Carabus variegatus Fabricius, 1792 (= Carabus serrator Forskål, 1775).

The Graphipterus serrator group is included in the genus Graphipterus based on the following combination of characters:

Clypeus concave at anterior margin, posteriorly well separated from front; labrum wide and short, with well-developed microsculpture and six setiferous pores. Mandibles broad at the base, sharp and strongly curved at tip; labial and maxillary palps long and slender, glabrous with exception of distal end of segments which bear a few hairs; last palpal segments slightly thicker than penultimate ones.

Pronotum transverse and cordiform, slightly convex, usually ornamented by colored scales at the lateral bead, disc with or without scales. Anterior and posterior angles obtuse.

Scutellum triangular, small and short, often hidden by the pronotal base. Flightless. Elytra wide and oval, slightly convex, coalesced along suture, humeri completely rounded; surface covered by dense or sparse scales, white scales creating longitudinal stripes on the radial field and spots on the disc; apex almost truncate. Pygidium not covered by elytra, last visible tergite with colored scales.

Legs long, usually black or brown, protibia with clypsetae (antenna cleaner) and dark parallel spurs, as long as ¾ of protarsomere 1. Mesotibia with two long and thin not serrated spurs, metatibia with one long and thin not serrated spur and one shorter, wide and obtuse spur. Claws of all legs long and smooth on median margin.

Holotype: ♂ (Blue label, black handwritten): <Barthelemyi. Solier/. in Barbaria. Tunis. D. Barthelemyi>. (White label with brown margin, brown letters, handwritten): <EX Musaeo/Chaudoir>. (Red label, black letters, type written): <TYPE>. Deposited in NHMB, Chaudoir collection [examined].

Medium-sized species with grayish or yellowish scales usually cover the elytra and sometimes also on the pronotum. Elytra pattern rarely visible with six lateral margin extensions and 18–24 isolated white circular to elongated spots occur on elytra.

BL male: 13.0–17.0 mm, average 15.8 ± 1.5 mm; BL female: 15.5–17.0 mm, average 16.3 ± 0.6 mm. Grayish with elytral white blurred spots and extensions.

Head slender: HW/PW: 0.72; EYL: 1.1-1.4 mm; EYL/EL: 0.15. Mentum without teeth (Fig. 3a). Frontal ridge reduced. In male, apical white frons stripes slenderer than exposed frons (cf. Fig. 4a).

Pronotum wide; PL/PW: 0.63; BPW/BPW/PW: 0.6; posteromedially concave and with white lateral margin, as wide as antennomere I long: white slushy scales cover disc sometimes.

Elytra wide, elytron margin almost continuously rounded from humeri to posterolateral angles; EL: 7.1–8.8 mm, average 8.2 mm; EW: 6.5–7.8 mm, average 7.3 mm; EL/EW: 1.1. Elytra longitudinally flat, usually with grayish scales, disc visible between scales (Fig. 6c); extensions and lateral margin blurred. Lateral margin nearly as wide as antennomere I long and with six extensions; extension I usually elongated; white posterior margin almost touches suture at apex. Disc with 18–24 rounded, usually elongated spots, anterior pair of spots elongate, as wide as extension I; posterior pair of spots rounded, located toward suture; round spots located posterior to third extensions laterally in imaginary lateral line as posterior spots. Apical sinuation slightly developed to straight, apex not protuberant, broadly rounded, especially medially (Fig. 7c). Suture inconspicuous.

Legs medium; MTIL: 4.5–6.5 mm, average 5.7 mm; El/MTIL: 1.7. Metatibial secondary spur brown. MTAL: 3.5–4.4 mm, average 4 mm; MTAL/MTIL: 0.8. Claws of hind legs brown at base.

Median lobe of aedeagus with short unbent tip (Fig. 9a).

Distinguished from all other species of the G. serrator group by white lateral margins merged at the posterior margin of the pronotum. Median lobe of aedeagus with short, straight tip.

Unknown. The species was found exclusively in coastal dune habitats.

Graphipterus barthelemyi lives in sympatry with G. luctuosus in Tunisia.

Restricted to north-east Tunisia (Fig. 17).

The restricted distribution range of the endemic species and the decline of the coastal sandy habitat as a result of increasing anthropogenic pressures (e.g., tourism activities, urbanization, etc.) threaten at least the long-term survival of the species.

Both Basilewsky (1977) and Lorenz (2005) note in error that G. barthelemyi was described by Dejean (1831).

Lectotype: ♂ (Blue label with black margin, black handwritten): <Heydeni Krtz./ luctuosus Guer./Tripolis. Oued>. (White label, print black): <Coll. Kraatz>. (White label, black print): <Tripolis>. (White label, black print): G. serrator/heydeni Kr>. (Green label, black print): <Muncheberg/Col – 01309>. (White label, black print): serrator/ heydeni Kz./P. Basilewsky det., 1975>. Deposited in ZSM [examined].

Paralectotype: two specimens – ♂, ♀ (White label, black black handwritten): <Tripolis>. (White label, black handwritten): <Call. Kraatz>. (White label, black handwritten): <Muncheberg/Col – 01310/01311>. Deposited in ZSM [examined]. Lectotypes and paralectotypes herewith designated.

Large species with 18–26 isolated white round spots on elytra, anterior and posterior discal spots larger than other spots; four marginal extensions, anterior extension triangular; median lobe of aedeagus with ventrally bent apex.

Graphipterus heydeni resembles G. valdanii from which it differs mainly by the following characters: G. heydeni: mentum with two teeth, margin between them clearly concave; EL/EW rounded (1.24); 18-26 spots on elytra; claws of hind legs dark; metatibial secondary spur brown. In Graphipterus valdanii, mentum with two teeth, margin between them slightly convex in middle; EL/EW elongated (1.31); 18-26 spots on elytra; claws of hind legs brown; metatibial secondary spur dark. Graphipterus heydeni also resembles G. magnus sp. n. from which it differs mainly by the following characters: G. heydeni: elytra shape oval; four elytral marginal extensions; anterior and posterior elytral spots larger than all other spots; median lobe of aedeagus with stout with ventrally bent tip. G. magnus sp. n.: elytra shape rounded; six elytral marginal extensions; all elytral spots with similar size; median lobe of aedeagus elongated with ventrally bent tip.

BL male: 17.1–20.9 mm, average 18.9 ± 1.6 mm; BL female: 18–20 mm, average 19.4 ± 1.4 mm.

Head slender; HW/PW: 0.71; EYL: 1.5–1.9 mm; EYL/EL: 0.16. Mentum with two teeth and concavity between them (Fig. 3b). Frontal ridge absent. In male, apical white frons stripes slenderer than exposed frons (Fig. 4a).

Pronotum slender; PL/PW: 0.65; BPW/PW: 0.7; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra oval, humeri rounded; EL: 10.0–12.1 mm, average 10.9 mm; EW: 8.4–9.9 mm, average 9.16 mm; EL/EW: 1.2. Lateral cross section convex. Black scales dense, disc not visible between them (Fig. 6a). White lateral margin almost as wide as antennomere I long and with four extensions; extension I triangular with rounded angels, slightly wider at margin of elytra, slightly elongated, wider and shorter than extension II; the latter one elongated at third quarter of elytra, imaginary line connecting the media ends of the extensions I and II parallel to the suture; white posterior margin forms a gap at suture, wider than lateral margin. Disc usually with 18–26 rounded spots; anterior pair of spots rounded, as wide as extension I, usually smaller than posterior spots, larger than spots on mid disc; mid disc spots usually asymmetrically smeared. Posterior pair of spots rounded, one or two small spots located laterally to posterior spots. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a). Suture inconspicuous.

Legs long; MTIL: 5.3–7.0 mm, average 6.5 mm; El/MTIL: 1.7. Metatibial secondary spur brown. MTAL: 4.4–5.2 mm, average 4.8 mm; MTAL/MTIL: 0.7. Claws of hind legs black at base.

Median lobe of aedeagus with apex bent ventrally (Fig. 9b).

Unknown.

Graphipterus heydeni lives in sympatry with G. luctuosus around Tripoli, Libya, and might live in sympatry with G. rotundatus in this region. It also lives in sympatry with G. piniamitaii sp. n. in Nefzaoua region in Tunisia.

Western Lybia (Tripolitania) and western Tunisia (Nefzaoua) (Fig. 18).

The restricted distribution range of the endemic species and the decline of the coastal sandy habitat as a result of increasing anthropogenic pressures (e.g., tourism, urbanization etc.) threaten at least the long-term survival of the species.

This taxon was first described by Guérin-Méneville (erroneously as luctuosus Dej.). As Kraatz already noted, Guérin-Méneville's and Dejean's specimens do not belong to the same taxon, and Kraatz substituted the name heydeni Kraatz, 1890 as a new replacement name (nomen novum) for the already available name luctuosus Guérin-Méneville. However, Kraatz never fixed the holotype (Jäger, pers. comm.), following the requirement of Article 72.2 (ICZN 1999). The type series of heydeni comprises three individuals from Tripoli (Kraatz 1890: 77) and not seven (holotype and six paratypes) as indicated by Basilewsky (1977: 451). The beetles were collected by Quedenfeldt, as this circumstance was indicated by Kraatz in the original description. These individuals have been transferred to the DEI (Kraatz was the director of this institution) and the syntypes are still preserved there. A lectotype is designated, labeled with a handwritten card indicating the taxon’s name, the name of the location, Tripoli, and the initial letters of the collector (Fig. 21c). The above description is based primarily on the three syntypes. The misinterpretation of the type material by Kraatz led Basilewsky to an incorrect interpretation of heydeni Kraatz. Consequently the distribution map given by Basilewsky (1977: page 450) is also incorrect.

