with the southern harvester termite, Microhodothermes viator (e.g.,
While the ventral habitus of this species is remarkably stable in colour, being predominantly reddish-brown, the dorsal surface ranges across two extreme varieties, one completely black (Fig.
Holotype (♂): South Africa, EC, Fullerton, 5 Jan 2017, R. Perissinotto & L. Clennell (
Xiphoscelis namibica can best be separated from both X. schuckardi and X. braunsi by the characteristics of its parameres, as it is the only species among the three to exhibit an apical protuberance on the inner margin of each dorsal lobe (Fig.
(Figs
Body.
Completely black and matte, except for small worn ridges on elytral umbones (Fig.
Head. Entirely black, with coarsely round sculpture throughout surface; ultrafine rugosity across entire surface; long, erect black setae on eye canthus and antennal pedicel; clypeus weakly bilobate but deeply concave, with both anterior and lateral margins equally elevated, lateral margins smoothly rounded all around; antennal clubs, flagellum and pedicel black and of normal cetoniine length; pedicel becoming lighter and brown towards base.
Pronotum.
Black and matte; regularly round in shape, except at antero-lateral margins, where sharp angles lead to medio-apical, moderately elevated transversal protuberance; posterior margin forming perfectly straight line in front of scutellum; round punctures regularly spaced across surface, but becoming more scattered on disc and denser at margins and on lateral declivities; thick, black, setae of medium length visible only on lateral margins (Fig.
Scutellum.
Completely black; isoscelic triangular with weakly rounded apex and lateral grooves absent on basal third but well developed along other two-thirds towards apex; with dense but irregularly shaped punctures on basal and baso-lateral margins, but absent on central part of disc and on apical third; few, thick and black erect setae scattered across basal third of surface (Fig.
Elytron.
Completely black and matte, narrower than abdomen leaving apical projection of sternites and pygidium partly exposed; subhumeral arch very low and postero-apical declivity extremely steep and abrupt; all costae subequally and weakly elevated, with 5th and 6th costae and umbones raised; surface densely sculptured with geminate striae or round to irregular punctures along intercostal spaces, becoming very sparse and occasional on surface of costae; sparse short, but thick and erect black setae across whole surface, except umbones; apex with short but distinct spinal projection (Fig.
Pygidium.
Completely black, narrow and broadly triangular in shape, with basal and lateral margins sharply upturned; with uniformly sparse horse-shoe sculpture across surface; convex with small baso-lateral depressions; without pubescence on general surface but with lining of black, long setae along lateral and apical margins (Fig.
Legs.
All legs black in dorsal view, with tarsal segments elongate, but tibiae of normal cetoniine thickness and length; protibia tridentate, with proximal tooth reduced and other two teeth severely worn; mesotibia exhibiting mid outer spine, two apical spines and two spurs of small to medium size; metatibia with slightly hypertrophic inner spine and spurs, inner spine much thicker but shorter than spurs (Fig.
Ventral surface. Shiny and black, with reddish-brown areas restricted to part of coxae and basal portion of femora; with thin and long dark setae on prosternum, coxae and all femoral margins, becoming shorter and more sparse on other surfaces, particularly abdominal sternites; all setae emerging at centre of small and round sculptures; mesometasternal process extremely small, not protruding forward or upwards and partly covered by coxal bases, black and with scattered round punctures and thin setae on surface; abdominal sternites slightly convex, becoming flat at middle particularly in area of sternites 5–7.
Aedeagus.
Parameres with dorsal lobes tapering abruptly towards apex, forming steeply elevated apical spine at inner end, covering completely ventral lobes in dorsal view (Fig.
Derivatio nominis. With the exception of one female from O’Kiep (South Africa), so far, all the specimens known for this species and representing the type series originate from the same locality, in south-western Namibia. Hence the obvious geographic link to its name.
Unlike with all the other species of the genus, it is virtually impossible to separate the two sexes of X. namibica on the basis of external morphology alone. This is because the metatibial internal apical spine and spurs of its male (Fig.
So far, the few specimens known for this species have been collected mostly in south-western Namibia, near the town of Rosh Pinah, in the Namuskluft area at about 1200 m asl (Fig.
The holotype and paratype series collected by Holm & Gebhardt in southern Namibia were all retrieved dead from middens of the southern harvester termite Microhodothermes viator. Given the very limited number of observations available for this species, it is not possible to establish whether or not this is a case of obligatory association, or again a rather opportunistic one.