Holotype: ♂ (Green label, black handwritten): <Luctuosus. mihi/ h. in Barbaria. Tripoli>. (White label, black typewritten): <P. Bedel/Visit 1905>. (White label with brown margin, brown typewritten): <EX Musaeo/Chaudoir>. (Red label, black typewritten): <TYPE>. Deposited in NHMB, Chaudoir collection [examined]. Neotype: ♂ (White label, black handwritten): <Tripolis>. (White label, black handwritten): <Coll: Kraatz>. (Green label, black handwritten): <DEI Muncheberg/ Call-01342>. (Red label, black typewritten): <Neotypus/Graphipterus reichei/ Guérin-Méneville, 1859/ des. I. Renan, 2018. Neotype: ♂ (White label, brown handwritten): <intermedius/♂ reichei>. (White label, brwon handwritten): <Tripolis>. (White label, black handwritten): <Call: Kraatz>. ♂ (Green label, black handwritten): <DEI Muncheberg/Call-01343>. (Red label, black typewritten): <Neotypus/Graphipterus intermedius/Guérin-Méneville, 1859/des. I. Renan, 2018>.

Medium-sized species with 22–30 isolated white, usually elongated elytral spots, six very short marginal extensions, and a series of 8–12 elongated spots along suture form a broken line. Median lobe of aedeagus with apex slightly bent ventrally.

Graphipterus luctuosus resembles G. peletieri from which it differs mainly by the following characters: G. luctuosus: apical white frons stripes, wider than exposed frons; six elytral extensions; 8–12 elongated spots along suture of elytra; elytral suture conspicuous; white posterior margin almost attached; median lobe of aedeagus with bent tip. G. peletieri apical white frons stripes slenderer than exposed frons; four elytral extensions; elytral spots scattered; elytral suture not conspicuous; white posterior margin forming gap; median lobe of aedeagus with unbent tip. Graphipterus luctuosus resembles also G. rotundatus from which it differs mainly by the following characters: G. luctuosus: 8–12 elongated spots along suture of elytra; scales of elytral disc brown, disc visible between; metatibial secondary spur brown. G. rotundatus: elytral spots scattered; scales of elytral disc black, disc not visible between them; metatibial secondary spur dark, not darker than the elytral scales.

BL male: 15.0–17.5 mm, average 15.8 ± 1.5 mm; BL female: 15–18 mm, average 16.3 ± 0.6 mm.

Head medium; HW/PW: 0.74; EYL: 1–1.6 mm; EYL/EL: 0.15. Mentum with two or three teeth. Frontal ridge absent. In male, apical white frons stripes wider than exposed frons, (cf. Fig. 4b).

Pronotum cordiform; PL/PW: 0.64; BPWBPW/PW: 0.61; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra oval, humeri rounded; EL: 7.3–9.9 mm, average 8.9 mm; EW: 5.8–8.4, average 7.7 mm; EL/EW: 1.16. Lateral cross section quite flat suture conspicuous. Scales brown, disc visible between them (Fig. 6b). White lateral margin narrow, as wide as half antennomere I long and with six extensions, rarely four; extension I usually elongated, sometimes constricted at the base to the lateral margin; extension II elongated, constricted or absent; extension III elongated; white posterior margin almost as wide as the lateral margin, gap at suture smaller than lateral margin or even absent. Disc usually with 22–30 mostly elongate small spots; anterior pair of spots slightly elongate, wide as extension I, lateral spots rounded, adjacent or sometimes fused to extension II, posterior pair of spots rounded, slightly larger than others, located toward suture, round spots slightly smaller than all others, located posteriori to third extensions and laterally to posterior spots; a series of 8–12 elongated spots along the suture. Apical sinuation developed, apex slightly protruded, strongly rounded (Fig. 7b).

Legs medium; MTIL: 4.5–5.9 mm, average 5.4 mm; El/MTIL: 1.7. Metatibial secondary spur brown. MTAL: 3.4–4.7 mm, average 4.4 mm; MTAL/MTIL: 0.8. Claws of hind legs brown at base.

Median lobe of aedeagus with ventrally bent tip (Fig. 9c).

Unknown.

Graphipterus luctuosus lives in sympatry with seven other species: G. peletieri in north-west Algeria, G. heydeni in Tripoli region, G. valdanii in north Algeria, G. rotundatus in Tunisia and Algeria, G. stagonopsis in the Ghardaia region, Algeria, G. piniamitaii sp. n. in Tunisia, and G. barthelemyi in north-east Tunisia.

Graphipterus luctuosus presents the widest distribution range of the group: from Laghouat, more than 300 km inland Algeria to the arid and semi-arid regions of north-east Algeria, over most of the Tunisian coast and east up to Sirte on the Libyan coast (Fig. 16).

The species does not seem to be endangered as it apparently lives in numerous habitats. Consequently, it might not be so strongly affected by human activities.

Graphipterus reichei and G. intermedius have been described by Guérin-Méneville (1859) as variants of G. multiguttatus. Unfortunately, the type material of both Guérin-Méneville's taxa has have been lost. Neotypes for both taxa are designated. Based on the original description and the type locality, Tripoli, the only other known species from the type locality is G. heydeni, which is clearly different in elytral pattern and body length. Basilewsky (1977), Huber and Marggi (2017) and Lorenz (2005) ranked both taxa as synonyms of G. luctuosus.

Holotype: ♂ (White label, black handwritten): <23.II 1942/Buq Buq/P.J. Gent/Egypt> (White label, black typewritten and black handwritten): <Brit. Mus./1952-180> (White label, black typewritten): <BMNH {E}/UIN989817>. (ae) Deposited in BMNH [examined].

Paratypes: (2 ♂), Egypt, Buq Buq: 14.11.1942, P.J. Gent, {E}/UIN989815 (♂); Egypt, E. of Buq Buq, 14.11.1942, P.J. Gent, {E}/UIN989815, Brit. Mus.952-180 (♂) (BMNH).

Large species with 20–24 white rounded and elongated elytra spots; six white marginal extensions, extension I elongated. Elytra wide, lateral margin strongly and continuous rounded. Aedeagus elongated, thin and with apex slightly bent ventrally (Fig. 9d).

Graphipterus magnus sp. n. resembles G. heydeni from which it differs mainly by elytra shape and pattern, and aedeagus shape (see comparisons in G. heydeni).

BL male: 18.3–20.1 mm, BL female: unknown. Average 19.4 ± mm.

Head slender; HW/PW: 0.72; EYL: 1.7 mm; EYL/EL: 0.17. Frontal ridge well developed. In male, apical white frons stripes slenderer than exposed frons (cf. Fig. 4a). Pronotum cordiform; PL/PW: 0.62; BPW/BPW/PW: 0.68; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra wide, rounded, rounded-like, humeri strongly narrowed; EL: 9–10.7 mm, average 9.7 mm; EW: 8.5–9.0 mm, average 8.7 mm; EL/EW: 1.1. Lateral cross section quite flat. Scales black, disc not visible between them (cf. Fig. 6a). White lateral margin nearly as wide as antennomere I long and with six extensions; extensions I slightly elongated, wider close to the margin; extensions II and III in front of middle. White posterior margin as wide as lateral margin or wider, sutural gap slenderer than lateral margin. Disc usually with 20 (rarely up to24) spots; anterior pair of spots rounded, wider than extension I, 6–8 spots adjacent elongated and parallel to suture, posterior pair of spots rounded, additional 1–3 small spots frequently present laterally to posterior ones. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a). Suture conspicuous.

Legs long; MTIL: 6.2–6.8, average 6.5 mm; El/MTIL: 1.53. Metatibial secondary spur brown. MTAL: 5.2mm; MTAL/MTIL: 0.8. Claws of hind legs brown at base.

Median lobe of aedeagus long and thin with apex hardly bent ventrally (Fig. 9d).

The species name is derived from Latin (magnus) and refers to the large body size.

Unknown.

No co-occurring species.

Unknown.

The only known records are from Buq Buq in north-east Egypt (Fig. 19).

Holotype: ♂ (White label with pencil handwritten) < luctuosus/(uc)>. (White label with black typewritten and handwritten): <OFFICE NATIONAL ANTIACRIDIEN/Azefal Mauritania/13 Fevirer 1950/J. Leroux>. (red label): <Holotype> (ae). Deposited in Colas collection, NHMB.

Paratypes: (3 ♂, 1♀), Azefal Mauritania: 13 Fevrier 1950, J. Leroux (♂) (NHMB, Colas collection): Mauritanie [Mauritania]: Chingvetti, 3.1951, L. Dekeyser and A. Villiers (2♂ (1-ae), ♀) (MRAC).

Medium-sized species with (18–) 22 white, mostly elongated spots on elytra, anterior and posterior spots larger than other spots; six marginal extensions, extension I usually triangular. Median lobe of aedeagus with short apex unbent ventrally (somewhat similar to that of G. barthelemyi).

Graphipterus mauretensis sp. n. resembles G. piniamitaii sp. n., from which it differs mainly by the following characters: G. mauretensis sp. n.: (18–) 22 spots on elytra; anterior and posterior elytral spots larger than all other spots; apical sinuation and apex developed and slightly protruded; median lobe of aedeagus with short bent tip. G. piniamitaii sp. n.: 24 spots on elytra; only posterior elytral spots larger than all other spots; apical sinuation and apex strongly developed and protruded; median lobe of aedeagus with ventrally bent tip.

BL male: 15.1–17.5 mm, average 16.6 ± 1.1 mm. Females were not available.

Head medium; HW/PW: 0.76; EYL: 1.3–1.5 mm; EYL/EL: 0.16. Mentum with two teeth (cf. Fig. 3b). Frontal ridge reduced. In male, apical white frons stripes wider than exposed frons (cf. Fig. 4b).

Pronotum cordiform; PL/PW: 0.66; BPW/PW: 0.63; posteriomedially concave and without white margin; white lateral margin as wide as antennomeres I+II long.

Elytra relatively elongated oval humeri slightly narrowed; EL: 8.6–9.7 mm, average 9.2 mm; EW: 6.8–8.0 mm, average 7.4 mm; EL/EW: 1.2. Lateral cross section quite flat. Dense black scales, disc not visible between them (Fig. 6a). White lateral margin as wide as half of antennomere I long and with six extensions; extension I triangular; extension II shorter than extension III. White posterior margin as wide as lateral margin, sutural gap slenderer than lateral margin. Disc usually with 18–22 rounded to elongate spots; anterior spot elongated, as wide as extension I, anterior and posterior spots larger than all other ones, posterior one rounded. Apical sinuation developed, apex slightly protruded, strongly rounded (Fig. 7b). Suture conspicuous.

Legs long; MTIL: 5.3–5.9 mm, average 5.7 mm; El/MTIL: 1.61. Metatibial secondary spur brown at base, MTAL: 3.8–4.5 mm, average 4.2 mm; MTAL/MTIL: 0.74. Claws of hind legs brown at base.

Median lobe of aedeagus with short apex, unbent ventrally (Fig. 9e).

The species name is derived from ancient Latin (Mauretania, -ensis).

Unknown.

No co-occurring species.

As we found in all collections only nine specimens of G. mauretensis sp. n., our knowledge of its distribution range is limited. Known from central coast of Mauritania to more than 400 km inland to Glebat el M’Boza Adrar (Fig. 16).

Unknown.

Lectotype: ♀ (blue label, black handwritten): <minutus. m/ h. in Egypt>. (blue label, black handwritten): <Olivier>. (White label with brown margin, brown typewritten): <EX Musaeo/Chaudoir>. (Red label, black typewritten): <TYPE>. Deposited in BMNH, Chaudoir collection [examined].

Paralectotypes: ♀ (blue label, black handwritten): <Graphipterus {minutus. Ol./minutus. Dej./Egypt. C. Olivier>. (Green circular label with black margin, black typewritten): <COLLECTION/OLIVIER/TYPE>. Deposited in BMNH, Chaudoir collection [examined]. ♀ (Green circular label with black margin, black typewritten): <COLLECTION/OLIVIER/TYPE>. Deposited in BMNH, Olivier collection [examined].

Additionally, two syntypes are deposited in Chaudoir’s collection, NHMB [examined].

The two subspecies of G. minutus are distinguished from all other species of the G. serrator group by smaller size, lack of the stridulatory structure, unique pronotum shape (G. serrator group excluding G. minutus: BPW/PW: 0.6-0.7, G. minutus: BPW/PW: 0.46) and flat tip of median lobe.

Figure 10. 

Spectrograms of two Graphipterus species: G. serrator and G. multiguttatus.

Graphipterus minutus minutus differs from G. minutus goryi mainly by the following characters: G. minutus minutus: frontal ridge not developed; 36–40 spots on elytra; two elytra marginal extensions; rounded and separated spots along median suture. G. minutus goryi: 28–30 spots on elytra; six elytra marginal extensions; elongated and fused spots along median suture.

BL male: 10.3–13.5 mm, average 12 ± 1.2 mm; BL female: 10.5–15.2 mm, average 13.1 ± 1.9 mm;

Head wide; HW/PW: 0.77; EYL: 1–1.8 mm; EYL/EL: 0.15. Frontal ridge slightly developed. Male with two short parallel frontal stripes of white scales usually diverging apically, became wispy, not reach the level of supraorbital setiferous pores. Mentum usually with two pronounced teeth (Fig. 3e). Pronotum strongly cordiform PL/PW: 0.54; BPW/PW: 0.46; posteromedially flat and without white margin; white lateral margin as twice as antennomere II long.

Elytra almost rounded, humeri stringly rounded, lateral margin continuously rounded; EL: 5.3–7.5 mm, average 6.6; EW mm: 4.8–7.6 mm, average 6.1 mm; EL/EW: 1.16. Suture inconspicuous. Scales black, disc not visible between them (cf. Fig. 6a). Lateral cross section convex. Apical sinuation almost lacking, apex almost absent, not rounded (Fig. 7d). White lateral margin usually nearly as wide as antennomere I long and usually with two extensions; extension I elongated from humeri posteriorly; extension II usually absent, sometimes indistinct wider section of lateral margin at its middle. White posterior margin becomes narrower toward the tip, usually disappearing in front of it; gap at suture wider than lateral margin. Disc with 36–40 mostly rounded spots; usually 12, sometimes ten or 14 rounded to elongated, not fused spots located parallel to suture; anterior spot as wide as extension I. Stridulatory structure absent.

Legs short; MTIL: 2.54–4.0 mm, average 3.3 mm; El/MTIL: 1.9 mm. Metatibial secondary spur brown, MTAL length: 2.5–3.3 mm, average 2.9 mm; MTAL/MTIL: 0.85. Claws of hind legs brown at base.

Median lobe of aedeagus with wide and flat tip (Fig. 9f).

Sparse populations in arid habitats with hallow sand dunes, and scant shrubs landscape (Fig. 11).

Figure 11. 

Habitat of Graphipterus minutus minutus: Shallow sand dunes in Wadi Ram, Jordan.

No co-occurring species.

Syria, east and south Jordan, north Saudi Arabia, Iraq, and Iran (Fig. 19).

The species does not seem to be endangered as it has a wide distribution range that is not strongly affected by human activities.

The type location of G. m. minutus, Egypt, is probably a labeling mistake. Only four specimens of this species were found with labels from Egypt; the three syntypes from Olivier’s collection and one specimen deposited in BMNH, collected by Bowring. Even though Olivier had a large amount of material from Egypt and Bowring collected in Egypt, we are convinced that G. minutus does not occur in Egypt: all the known populations from collections and field observations are from Asia and not from Africa. Furthermore, no specimen has been ever collected in Israel, despite intensive collecting in the potential habitats. Basilewsky (1977) noted that although several researchers had contended that G. minutus does exist in Egypt, they were wrong, but he does not refer to the problem of types.

By applying other species concepts (e.g., Evolutionary or Phylogenetic Species Concept, Claridge et al. 1997) or by using another approach to delineate species, the two taxa minutus and goryi might be ranked as two species. However, our numerical approach to delineate species results in a value for both minutus and goryi that is clearly below the threshold of the least differentiated sympatrically occurring species of the Graphipterus serrator group. Therefore these two taxa must be ranked as one species. Nonetheless both taxa differ clearly from each other and are well established in the literature as subspecies (Basilewsky 1977; Lorenz 2005; Huber and Marggi 2017). Therefore we prefer a conservative taxonomic approach which avoids taxonomic inflation (cf. Zachos et al. 2013, Assmann et al. 2008) and preserve the rank of subspecies for both taxa.

Holotype: ♂ (White label with brown margin, brown typewritten): <EX Musaeo/Chaudoir>. (Red label, black typewritten): <TYPE>. Deposited in BMNH, Chaudoir collection [examined].

Small-sized taxon with 28–30 mostly elongated white spots, usually with several spots fused with lateral margin, and with series of usually ten elongated spots, regularly at least several are fused to each other along median suture. Two marginal extensions elongated from humeri posteriorly. Median lobe of aedeagus with wide and flat tip.

Graphipterus minutus goryi resembles G. minutus minutus, for further details see Graphipterus minutus minutus.

BL male: 11.2–11.8 mm, average 11.5 ± .02 mm; BL female: 11.4–13.6 mm, average 12.2 ± 0.9 mm.

Head wide; HW/PW: 0.78; EYL: 1.1–1.3 mm; EYL/EL: 0.19. Frontal ridge absent. Male with two short parallel frontal stripes of decumbent white scales usually diverging apically became wispy, not reaching the level of supraorbital setae. Mentum with two pronounced teeth (cf. Fig. 3e).

Pronotum strongly cordiform; PL/PW: 0.57; BPW/PW: 0.5; posteriomedially flat and without white margin; white lateral margin as twice as antennomere II long.

Elytra almost rounded, humeri strongly rounded, lateral margin continuously rounded; EL: 5.9–7.3 mm, average 6.2 mm; EW: 5.2–6.8 mm, average 5.7 mm; EL/EW: 1.1. Lateral cross section convex. Scales black, disc not visible between them (cf. Fig. 6a). White lateral margin nearly as wide as antennomere I long and with six, rarely fewer, elongated extensions; extensions I elongated from humeri posteriorly; extensions II and III in front and behind the middle of lateral margin, usually much longer than lateral margin wide. White posterior margin becomes narrower toward the tip, usually disappearing in front of it; gap at suture as wide as lateral margin. Disc with 28–30 mostly elongated spots, several spots fused with lateral margin resulting in extensions II and III, a series of 10, (rarely 12–14), elongated spots fused to each other parallel to suture, anterior spot as wide as extension I. stridulatory structure absent. Apical sinuation almost lacking, apex almost absent, not rounded (Fig. 7d). Suture inconspicuous.

Legs short; MTIL: 2.5–3.3 mm, average 3.0 mm; El/MTIL: 1.9. Metatibial secondary spur brown. MTAL: 2.1–3.8 mm, average 2.9 mm; MTAL/MTIL: 0.87. Claws of hind legs brown at base.

Median lobe of aedeagus with wide and flat tip (Fig. 9g).

Unknown.

No co-occurring species.

Saudi Arabia and Iraq (Fig. 19). There are old records from Iran (Perse), but without indication of exact locality.