Holotype (♂): Namibia, Namuskluft, 1200 m, 27°45'S, 16°53'E, 2–6 Apr 2002, in Microhodothermes viator middens, E. Holm & H. Gebhardt (
1 | Dorsal habitus completely black and shiny; elytral costae markedly elevated and visible; general dorsal sculpture dense and deep; body length 14–22 mm, larger than all other species; distribution: west coast lowlands of Western and Northern Cape |
X. shuckardi Burmeister, 1842 (Fig. |
– | Dorsal habitus black or brown and matte or velutinous; elytral costae weakly elevated and dorsal sculpture scattered and shallow; body length 11–16 mm | 2 |
2 | Dorsal habitus exhibiting cretaceous markings | 3 |
– | Body without cretaceous markings | 4 |
3 | Cretaceous ornamentation extensive on elytral surface, pronotal margins and protruding areas of abdominal sternites; body covered in long, scattered black setae; distribution: high mountains of Eastern Cape Karoo, above Great Escarpment |
X. sneeubergensis Perissinotto, Villet & Stobbia, 2003 (Fig. |
– | Cretaceous markings moderately developed on pronotal margins and occasionally present also on elytra, but very faintly; body velutinous and covered in dense, medium to long golden-brown or orange setae; distribution: eastern areas of Eastern Cape Karoo |
X. lenxuba Perissinotto, Villet & Stobbia, 2003 (Fig. |
4 | Dorsal habitus completely black and matte; metatibial internal apical spine as long as spurs in both sexes; anterior clypeal margin weakly sinuate to straight; distribution: south-western Namibia and adjacent areas of Northern Cape |
X. namibica Perissinotto, sp. nov. (Fig. |
– | Dorsal habitus black or brown to reddish-brown; metatibial spurs and particularly inner apical spine hypertrophic in male; anterior clypeal margin strongly sinuate; distribution: Cape Fold Belt of the Western and Eastern Cape provinces |
X. braunsi Perissinotto & Šípek, sp. nov. (Figs |
Figs
Differential diagnosis. The larvae of Xiphoscelis braunsi sp. nov. are characterised by the following characters: long and prolific chaetotaxy on cranium; frons with 2–3 posterior frontal setae, anterior and lateral frontal setae long; epipharynx with sensorial cone (left nesium) desclerotised, low, obtuse, reduced and plate-shaped; mandibles with an external tooth on lateral margin; lacinial unci unequally fused at their base, smaller uncus about three times shorter than the longer one; hypopharyngeal scleroma without truncate process; pretarsi cylindrical with 11–12 setae, raster composed of 2 slightly subparallel rows of 14–20 pali, septula open posteriorly, narrow subtriangular to elliptical.
Larvae of X. braunsi sp. nov. differ from all known larvae of Cetoniinae sensu stricto by the absence of the hypopharyngeal truncate process. From those of Meridioclita capensis and Heteroclita haworth larvae of X. braunsi sp. nov. differ by the absence of the minute tip on the cylindrical pretarsi. Also, they can be separated from those of Ichnestoma rostrata by their only slightly emarginate frontal suture, by the trapezoidal clypeus and by the number of posterior frontal setae.
Material studied. Eight last instar larvae: EC, Sarah Baartman District (Dr Beyers Naudé Municipality), 15 km NW of Willowmore, 870 m, 5.I.2017. Fifteen larvae were collected in soil with organic debris under a shrub of Psilocaulon sp. (Mesembryanthemaceae), where also adult speciemens were found together with larvae. No termites or ants were observed in the vicinity of the shrub. The identity of the larvae has been confirmed by rearing the remaining specimens to adulthood and by molecular match (COI) with adult specimens.
General body.
Scarabaeiform (Fig.
Head capsule
(Fig.
Cranial chaetotaxy of the larva of Xiphoscelis braunsi sp. nov. Abbreviations: AAS = setae of anterior frontal angle; ACS = anterior clypeal setae; AES = anterior epicranial setae; AFS = anterior frontal setae; DES = dorsoepicranial setae; LCS = lateral clypeal setae; LES = lateral epicranial setae; LFS = lateral frontal setae; PES = posterior epicranial setae; PFS = posterior frontal setae.