The species does not seem to be endangered as it has a wide distribution range in desert regions that are not strongly affected by human activities.

Lectotype: ♂ (blue label, black handwritten): <Graphipterus/multiguttatus. Ol./Egypt. G. Olivieir> (Green circular label with black margin, black typewritten): <COLLECTION/OLIVIER/TYPE>. Deposited in NHMB, Olivier collection [examined]. Syntypes: NHMB (Olivier collection): Egypte Olivier, multiguttatus, (uc), TYPE (♂); (Olivier collection) Collection Olivier, TYPE (♂); (General collection) Egypte Olivier, multiguttatus, Egypt, Oliv., Bedel et (uc), p. 339, 1909, vid. (♀).

Small species with 16–20 white, mostly elongated spots on elytra, only posterior discal spots rounded; 4–6 marginal extensions, extension I oriented slightly posteriorly. Median lobe of aedeagus with ventrally short, unbent tip.

Graphipterus multiguttatus resembles G. rotundatus from which it differs mainly by the following characters: G. multiguttatus: average body length of 13.2 mm; El/MTIL, 1.6; all elytral spots with similar size; MTAL/MTIL, 0.84; median lobe of aedeagus with ventrally short unbent tip. G. rotundatus: average body length of 17.4 mm; El/MTIL, 2.08; posterior elytral spots larger than all other spots; MTAL/MTIL, 1.28; median lobe of aedeagus with longer (than G. multiguttatus) slightly bent tip. Graphipterus multiguttatus resembles also G. sharonae sp. n., from which it differs mainly by body length, elytral pattern, and shape of median lobe of aedeagus (see full comparisons under G. sharonae sp. n.).

BL male: 10.0–15.0 mm, average 13.0 ± 1.3 mm; BL female: 11.5–16.0 mm, average 14.0 ± 1.2 mm.

Head wide; HW/PW: 0.76; EYL: 1.0–1.6 mm; EYL/EL: 0.17. Frontal ridge slightly developed. In male, apical white frons stripes wider than exposed frons (Fig. 4a); stripes elongate, reaching the level of supraorbital setae (populations east of the Dead Sea-Rift Valley), or being shorter (populations west of the Dead Sea-Rift Valley). Mentum with 2–3 teeth.

Pronotum cordiform; PL/PW: 0.66; BPW/PW: 0.64; posteromedially concave and without white margin; white lateral margin as wide antonomer 1 long.

Elytra oval, humeri rounded; EL: 4.5–9.1 mm, average 7.7 mm; EW: 4.1–8.0 mm, average 6.4 mm; EL/EW: 1.2. Lateral cross section quite flat. Elytra with Dense black scales, disc not visible between them (cf. Fig. 6a). White lateral margin nearly as wide as half of antennomere I long and with 6, sometimes four extensions; extension I medium long, shorter than anterior spot, but longer than extension II and shorter than extension III, which is wider than lateral margin; extension II sometimes constricted, rarely absent or fused with lateral disc spot. White posterior margin as wide as lateral margin, gap at suture smaller than lateral margin. Disc usually with 16 sometimes 18 rounded to elongate spots; anterior spot slightly elongate, longer than extension I; lateral spots rounded, adjacent, or sometimes fused to extension II, six spots forming an arch pattern anteriorly and laterally to posterior rounded larger spots. Apical sinuation slightly developed to straight, apex not protuberant, broadly rounded, especially on the medial side (Fig. 7c). Suture conspicuous.

Legs long; MTIL: 3.7–5.5 mm, average 4.7 mm; El/MTIL: 1.6. Metatibial secondary spur brown at base, MTAL: 3–4.5 mm, average 3.7 mm; MTAL/MTIL: 0.8. Claws of hind legs black at base.

Median lobe of aedeagus with short, unbent tip (Fig. 9h).

In the western Negev (Israel), the species shows a significant habitat preference for stabilized interdunes and for the semi-stabilized slopes. In this region it is completely absent from the crest of shifting sand dunes. On the dunes it prefers the lower part of the north-facing slope, which is the part of the dune being most humid and most vegetated by annual plants (Fig. 12). Large populations inhabit the loamy and more humid region in the northern Negev. In spring, after an extremely dry winter, specimens might also be found on the margins of irrigated agriculture fields.

Figure 12. 

Habitat of Graphipterus multiguttatus: Sand rich with loess soil, relatively rich in annual plants. Western Negev sands, Israel.

Graphipterus multiguttatus lives in sympatry with G. serrator in Egypt and Israel.

Egypt, Israel, and Jordan (Fig. 19). The only Jordanian population of which we are aware lives between Aqaba to Ma’an, and inhabits a flat sand dune area without or only slightly developed crust. In the same habitat Anthia (Thermophilum) duodecimguttata (Bonelli, 1813) and Amara maindromi Bedel, 1907 occur.

The species does not seem to be endangered as it has a wide distribution range and it prefers habitats that are not strongly affected by human activities. However, in Israel, in the Dead Sea region and the Arava Valley, G. multiguttatus has been collected in the past, but no longer exists there. The latest records from these regions are Ein Gedi, 1976; Ein Husub, 1956 (leg. unknown, both specimens preserved in KCE); Sedom road, 1953; Ein-Radian, 1958 (leg. Ch. Lewinsohn, both specimens preserved in SMNHTAU). Habitats for G. multiguttatus on the Israeli side of the Arava valley may have disappeared. Anthia (Thermophilum) duodecimguttata (Bonelli, 1813), one of the co-occurring ground beetle species of the Jordanian population of G. multiguttatus, was last found in 2003 in Israel (coll. U. Shanas, V. Chikatunov, SMNHTAU; pers. obs.).

Specimens from Jordan and the central Negev in Israel are usually larger than those from the western Negev. Specimens from the HaBesor National Park are smaller than those from the western Negev. The latter populations of G. multiguttatus which co-occur with G. serrator populations (Renan et al. 2011) have individuals with intermediate body lengths.

Graphipterus kindermanni has to be ranked as a junior synonym of G. multiguttatus. We checked for comparison the types of Basilewsky in MRAC (but did not find the type in NHMB that Basilewsky noted he had checked there) and did not find any morphological differences, with the exception of white setae on the elytral base. Both Basilewsky (1977) and Lorenz (2005) contended that G. kindermanni is a synonym of G. luctuosus.

Seasonality and daily activity time are in the same as in G. serrator (see there), but the species seems to spend more time under shrubs. Graphipterus multiguttatus prefers stabilized and semi-stabilized sand with high vegetation. The population densities in the sympatric areas of the distribution ranges are lower than those for G. serrator. The beetles dig burrows between the hard crust layer and the soft sand, sometimes close to the dwarf-shrubs. Frequently, the openings do not collapse or become covered by sand. The beetles sometimes close the openings with sand from inside. Diet, intraspecific behavior including copulation and the chirping sounds produced by the stridulatory structure, are same as in G. serrator.

In comparison to the co-occurring G. serrator, the scraping spectrograms of G. multiguttatus show clear differences in pulse interval as well as in the sound pressure level (Fig. 10b).

Holotype: ♂ (Blue label, black handwritten): <Pletieri. Chevrolat./Oran. D.S Fargeau>. (White label, black typewritten): <P. Bedel/Visit 1905>. (White label with brown margin, brown typewritten): <EX Musaeo/Chaudoir>. (Red label, black typewritten): <TYPE>. Deposited in NHMB, Chaudoir collection [examined].

Predominantly dark, medium-sized species with 18-24 small, mostly rounded white spots on elytra, four usually short marginal extensions. Median lobe of aedeagus with ventral, short, unbent tip.

Graphipterus peletieri resembles G. luctuosus (see comparisons in G. luctuosus).

BL male: 13.9–14.8 mm, average 14.3 ± 0.4 mm; BL female: 11.5–16.1 mm, average 13.6 ± 1.8 mm.

Head medium; HW/PW: 0.76; EYL: 1–1.5 mm; EYL/EL: 0.17. Mentum with usually three teeth (Fig. 3d). Frontal ridge absent. In male, apical white frons stripes slenderer than exposed frons (cf. Fig. 4a).

Pronotum strongly cordiform, PL/PW: 0.63; BPW/PW: 0.6; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra oval, humeri rounded; EL: 7.4–9.1 mm, average 8; EW: 5.7–7.9 mm, average 7.0 mm; EL/EW: 1.15. Elytra with brown scales, disc of elytra visible between scales (cf. Fig. 6b). White lateral margin wide as half antennomere I long and with four extensions; extensions often constricted; extension I elongated, shorter than extension II; sutural gap of white posterior margin wider than lateral margin. Disc usually with 18–24 mostly rounded spots; anterior pair of spots slightly elongate, as wide as extension I, lateral spots rounded, adjacent or sometimes fused to extension II, posterior pair of spots rounded, slightly larger than others, located toward suture. Lateral cross section quite flat. Apical sinuation developed, apex slightly protruded, strongly rounded (cf. Fig. 7b). Suture inconspicuous.

Legs long; MTIL: 3.8–5.3 mm, average 4.8 mm; El/MTIL: 1.7. Metatibial secondary spur brown. MTAL: 2.8–4.0 mm, average 3.5 mm; MTAL/MTIL: 0.7 (all other species of the G. serrator group El/MTIL: 0.8). Claws of hind legs brown at base.

Median lobe of aedeagus with ventrally short, not bent tip (Fig. 9i).

Unknown.

Graphipterus peletieri lives in sympatry with five other species in north-west Algeria: G. luctuosus, G. rotundatus, G. valdanii, G. stagonopsis sp. n., and G. piniamitaii sp. n.

North-west Algeria and north Morocco (Fig. 17).

The species does not seem to be threatened as it has a wide distribution range that appears to be mostly not strongly affected by human activities.

Alluaud (1926) initially erroneously named this species Graphopterus lepeletieri and since then this spelling has commonly been used by many authors (e.g., Basilewsky 1977). Graphipterus luctuosus was ranked as a subspecies of G. peletieri by Basilewsky (1977), but as a “good” species by Lorenz (2005) and Huber and Marggi (2017).

Holotype: ♂ (White label, black handwritten): <Kebili>. (White label, black typewritten): <Ex Museo/L. Vibert>. (ae). Deposited in NHMB, general collection.

Paratypes. (20♂, 4♀). El Hammama, Tunis: (Gabès), I. 1889, Alluaud (♂) (ZMUC); Gafsa Tunis, Vibert Lyon (♂) (NMP). Kebilli, Tunis: 1906, EX Call. Maindron M., Call G. Babault 1930 (♂ae) (NHMB, General collection); 1950, Cobos Sa’nchez, (uc) (♂) (NHMB, Negre collection); L. Vibert, Ex Musaeo (♂ae) (MRAC); Call. Mus Congo, Col. P. Basilewsky (5♂) (RMRAC); Tunisia, Kebili 15 km N.W, 17.III. 1986, Zool. Mus. Copenhagen Exp. (3♂) (ZMUC); Kebili 2 km s, W. Ziegler, 30 m, Dünen, 5.3.2012, (♂, 2♀) (DWC, CAB); Douz, south Tunisia, Zaafrane (Sahara), 02.04.1992 (♂). S. Tunisia (Kebili), Zaafrane, 12 km SW Douz, 21.IV.2007 M. Liebscher (♀); S. Tunisia (Kebili), Zaafrane, 12 km SW Douz, 21.IV.2007 M. Liebscher (♀); C. Tunisia, 2 km E. Kairouan, 23.4.2005, M. Liebscher Sammlung (♂,♀) (DWC). Oasis Gafsa: Tunis, B v. Bodemeyer (♂) (DEI Muncheberg Call- 01314); B v. Bodemeyer, O. Leonhard, (uc) (♂ae) (DEI Muncheberg Call- 01315); B v. Bodemeyer, O. Leonhard (♂) (DEI Muncheberg Call- 01316).

Medium-sized species with usually 24 white large rounded and elongated spots on elytra; posterior discal spots slightly larger than other spots; six marginal extensions (Fig. 25a). Median lobe of aedeagus with slightly bent tip.

Graphipterus piniamitaii sp. n. is easily distinguished from all other species of the group by its large white spots on the elytra. The new species resembles G. mauretensis sp. n. (see comparisons in G. mauretensis sp. n.).

BL male: 15.5–19.8 mm, average 17.5 ± 2.1 mm; BL female: 17–17.9 mm, average 17.5 ± 0.3 mm.

Head medium; HW/PW: 0.76; EYL: 1.5–1.8 mm; EYL/EL: 0.17. Mentum with two or three teeth. Frontal ridge absent. In male, apical white frons stripes wider than exposed frons (cf. Fig. 4b).

Pronotum cordiform; PL/PW: 0.63; BPW/PW: 0.66; posteromedially concave and without white margin; white lateral margin as wide antennomere I long.

Elytra oval, humeri rounded, but slightly protruding; EL: 8.2–11.0 mm, average 9.4 mm; EW: 6.5–9.2 mm, average 8.1 mm; EL/EW: 1.2. Lateral cross section quite flat. Suture conspicuous. Black scales dense, disc not visible between scales (cf. Fig. 6a). White lateral margin nearly as wide as 1½ antennomere I long and with six extensions; extension I triangular, slightly elongated and posteriori oriented; extensions II and III frequently constricted at base, usually wider than lateral margin. White posterior margin as wide as lateral margin or wider, not becoming narrower towards the suture; gap at suture smaller than lateral margin. Disc usually with 24 rounded to elongate, moderate large spots; anterior spot elongated, as wide as extension I, posterior discal spots slightly larger than other spots, series of six elongated spots along suture, sometimes fused to each other; posterior discal spots larger than other spots. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (cf. Fig. 7a).

Legs long; MTIL: 5.3–6.6 mm, average 6.1 mm; El/MTIL: 1.54. Metatibial secondary spur brown at base, MTAL: 4–5.7.0 mm, average 4.9 mm; MTAL/MTIL: 0.8. Claws of hind legs brown at base.

Median lobe of aedeagus with slightly bent tip (Fig. 9j).

The species is dedicated to Pinchas (Pini) Amitai, an inspiring entomologist and mentor who wrote the first Hebrew photographed insect guide.

The species dwells in the vicinity of Kebili on intensively grazed dunes, together with Anthia (Thermophilum) sexmaculata (Fabricius, 1787) and A. venator (Fabricius, 1792) (Fig. 13). The dunes have a diverse vegetation of shrubs and dwarf-shrubs.

Figure 13. 

Habitat of Graphipterus piniamitaii sp. n.: Shifting sand dunes with vegetated Nebka hills, Kebili, Tunisia.

Graphipterus piniamitaii lives in sympatry in Kebili and Gabès region in Tunisia with G. luctuosus, G. peletieri, and G. heydeni.

Restricted to Central Tunisia, from the vicinity of Kebili to Gabès (Fig. 18).

The species does not seem to be endangered, as the preferred habitat is not strongly affected by human activities.

Holotype: ♂ (White label, brown handwritten): <Reymondi m./ (same label, black typewritten): Antoine det.>. (White label, black handwritten): <Inhamid/ Sahara septe./(Reymond)>. (Red label, black handwritten): <HOLOTYPE>. Deposited in NHMB, General collection, box 31[examined].

Large species with 20-24 isolated white round spots on elytra, six marginal extensions, extension II short, almost triangular. Humeri very narrowed, maximum width of elytra at interior rear third. The discal elytra pattern comprises a group of 8–12 elongated spots in an order parallel to the suture. Median lobe of aedeagus with ventrally bent tip.

Graphipterus reymondi resembles G. sharonae sp. n., from which it differs mainly by mentum and humeri morphology, pattern, color and morphology of elytra (see full comparisons under G. sharonae sp. n.). Graphipterus reymondi resembles also G. stagonopsis sp. n., from which it differs mainly by mentum morphology, pattern, and morphology of elytra, and color of claws and spurs (see full comparisons under G. stagonopsis sp. n.).

BL male: 17–18, average 17.6 ± 0.4 mm; BL female: 17.4–21.4, average 19.3 ±2 mm.

Head medium; HW/PW: 0.76; EYL: 1.6–1.8 mm; EYL/EL: 0.17. Mentum with three teeth (cf. Fig. 3d). Frontal ridge absent. In male, apical white frons stripes wider than exposed frons (Fig. 4b).

Pronotum wide; PL/PW: 0.72; BPW/PW: 0.63; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra with strongly narrowed humeri; EL: 9.4–10.3 mm, average 9.75 mm; EW: 8.0–8.5, average 8.3 mm; EL/EW: 1.2. Lateral cross section convex. Suture conspicuous. Scales brown, disc visible between them (Fig. 6b). White lateral margin nearly as wide as antennomere I long and with six extensions; extension I triangular with rounded tip, slightly more elongated than in G. serrator, wider and shorter than extensionII; the latter one elongated, at third quarter of elytra. White posterior margin commonly slightly wider than lateral margin, gap at suture smaller than lateral margin, usually with a small, indistinct tip anteriorly. Disc usually with 20–24 (rarely 18), mainly rounded spots; anterior pair of spots rounded, wide as extension I, usually smaller than posterior spots, but larger than spots on central disc; central disc spots usually asymmetrically smeared; posterior pair of spots rounded; one or two small additional spots adjacent laterally to the posterior ones. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a).

Legs long; MTIL: 5.8–6.1 mm, average 5.9 mm; El/MTIL: 1.6. Metatibial secondary spur brown. MTAL length: 4.2–5, average 4.7 mm; MTAL/MTIL: 0.8. Claws of hind legs black at base.

Median lobe of aedeagus with ventrally bent tip (Fig. 9k).

Unknown.

No co-occurring species.

Morocco (Fig. 16).

Unknown.

Lectotype: ♀ (blue label, black handwritten): <rotundatus/Klug*/x.118-21./ Bir Hamam El Eherenberg> (White label, black handwritten): <Zwischen Bir-Lebuck/and Bir Hamam/(Libye)> (White label, black typewritten): <Type> (White label, black typewritten): <Hist. –Coll. (Coleoptera)/Nr. 1299/ Graphipterus rotundatus/ Klug*/Bir Hamam El Eherenberg/Zool. Mus. Berlin>. Deposited in ZMHB [examined]. Paralectotype: ♂ (Red label, black typewritten): <Type> (White label, black handwritten): <1299> (white label, black typewritten): <Hist.–Coll. (Coleoptera)/Nr. 1299/Graphipterus rotundatus/Klug*/Bir Hamam El Eherenberg/Zool. Mus. Berlin>. Deposited in ZMHB [examined].

Small species with large distribution range, high variation in size (15–19 mm) and variation in elytra pattern (4–6 extensions, 16–22 spots). posterior discal spots larger than other spots; six spots usually forming an arc pattern anterior and lateral to posterior spots; Median lobe of aedeagus with short, slightly bent tip.

Graphipterus rotundatus resembles G. multiguttatus (see comparisons in G. multiguttatus) and G. luctuosus (see comparisons in G. luctuosus).

BL male: 15.0–19.0 mm, average 17.4 ± 1.5 mm; BL female: 15.4–17.1 mm, average 16.1 ± 1.3 mm.