Group of setae | Epicranium | Frons | Clypeus | |||||||
---|---|---|---|---|---|---|---|---|---|---|
DES | PES | AES | LES | PFS | LFS | AFS | AAS | ACS | LCS | |
Long/medium setae | 5–9 | 1–5 | 1 | 9–15 | 2–30 | 1 | 1 | 1 | 1 | 2 |
Minute/short setae | 13–23 | 3 | – | 3–6 | – | – | – | – | – | – |
Antennae
(Fig.
Labrum
(Fig.
Epipharynx
(Figs
Mandibles
(Fig.
Maxilla
(Figs
Hypopharyngeal sclerome
(Figs
Ligula
(Figs
Thorax
(Fig.
Abdomen
(Figs
Raster
(Fig.
Apart from a number of characters that have led to believe that the genus Xiphoscelis may occupy a very primitive position in the phylogenesis of the Cetoniinae, species of the genus also exhibit a most unusual feature, represented by the extreme hypertrophy of structures in their metalegs. This is very prominent in males but less so in females and includes the femora, spurs and especially the inner spines, which are truly extraordinary in their thickness, length and sharpness in virtually all the species of the genus, possibly with the exception of X. namibica sp. nov. (Fig.
Males of the largest species, X. schuckardi (Fig.
As reported earlier, all species of the genus are restricted to the south-western arid and semi-arid regions of southern Africa (
Following the description of the two new species here, it is evident that X. schuckardi is restricted to the lowlands of the South African west coast, from just north of Cape Town to the Namaqualand area of the Northern Cape (Fig.
The frequently reported association with the termite Microhodothermes viator (
The Xiphoscelidini, or more accurately Xiphoscelidina (refer to
Based on the description of the larval morphology of X. braunsi, we can conclude that the larvae of the genus Xiphoscelis are characterised by a remarkable subset of morphological characters: i.e., long and dense chaetotaxy of cranium; reduction of sense cone and plate-shaped sclerite of epipharynx; presence of an external tooth on lateral mandibular margin; shape and size of lacinial unci; shape of hypopharyngeal scleroma (especially the reduction of the truncate process); and shape of pretarsi. To allow an easy comparison of the morphological similarities of Xiphoscelis with other Cetoniinae larvae, a parsimony analysis of the larval morphology of 13 cetoniine genera, including Xiphoscelis, was performed (Fig.
Strict consensus topology of 13 representatives of the Cetoniinae, based on a larval morphology dataset. Character states are marked on clades, with character number above each circle and state number below it; black circles indicate unique evolutionary events, while white circles denote reversals or parallelisms. Partitioned Bremer Support (PBS) values above 50% are indicated. The analysis was performed in order to highlight morphological similarities among representatives of distinct cetoniine clades.
The outcome shows that the larva of X. braunsi sp. nov. falls within a clade shared with M. capensis and H. haworth (Fig.
In the early taxonomic classifications of the Cetoniinae, the genera Xiphoscelis, Ichnestoma, Heteroclita and Meridioclita were all classified under the Xiphoscelidini (e.g.,
The following museum curators, researchers and owners of private collections are thanked for kindly providing photos, data and material for analysis: Riaan Stals & Werner Strumpher (
Character states of the 77 morphological characters used in the analysis of larval similarity of 13 Cetoniinae genera (refer also to Fig.
1. Color of cranium: 0 – black to dark brown, 2 – brown, 3 – light brown to yellowish
2. Surface of cranium: 0 – smooth or with very fine microsculpture, 1 – light to medium microsculpture, 2 – with coarse microsculpture
3. Texture of cranial microsculpture: 0 – more or less pitted, 1 – more or less linear
4. Chaetotaxy of cranium: 0 – light (short and/or very sparse setae), 1 – “standard Cetoniinae” type, 2 – dense (setae numerous and/or extraordinary long)
5. Anterior end of epicranial suture: 0 – originating at posterior end of frontal sutures, 1 – extending into frons, 2 – irregular, feebly visible
6. Epicranial suture: 0 – distinct, 1– indistinct
7. Frontal suture: 0 – straight, 1 – straight, slightly emarginate, 2 – convex, 3 – concave, 4 – bisinuate
8. Frontal suture: 0 – distinct, 1 – indistinct
9. Posterior angle of frontal sutures: 0 – acute, 1 – right, 2 – obtuse
10. Widest part of cranium: 0 – at base of antennae, 1 – at approx. one-third of cranium, 2 – at middle of cranium
11. Number of antennomeres: 0 – 3, 1 – 4, 2 – 5
12. Sensory spot on third antennomere: 0 – present, 1 – absent
13. Third antennomere: 0 – with well-developed ventral and apical projection, 1 – ventroapical projection only feebly developed, 2 – ventroapical projection absent
14. Stemmata: 0 – present and pigmented, 1 – present without pigment, 2 – reduced, 3 – absent
15. Setae on antennomeres: 0 – present (at least on a single joint), 1 – absent
16. Number of sensory spots on dorsal side of antenna: 0 – 1, 1 – 2 or 3, 2 – 4 or more, 3 – 1 or 2 (if both character states 1 and 2 are present)
17. Number of sensory spots on ventral side of antenna: 0 – 0 to 3, 1 – 4 and more, 2 – 3 to 5 (if both character states 1 and 2 are present)
18. Shape of clyepus: 0 – trapezoidal, 1 – rectangular
19. Preclypeus: 0 – about ½ of area of entire clypeus, 1 – about 1/3 of area of entire clypeus, 2 – about ¼ of area of entire clypeus, 3 – less than ¼ of area of entire clypeus
20. Anterior and lateral frontal setae: 0 – medium long to long, 1 – minute, 2 – absent
21. Posterior frontal setae: 0 – absent or minute, 1 – one medium long to long seta, 2 – 2 or more medium to long setae, 3 – both long and short setae
22. Number of long lateral epicranial setae: 0 – absent, 1 – 1 or 2, 2 – 3 or more
23. Dorso-epicranial setae: 0 – 1 to 3 medium to long setae together with some minute setae, 1 – only minute setae, 2 – only medium to long setae, 3 – both long and short setae in abundance
24. Shape of anterior labral margin: 0 – asymmetric, 1 – slightly convex, 2 – strongly convex, 3 – slightly concave, 4 – trilobed
25. Shape of sensory cone (second nesium): 0 – ovoid, 1 – conical, 2 – not applicable
26. Size of sensory cone (second nesium): 0 – well developed, 1 – more or less reduced, 2 – absent, indistinct
27. Size of third nesium (plate-shaped sclerome): 0 – largely reduced, 1 – feebly sclerotized, reduced, 2 – absent
28. Shape of third nesium (plate-shaped sclerome): 0 – transverse, 1 – present on right and in front of sensory cone (shape of upturned and reversed capital letter “L”), 2 – present on left side of sensory cone, 3 – present in front and on both sides of the sensory cone (shape of reversed capital letter “U”), 4 – not applicable
29. Dexiotorma: 0 – straight and wide, 1 – bent at inner end, 2 – straight, narrow, 3 – reduced
30. Right pternotorma: 0 – present, 1 – absent
31. Laeotorma: 0 – present, 1 – reduced, 2 – only feebly developed
32. Left pternotorma: 0 – present, 1 – absent
33. Number of sensilla on haptolachus: 0 – four, 1 – five, 2 – six or more
34. Clithra: 0 – absent, 1 – only single clithrum present, 2 – two clithra present
35. Setae of acanthoparia: 0 – directly on lateral margin, with swollen base, 1– directly on lateral margin, without distinctly swollen base, 2 – away from margin and slightly towards center of epipharynx
36. Pedium: 0 – absent, 1 – reaching < 25% of central part of epipharynx, 2 – reaching 25–50% of central part of epipharynx, 3 – reaching 51–75% of central part of epipharynx, 4 – reaching > 75% of central part of epipharynx
37. Chaetoparia of epipharynx: 0 – reaching only to level of laeo- and pternotorma, 1 – extending posteriorly behind level of laeo- and pternotorma
38. Crepis: 0 – well developed, 1 – reduced, 2 – absent
39. Zygum: 0 – flat or transverse, 1 – high and conical, 2 – extending into tylus, 3 – absent
40. Sensilla of zygum: 0 – not grouped on projection, 1 – grouped on distinct projection, 2 – intermediate