Head slender; HW /PW: 0.72; EYL: 1.4–1.7 mm; EYL/EL: 0.16. Mentum with 2–3 teeth. Frontal ridge slightly developed. In male, apical white frons stripes wider than exposed frons (Fig. 4b).

Pronotum cordiform; PL/PW: 0.65; BPW/PW: 0.69; posteromedially concave and without white margin; white lateral margin as wide antonomer 1 long.

Elytra oval, humeri rounded; EL: 8.9–11.0 mm, average 9.7 mm; EW: 7.0–8.7 mm, average 7.8 mm; EL/EW: 1.25. Lateral cross section quite flat. Dense black scales, disc not visible between scales (Fig. 6a). White lateral margin nearly as wide as half antennomere I long and with six, sometimes fouor extensions; extension I triangular to slightly elongated; extension II absent or only weakly developed, rarely fused with lateral disc spot. White posterior margin becomes narrower towards the suture, gap at suture smaller than lateral margin. Disc usually with 18, sometimes 16 or 22 rounded to weakly elongate spots; anterior spot slightly elongated, wide as extension I, six spots usually forming an arc pattern anterior and lateral to posterior rounded, larger spots. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a). Suture conspicuous.

Legs long; MTIL: 4.3–5.2 mm, average 4.7 mm; El/MTIL: 1.63. Metatibial secondary spur dark at base, MTAL: 5.4–6.9 mm, average 6 mm; MTAL/MTIL: 0.8. Claws of hind legs brown at base.

Median lobe of aedeagus with slightly bent tip (Fig. 9l).

Unknown.

Graphippterus rotundatus lives in sympatry with G. luctuosus, G. peletieri, and G. valdanii in Algeria and Tunisia.

Algeria, Tunisia, and the coastal region of west Libya (Fig. 18).

The species does not seem to be endangered as it has a wide distribution range which is not strongly affected by human activities.

On the label of the Graphipterus rotundatus type, “Libye” is written; however, as far as it is known, C.G. Ehrenberg never succeeded in reaching Libya (Baker, 1997). There is only a very small chance that any other entomologist had collected Graphipterus in Lybia earlier than 1830.

The three larval stages develop during the summer inside ant nests. The first larval instar is nearly 4 mm long and creeps into nests of large ant species, digs there a chamber, preys on the ant’s brood and pupates within the nest. When the first larval instar tries to enter nests of small ants, it is attacked by the ants (Paarmann 1985; Dinter et al. 2002). The larval instars have a mandibular suctorial tube to suck hemolymph from their prey (Brandmayr 1994a, 1994b). Four specimens from the species studied by Wilfried Paarmann, Pietro Brandmayr, and their co-workers were examined; the material belongs to G. rotundatus and not to G. serrator as noted in their publications.

Holotype: ♀ (White label with blue margin, black handwritten): <Graphipterus Latr./serrator Forsk./Aegypten>. Deposited in ZMUC [examined].

Holotype: gender unknown (only fragments of a beetle preserved). (White label with black margin, black handwritten): < variegatus/ 824>. Deposited in ZMK [examined] (Fig. 28d).

Large species with 10–12 isolated white round spots on elytra: anterior and posterior discal spots larger than other spots, six smaller spots near suture form circular pattern on disc; four white marginal extensions present, extension I triangular. Median lobe of aedeagus with ventrally bent tip.

Graphipterus serrator resembles G. valdanii from which it differs mainly by the following characters: G. serrator: mentum with three teeth, mid tooth shallow; PL/PW (0.72); BPW/HW (0.8); EL/EW rounded (1.18); elytra lateral margin wide as antennomere I long; Claws of hind legs dark. G. valdanii: mentum with three teeth, merges shallow and mid tooth bolt; PL/PW (0.64); BPW/HW (1); EL/EW elongated (1.3); elytra lateral margin wide as half antennomere I long; Claws of hind legs brown.

BL male: 17–18 mm, average 17.6 ± 0.4 mm; BL female: 17.4–21.4 mm, average 19.3 ± 2 mm.

Head medium; HW/PW: 0.76; EYL: 1.6–1.8.0 mm; EYL/EL: 0.16. Mentum with three teeth, mid tooth shallow (Fig. 3f). Frontal ridge absent. In male, Apical white frons stripes slenderer than exposed frons (Fig. 4b). Pronotum wide; PL/PW: 0.58; BPW/PW: 0.65; posteromedially concave and without white margins; white lateral margin as wide as antennomere I long.

Elytra oval, humeri rounded; EL: 9.3–11.3 mm, average 10.3 mm; EW: 7.0–9.8 mm, average 8.4 mm; EL/EW: 1.2. Lateral cross section convex. Elytra with dense black scales, disc of elytra not visible between scales (Fig. 6a). White lateral margin nearly as wide as antennomere I long and with four extensions; extension I triangular with rounded angels, margin of elytra wider and shorter than extension II; the latter one elongated; at third quarter of elytra, imaginary line connecting the medial ends of the extension I and I parallel to the suture. White posterior margin forming gap at suture which is wider than lateral margin. Disc usually with 10, sometimes 12 round spots; anterior pair of spots circular to slightly elongate, narrower than extension I, larger than the six central spots forming a circular pattern; anterior and posterior pair of spots circular rounded, larger than other spots; small additional spots frequently present laterally to the posterior spots. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a). Suture inconspicuous.

Legs long; MTIL: 4.8–7.4 mm, average 6.1 mm; El/MTIL: 1.7. Metatibial secondary spur dark. MTAL: 4.0–5.3 mm, average 4.6 mm; MTAL/MTIL: 0.8. Claws of hind legs black at base. Median lobe of aedeagus with ventrally bent tip (Fig. 9m).

Very common in arid sandy habitats, it shows a significant habitat preference for the crest of shifting sand dunes (Fig. 14). It avoids stabilized interdunes and half-stabilized dune slopes (Renan et al. 2011). The sandy habitat in the western Negev sand dunes is poor in perennial woody plants with maximal coverage of 10–15% (Perry 2008; Siegal et al. 2013). The dominant perennial plants are Retama raetam (Fabaceae) and Stipagrostis scoparia (Poaceae).

Figure 14. 

Habitat of Graphipterus serrator: Shifting sand dunes in the Western Negev Sands, Israel.

Graphipterus serrator lives in sympatry with G. multiguttatus in Egypt and Israel.

North-east Egypt (incl. Sinai) and Israel. In Israel it is restricted to the western Negev sand dunes (Fig. 19).

The sand dunes in the western Negev suffer from two major threats: agricultural development that has caused a significant loss of the sands’ range (Ben David and Avni 2013), and a stabilizing process of the shifting sand resulting from a bio-crust (Kidron and Abeliovich 2009). In the Sinai Peninsula, a lack of shrubs as a result of overgrazing threats the population.

The female holotype of Carabus serrator has been considered lost (Basilewsky 1977), but it was recently found by us in ZMUC (Fig. 28b–c). After studying the type material, we agree with Hůrka (2003), Lorenz (2005) and Huber and Marggi (2017), that variegatus (Fabricius 1792) falls within the morphological variability of serrator. Therefore variegatus is confirmed as a junior synonym of serrator (Fig. 28d). Graphipterus serrator lobatus and G. serrator sexmaculatus were considered by Alfieri (1976) as variations of G. serrator. Following the ICZN (1999, Article 45.6.3), a taxon that is described as a variation after 1960 is not valid. Moreover, no holotype has been designated. Therefore, both lobatus Alfieri and sexmaculatus Alfieri are not available names. One specimen from the western Negev sands was found with intermediate characters of G. serrator serrator and G. multiguttatus, and this specimen seems to be a hybrid between them: ♂ Israel, Holot Agur, May 2012, leg. I. Renan.

Adults emerge immediately after the first significant rainfall and inhabit sandy dunes or sand and loess plains and edges of salt lakes. In the spring following an average rainy winter, the species can densely populate the dunes (one observer can locate up to 40 individuals within one hour). Their diet is based mainly on ants and occasionally on other small insects, as well as on dead insects and dead reptiles. Activity is limited by temperature: it begins at a soil temperature of approximetly 18 °C, and ceases at a soil temperature of approximetly 39 °C. By moving between sun-exposed microhabitats and the shadow of dwarf-shrubs can prolong the activity period. Strong wind halts activity due to the beetle's sensitivity to dehydration. Some activity also occurs in the afternoon, but it is significantly lower than in the morning peak hours.

Prior to commencing inactivity, the beetle digs a short burrow with a narrow elliptic cross-section into the dune’s slope. The digging is performed mainly with the hind legs and secondarily with the middle legs. The well-developed, spoon-shaped metatibial spurs (see fig. 4a in Assmann et al. 2015) seem to function as a shovel. The burrow’s opening usually collapses behind the beetle or is covered by shifting sand. In the burrow, a few centimeters below the sand surface, the beetle is relatively protected from predation and can probably still detect the outside temperature and light conditions. In enclosure experiments with individual markings and variation in population density, one of us found that even during the peak activity season, most of the specimens spend most of the days without displaying epigeic activity. An encounter between two individuals of any gender immediately develops into a short, hasty, bite battle and the escape of the loser. In some regions, shade is a limited resource and the battle occurs mainly under bushes and dwarf-shrubs. An encounter between male and female starts with an aggressive fight. The persistent male will then mount the back of the female. His forelegs grasp the female between the basal part of the pronotum and the elytral humeri, while the female tries to grab the male with her hind legs. The copulation lasts for approximately 30 minutes and occurs mostly beneath perennial vegetation. During the fight, the beetles stridulate. This sound is produced when the beetles are threatened by other individuals or by potential predators (Renan unpublished data, based on field observations and arena experiments).