41. Number of setae on zygum: 0 – six, 1 – seven, 2 – eight and more, 3 – intermediate
42. Transverse row of prominent setae on zygum: 0 – arcuate, 1 – right angled, 2 – angulate, 3 – not applicable
43. Setae at inner margin of zygum: 0 – approximately of same size and shape as setae in prominent row, 1 – different from setae in transverse row, 2 – not applicable, 3 – absent
44. Setae of chaetoparia: 0 – all setae of approximately same size and shape, 1 – chaetoparia with several rows of longer and stouter setae
45. Number of teeth on scissorial area of left mandible: 0 – two, 1 – three, 2 – four
46. Last two teeth of left mandible: 0 – fused, 1 – separated
47. Number of teeth on scissorial area of right mandible: 0 – two, 1 – thee, 2 – four
48. Second and third teeth of left mandible (number from the apex): 0 – separated, 1 – fused, 2 – not applicable
49. Mandibular teeth: 0 – in single plain, 1 – second and third tooth oriented dorsoventrally
50. Dorsomolar setae: 0 – absent on both mandibles, 1 – absent on left mandible, 2 – present on both mandibles
51. Tooth on external mandibular margin: 0 – present, 1 – absent
52. Ventromolar setae: 0 – present on both mandibles, 1 – absent on both mandibles
53. Mandibular shape: 0 – falcate at least at apex, 1 – more or less straight
54. Mandibular stridulatory area: 0 – well developed with transverse ridges on entire area, 1 – developed, ridges only feebly developed, 2 – absent
55. Shape of stridulatory area: 0 – oval, 1 – oblong, 2 – intermediate
56. Number of stridulatory ridges on mandible: 0 – less than 10, 1 – 10 to 20, 2 – 20 or more, 3 – intermediate
57. Extent of ventral asperities: 0 – reaching towards ventral process of mandibles, 1 – not reaching towards ventral process of mandibles, 2 – reaching towards ventral process only on left mandible, 3 – reaching towards ventral process only on right mandible
58. Shape of ventral asperities (obr. 17): 0 – small tubercle, 1 – tooth-like process
59. Hypopharyngeal scleroma: 0 – symmetrical, without processes, 1 – asymmetric, with large process on right side
60. Number of maxillar stridulatory teeth: 0 – 0, with indistinct transverse ridges, 1 – 4 to 9, reduced, 2 – 4 to 9, well developed, 3 – 10 to 20
61. Shape of stridulatory teeth: 0 – short, with swollen base, 1 – long and sharp, 2 – obtuse tubercles, 3 – more than one type, 4 – intermediate shape
62. Lacinia and galea: 0 – fused, forming mala, 1 – separated
63. Number of lacinial unci: 0 – three, 1 – two, 2 – one or none
64. Shape of legs: 0 – short and stout, 1 – slender and long
65. Relative length of legs: 0 – each pair increasing in size from protorax to metathorax, 1 – all pairs (sub)equal
66. Legs: relative length of tibiotarsus and trochanter joints: 0 – trochanter longer than pretarsus, 1 –- trochanter shorter than pretarsus
67. Shape of pretarsus: 0 – falcate, pointed with pair of setae, 1 – conical, setae in apical half, 2 – conical, with numerous setae and small claw on apex, 3 – narrow, pointed and slightly bent, with swollen base and a pair of basal setae, 4 – almost reduced with pair of setae
68. Relative size of pretarsi: 0 – all pairs equal in length, 1 – increasing in size from prothorax to metathorax, 2 – decreasing in size from prothorax to metathorax.
69. Tibiotarsi: 0 – all pairs equal in shape and size, 1 – first two pairs similar, last pair of different size, 2 – all three pairs same in size but of different shape
70. Size of spiracles: 0 – all spiracles (sub)equal, 1 – first or first and second abdominal spiracles distinctly smaller than thoracic spiracles, 2 – size decreasing posteriorly
71. Shape of thoracic spiracles: 0 – concealed, arms of respiratory plate almost closed, 1 – open but opening between arms narrow, 2 – open with arms of spiracular plate well separated
72. 9th and 10th abdominal segments: 0 – entirely fused, 1 – separated, 2 – partially fused
73. Palidium: 0 – present, 1 – absent
74. Size of Palidium: 0 – absent, 1 – short, 2 – of medium length, reaching middle of last abdominal segment, 3 – long, reaching almost end of last abdominal segment
75. Hammate setae: 0 – present, 1 – absent
76. Setae of last abdominal segment (excluding raster): 0 – long, hair-like, 1 – short, 2 – long, dorsoventrally flattened at distal margin, 3 – absent, 4 – mixture of long hair-like and short stout setae
77. Chaetotax (general appearance) of larva: 0 – hairy, with numerous long setae. 1 – dense, covered in numerous short setae, 2 – standard “Cetoniinae” type, 3 – sparse