Comparing G. serrator’s scraping spectrograms with those from its co-occurring species, G. multiguttatus, reveals clear differences in pulse intervals as well as in the sound pressure level (Fig. 10).

Holotype, ♂ (White labe, black typewritten): <51780 ISRAEL/ Karmiya N.P/ 7.4.2011/ I. Renan>. (red label): <Holotype>(ae). Deposited in SMNHTAU [examined].

Paratypes: (79♂, 70♀): All material collected in Israel. Ashdod: 6.V.2015, I. Renan (7♂, 14♀); 5.XII.2014, I. Renan (6♂, 3♀); 16.III.2011, (♂) (CAB); 3.IV.1998, H. Ackerman (♂) (SMNHTAU); 16.III.2011 leg. Th. Assmann, (♂,♀), W. Starke leg. (♂,♀) (CAB). Ashkelon [Ashqelon]: 7.IV.2017, I. Renan (6♂, 4♀) (SMNHTAU). Avshalom: 24.III.2012, M. Bologna (2♂) (AVTC). Ayalon: 1.IV.1943 (♂) (KCE). Bat Yam: 14.III.1940, Bytinski- Salz (3♂); 24.III.1940, Bytinski- Salz (2♂); 23.IV.1959, J. Wahrman (2♂, 2♀) (SMNHTAU). Bene’ Berack [Bene Beraq]: 26.II.1954 (♂) (SMNHTAU). ‘En Sarid: 22.IV.2015, I. Renan (2♀) (SMNHTAU). Holon: 14. IV.1981, A. Freidberg (♂) (SMNHTAU). Jaffa [Yafo]: 21.I.1900 (♀) (BMNH). Jaffa-Rehoboth [Rehovot]: 14.VII.1913, S.G.J. Aharoni (♂) (RMRAC). Karmiyya N.P: 07.IV.2011, I. Renan (4♂, 2♀) (SMNHTAU). MiqWeYisra’el: 14. IV.1934, F.S. Bodenheimer (♂, 3♀); 11.IV.1946, J. Wahrman (♀); 20. IV.1934, F.S. Bodenheimer (♂) (SMNHTAU). Nachalat Jischack, Palestina [Tel Aviv, Nahalat Yizhaq], 5.VI.1942, Housk (4♂) (NMP). Netanya: III.-IV. 1996, R. Rod (♀) (DWC); 15.II.1955, S. Nothiltz (♂); 11. IV.1957, J. Machlis (♂, ♀); 03.V.1997, R. Hoffman (♂, ♀); 04. IV.2010, I. Renan (5♂, 1ae, ♀) (SMNHTAU); III.IV.1996, leg. R. Rod (♀) (CWD); III.2016, leg. Th. Assmann (3♂, 4♀) (CAB). Nizzanim N.P: 29. IV.2015, Renan I. (14♂, 10♀); 19.V.2009, I. Renan (3♂,2ae, ♀); 15.V.2009, I. Renan (5♂, 2ae, 4♀,) 7.4.2011, I. Renan (♂ae) (SMNHTAU). 22.III.2012, M. Bologna (♂) (AVTC). 25.II.2009, L. Friedman (♀) (BMNH); 07.VI.2007, leg. J. Buse (♀) (CAB). Palmahim: 25.III.1978, Tedeschi (3♂, 1♀) (AVTC). Porat: 22.I.2015, I. Renan (♂); 09.IV.2014, I. Renan (5♂); 19.IV.2015, I. Renan (2♂); 22.IV.2015, I. Renan (8♂, 8♀) (SMNHTAU). Ra’ananna: 11.IV.1947, Bytinski-Salz (♂) (SMNHTAU). 13.VI.1940, Bytinski-Salz (♀) (SMNHTAU). Rafha [Rafiah]: (♂) (ae) (BMNH). Rishon Leziyyon: 17.III.2003, M. Yogev (♂) (BMNH); 10.III.1942, Bytinski-Salz (♀); 29.VI.1979, D. Furth (♂); 1.III.1938 (♂) (SMNHTAU). Tel Aviv: 11.I.1900 (♂) (BMNH); 2.I.1900 (♂) (KCE). Ziqim N.P: 4.VI.2015, I. Renan (2♂, 4♀); 5.V.2015, I. Renan (♂, ♀); 7.IV.2011, I. Renan (2♂, 1♀) (SMNHTAU).

Medium-sized species with 12–18 white elytral spots; the anterior and central ones usually elongated, the posterior ones rounded; six marginal extensions, extension II triangular. Median lobe of aedeagus with bent tip.

Graphipterus sharonae sp. n. resembles G. multiguttatus, from which it differs mainly by the following characters: G. sharonae average body length 13mm; extension slightly elongated 1; median lobe of aedeagus short, unbent tip. G. multiguttatus average body length 17.05 mm; extension I triangular; median lobe of aedeagus with bent tip. Graphipterus sharonae resembles also G. reymondi, from which it differs mainly by the following characters: G. sharonae sp. n. mentum with two teeth, humeri rounded, 12–18 spots on elytra, widest line of elytra located at middle, elytra disc not seen, and elytra scales black while G. reymondi has the mentum with three teeth, humeri narrowed, 20–24 spots on elytra, widest line of elytra located at interior rear third, elytra disc seen, and elytra scales brown.

BL male: 15.0–18.0 mm, average 16.5 ± 0.8 mm; BL female: 16.0–193 mm., average 17.6 ± 0.8 mm;

Head medium; HW/PW: 0.74 mm; EYL: 1.2–1.7 mm; EYL/EL: 0.15. Mentum with two teeth and shallow depression between (cf. Fig. 3b). Frontal ridge slightly developed. In male, apical white frons stripes wider than exposed frons (Fig. 4b).

Pronotum cordiform; PL/PW: 0.66; BPW/PW: 0.66; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra oval, humeri rounded; EL: 8.1–10.3 mm, average 9.2 mm; EW: 6.2–8.8 mm, average 7.8 mm, (EL/EW: 1.3). Lateral cross section flat. Dense black scales, disc not visible between scales (Fig. 6a). White lateral margin as wide as half antennomere I long and with six extensions; extension I triangular with rounded angels, as wide as or wider than elytra margin, wider and shorter than extension III; extension II smaller and usually shorter than two other ones; extension III often constricted at base. White posterior margin almost continuously rounded, only slightly becoming narrower, gap at suture smaller than lateral margin. Disc with 14, sometimes 12 or 18 spots; most anterior pair of spots slightly elongate to rounded, usually wide as extension I, second anterior pair of spots strongly elongate, nearly two times as long as wide; the two lateral pairs of spots rounded, adjacent or sometimes fused to extension II; the tow to four posterior pairs of spots rounded; the medial, most posterior pair of spots larger than all other spots; the outer most posterior pair of spots much smaller than the latter one. Apical sinuation slightly developed to straight, apex not protuberant, broadly rounded, especially on the medial side (Fig. 7c). Suture conspicuous.

Legs long; MTIL: 4.9–6.1 mm, average 5.6 mm; El/MTIL: 1.7. Metatibial secondary spur brown. MTAL: 4.0–5.0 mm, average: 4.5 mm; MTAL/MTIL: 0.8. Claws of hind legs black at base.

Median lobe of aedeagus with bent tip (Fig. 9n).

The species is dedicated to Sharon Renan, biologist, conservationist, and the first author’s wife.

In sand dunes and on calcareous sandstone habitats along the coast. Low, mostly vegetated and stabilized sand dunes are the preferred habitat (Ramot 2008). Individuals are active as far as 50 meters from the shoreline, but seem to be more common further inland. The average annual rainfall in the coastal plain is approxemetly 450 mm (I.M.S, 2016). The dominant perennials of the habitats in Israel are Artemisia monosperma and Helianthemum stipulatum (Fig. 15).

Figure 15. 

Habitat of Graphipterus sharonae sp. n.: Stabilized sand dunes with rich vegetation. Nizzanim, Israel.

No sympatrically occurring species.

Endemic to the Mediterranean coastal plains from north-east Sinai (El Arish) to central Israel south of Nahal Alexander (Fig. 19).

The coastal plain sand dunes of Israel form the largest part of the entire distribution range of G. sharonae sp n. As a result of land use changes and urbanization, less than 25% of the Israel sandy habitats remain and a further decline can be expected. In addition, the remaining dune habitats are under extreme anthropogenic disturbance and highly fragmented (Achiron-Frumkin et al. 2003). The following records are examples of sites that were populated by G. sharonae sp. n. in the past, but where their populations no longer exist: Kefar Bilu, Rehovot, Nes Ziyyona, Bat Yam, Holon, Tel Aviv, Ramat Gan, Bene Beraq, Ra’ananna, Yafo (based on SMNHTAU collection and the authors’ experience).

Despite having no precise data, the habitats in the Gaza Strip and north-eastern Egypt seem also to have declined as these areas feature a strong increase in human population density. In a faunistic survey of the ground beetles of the Sinai Peninsula, Abdel-Dayem et al. (2004) did not record Graphipterus from El-Arish, where it had been present nearly a century ago (records in London, cf. Schatzmayr 1936). El Surtasi et al. (2012) demonstrated the negative effect of urbanization on G. serrator population in Egypt. Both the restricted distribution range of the endemic species G. sharonae sp. n. and the decline in coastal sandy habitats threaten the long-term survival of the species.

Seasonal and daily activity time, as well as diet, intraspecific behavior, including copulation and the chirping sounds produced by the stridulatory structure, are as in G. serrator. Graphipterus sharonae sp. n. prefers stabilized sands with high vegetation cover, and its population density is higher than that of G. serrator.

Holotype.♂ (White label, black handwritten): <Beni Abbes/23.III.48 F. Pierre>. (red label): <Holotype> (ae). Deposited in NHMB [examined].

Paratypes.(11♂, 3♀), NHMB (Colas collection): Gardhaia (Ghardaia), Sahara, G. Mahoux, 19.5.60 (2♂); Beni Abbes, 23.3.48, F. Paiu (2♂, 1- ae). (Negre collection): Beni Abbes, Sahara argelino, J. Mateu (3♀). (Antoine collection): Beni Abes, south Algerien (reymondi) (♂ae). ZMUC Algerie, Beni Abbes, 11.3.1984, Tilg. 4-12.1948, Tentens-Nielsen [G. serratorvaldani Guer. P. Basilewsky 1985] ♂. NMP: Algeria, Igli, 12.IV. 1988, Igt. Kepler, 11/1988. Ex call. M. Dvorak, National Museum, Prague, Czech Republlic. MRAC: Aoulel el Arab Tidicelt Sahara Cen., J. Mateu (♂); Pozo zug (R.O.) Sa’hara espanol, J. Mateu (2♂); Oasis de la-Salah Tidikelt Sahara Cen, J. Mateu (3♂).

Large species with 16 white rounded to elongated spots on elytra, anterior and posterior pair of spots larger than others; six marginal extensions, extension I triangular, extension I and II elongated. Elytra widest at interior rear third, drop-like shape. Median lobe of aedeagus with slightly bent tip.

Graphipterus stagonopsis sp. n. resembles G. reymondi from which it differs mainly by the following characters: G. stagonopsis sp. n.: mentum with two teeth; eight spots on elytra; scales of elytral disc brown; claws of hind legs and metatibial secondary spur dark . G. reymondi: mentum with three teeth; 10–12 spots on elytra; scales of elytral disc black; claws of hind legs and metatibial secondary spur brown .

BL male: 17.2–20.1 mm, average 18.8 ± 1 mm; BL female: 18.4–19.8 mm, average 18.9 ± 0.6 mm.

Head slender; HW/PW: 0.7; EYL 1.1–1.7 mm; EYL/EL: 0.16. Mentum with two teeth (Fig. 3b). Frontal ridge slightly developed. Male, apical white frons stripes wider than exposed frons (Fig. 4b).

Pronotum strongly cordiform; PL/PW: 0.66. BPW/PW: 0.64; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra droplet-like, humeri strongly narrowed; EL: 9.1–11.1 mm, average 8.4 mm; EW: 7.5–9.0 mm, average 8.4 mm; EL/EW: 1.1–1.5. Lateral cross section convex. Dense black scales, disc not visible between them (Fig. 6a). White lateral margin nearly as half wide as antennomere I long and with six extensions; extension I triangular with rounded angels, as wide as lateral margin, posteriorly oriented; extension II small, often constricted at base, as wide as lateral margin; extension III large, elongated, posteriorly oriented. White posterior margin becomes narrower towards suture; gap at suture wider than lateral margin. Disc usually with 16 spots; anterior pair of spots elongate, as wide as extension I; posterior pair of spots rounded and larger than other ones; six spots forming arch pattern anterior and lateral to posterior rounded larger spots. Apical sinuation slightly developed to straight, apex not protuberant, broadly rounded, especially on the median side (Fig. 7c). Suture inconspicuous.

Legs long; MTIL: 6.0–7.0 mm, average 6.5 mm; El/MTIL: 1.6. Metatibial secondary spur black. MTAL: 4.7–5.3 mm, average 4.9 mm; MTAL/MTIL: 0.8. Claws of hind legs black at base.

Median lobe of aedeagus with bent tip (Fig. 9o).

The name is derived from ancient Greek (σταγών, óψις) and means ”drop-like” which refers to the shape of the elytra.

Unknown.

Graphipterus stagonopsis lives in sympatry with G. luctuosus, G. peletieri, and G. valdanii in Ghardaia, Algeria.

Central and west Algeria (Fig. 18).

Unknown.

Neotype. ♂ (White label, black handwritten): < Bou saada/ Oherthur R.>. (ae). Deposited in NHMB, General collection. (Red label, black typewritten): < Neotypus Graphipterus valdanii Guérin-Méneville, 1859/ des. I. Renan, 2018>.

Neoparatypes. NHMB (General collection): Baniou, Vibert L. (♂,♀); Bou saada, 1875, Oberthur R. (2♂); Bou saada, Oberthur R. (♂); Bou saada, Dr Martin (♂). (Negre collection): Algeria (♂); Bou Saada (♂); Bou Saada, Dr Martin (♀); BMNH: Bou-Saada, 1875, Oberthur R. (2♂). MRAC: Bou Saada, P. Basilewsky (♂); Bou saada, Dr Martin (♂). DEI: Bou Saada, O. Leonhard / Dr Martin (♂); ZMUC: Bou Saada, 28.4.1927 (♀); (uc) (♂);

Large species with 10–16 white round spots on elytra; anterior and posterior discal spots larger than other spots; four white marginal extensions, oval elytra, extension I triangular. Median lobe of aedeagus with bent tip.

Graphipterus valdanii resembles G. serrator (see comparisons in G. serrator) and G. heydeni (see comparisons in G. heydeni).

BL male: 14.8–19.0 mm, average 17.1 ± 1.7 mm; BL female: 18.6–20.5 mm, average 18.6 ± 1.9 mm.

Head slender; HW/PW: 0.71; EYL: 1.4–1.8 mm; EYL/EL: 0.15. Mentum with mentum with two teeth as margin between them slightly convex in middle (Fig. 3c). Frontal ridge slightly developed. In male, apical white frons stripes slenderer than exposed frons (Fig. 4a).

Pronotum cordiform; PL/PW: 0.64; BPW/PW: 0.7; posteromedially concave and without white margin; white lateral margin as wide as antennomere I long.

Elytra oval, relatively elongated, humeri rounded; EL: 8.1–12 mm, average 10.6 mm; EW: 6.5–9.1 mm, average 8.0 mm; EL/EW: 1.3. Lateral cross section convex. Dense black scales, disc not seen between scales (Fig. 6a). White lateral margin as wide as half antennomere I long and with four extensions; extension I triangular with rounded angels, much wider than lateral margin, but shorter than extensionII; the latter one elongated, positioned at second third of elytra. Apical sinuation strongly sinuated, apex strongly protruded, forming almost a rectangular. White posterior margin evenly rounded, not becoming narrower towards the suture; gap at suture wider than lateral margin. Disc with (10–) 14 (–16) spots; anterior pair of spots rounded to slightly elongate, much smaller than extension I; anterior and posterior pair of spots round, larger than other spots; small additional spots located medially to extension II and laterally to posterior spots. Apical sinuation strongly developed, apex protruded, almost rectangular, only slightly rounded at most distant tip (Fig. 7a). Suture inconspicuous.

Legs long; MTIL: 4.9-7.0 mm, average 6.3 mm; El/MTIL: 1.7. Metatibial secondary spur dark. MTAL: 5.4-6.9 mm, average 6.0 mm; MTAL/MTIL: 0.72. Claws of hind legs brown at base.

Median lobe of aedeagus with bent tip (Fig. 9p).

Unknown.

Graphipterus valdanii lives in sympatry with G. peletieri, G. luctuosus, G. rotundatus, and G. stagonopsis in Algeria.

The arid and semi-arid regions of north-east Algeria from Ghardaia, to Bou-Saada and Tebessa (Fig. 17).

Figure 16. 

Distributional records of G. luctuosus, G. mauretensis sp. n., and G. reymondi.

Figure 17. 

Distributional records of G. barthelemyi, G. peletieri, and G. valdanii.

Figure 18. 

Distributional records of G. heydeni, G. piniamitaii sp. n., G. rotundatus, and G. stagonopsis, sp. n.

Figure 19. 

Distributional records of G. minutus minutus, G. minutus goryi, G. magnus sp. n., G. multiguttatus, G. serrator, and G. sharonae sp. n.

Unknown.

Guérin-Méneville described G. valdanii as a new species in 1859. Chaudoir (1870: 296), having seen the types that had been collected in Algeria and compared them with G. serrator from Egypt, contending that they differed in elytral shape and were a local variation of G. serrator.

The type of valdanii is lost. Our attempts to find any specimen from the typical series in several museums (incl. NHMB) was unsuccessful, as Guèrin's collection was sold and his material appears to be unavailable (Thierry Deuve pers. comm.). Chaudoir (1870) used G. valdanii, but Guérin-Méneville introduced the species as G. valdani (sic). Since then, Chaudoir’s spelling has appeared in most of the literature that deals with this species.

Figure 20. 

Dorsal habitus of Graphipterus: a G. barthelemyi with greyish scales phase b G. barthelemyi without grayish scales.

Figure 21. 

Dorsal habitus of Graphipterus: a G. heydeni b G. luctuosus c G. heydeni lectotypes' lables (ZSM).

Figure 22. 

Dorsal habitus of Graphipterus: a G. magnus sp. n. b G. mauretensis sp. n.

Figure 23. 

Dorsal habitus of Graphipterus: a G. minutus minutus b G. minutus goryi.

Figure 24. 

Dorsal habitus of Graphipterus a G. multiguttatus b. G. peletieri.

Figure 25. 

Dorsal habitus of Graphipterus: a G. piniamitaii sp. n. b G. reymondi.

Figure 26. 

Dorsal habitus of Graphipterus: a G. rotundatus b G. serrator.

Figure 27. 

Dorsal habitus of Graphipterus: a G. sharonae sp. n. b G. stagonopsis sp. n.

Figure 28. 

Dorsal habitus of Graphipterus: a G. valdanii b G. serrator holotype (ZMUC) c G. serrator holotypes’ label d Graphipterus variegatus holotype (ZMUC).