Research Article |
Corresponding author: Mario García-París ( chalcabris@yahoo.es ) Academic editor: Christopher Majka
© 2019 José L. Ruiz, Alexandre François, Mario García-París.
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:
Ruiz JL, François A, García-París M (2019) A new singular species of Croscherichia Pardo Alcaide, 1950 (Coleoptera, Meloidae, Mylabrini) from arid zones of eastern Morocco. ZooKeys 885: 27-50. https://doi.org/10.3897/zookeys.885.34308
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A new species of blister beetle (Coleoptera, Meloidae, Mylabrini), Croscherichia armass Ruiz, François & García-París, sp. nov., is described from the arid steppes of eastern Morocco (Missour, Boulemane Province). The new species presents traits shared with both Croscherichia and desert species of the genus Ammabris, making it morphologically singular. Conspicuous external similarities (coloration pattern, shape of the mandibles, setation) between C. armass sp. nov. and Ammabris allow the two to be easily confused. However, C. armass sp. nov. can be readily distinguished from all other Croscherichia species by the following traits: reddish-orange legs with dark tarsi; relatively short black antennae with the proximal-most three to four antennomeres of each antenna having a reddish-brown coloration; dense and silvery body setation that lies over most of the body integument; straight and pointed outer mandible margins that protrude from the labrum; a mesosternum with an angulate anterior margin; a short, subcylindrical, and weakly spatulate external metatibial spur that is truncated obliquely at the apex. Croscherichia armass sp. nov. is only known from three localities in the arid Hammada steppes, which are located within the Quaternary alluvial plains of the Muluya river valley. Live specimens of C. armass sp. nov. were found in flight and actively feeding on Atriplex halimus (Chenopodiaceae) flowers at the end of summer (mid-September). The phenology of C. armass sp. nov. is exceptional as no other Mylabrini species known from eastern areas of Morocco are active in late summer.
Ammabris, arid steppes, Atriplex, biodiversity, blister beetle, Hammada, morphology, North Africa, taxonomy
Mylabrini Laporte 1840, which is distributed over most of the Old World (Palaearctic, Oriental and Afrotropical regions), is the most diversified tribe within the family Meloidae. Mylabrini is comprised of about 760 species distributed in 12 genera (
In contrast to Hycleus and Mylabris, Croscherichia Pardo Alcaide, 1950 is a medium-sized genus currently comprised of 18 described species, whose internal phylogeny based on morphological traits and a largely undisputed taxonomy is well recognized (
In its current sense, Croscherichia [type species: Mylabris circumflexa Chevrolat, 1840 (= Mylabris paykulli Billberg, 1813), by monotypy] is monophyletic. This genus, with its sister group Mimesthes Marseul, 1872, is characterized by the synapomorphic condition of its external metatibial spur, which is spatulate in both groups (
In this work, a new species of Croscherichia is described from arid zones of central-eastern Morocco (Missour region) (Figs
Habitat and live specimens of Croscherichia armass sp. nov. A thickets of saltbush (Atriplex halimus) in an alluvial silt plain of the Al Baten area between Missour and Outat el Haj B detail of Atriplex halimus L. where the species is usually found. Emirates Center for Wildlife Propagation (
The fauna of Mylabrini is relatively well known in Morocco (e.g.,
A total of 19 dry-preserved (5 males and 14 females) and 4 ethanol-preserved (2 males and 2 females) specimens were used to describe the new species of Croscherichia. The specimens were collected from three nearby localities (separated by a maximum distance of 35 km) in the region of Missour (Fès-Boulemane, Morocco) (Fig.
Interspecific comparisons were performed using the material indicated below (308 specimens), including specimens of all known species of Croscherichia except C. femorata (Klug, 1845) from Arabia, C. quadrizonata (Fairmaire, 1875) from eastern Algeria, Tunisia, and Libya, and the aforementioned problematic C. sonyae. The diagnostic morphological traits of these three species were extracted from
Material examined: Croscherichia albilaena (Bedel, 1899): ALGERIA: Biskra, L. Bleuse, Mai 1885 (Slg. R. Oberthür, Coll. E. Martin, Eing. Nr. 4 1956, Croscherichia albilaena Bedel, Dr Kaszab det. 1957): 1 ex. (HNHM). Croscherichia bedeli (Bleuse, 1899): ALGERIA: Aïn-Sefra, Mai 1896, A. Chobaut (Coll. Reitter, Mylabris bedeli Bleuse, det. Dr Kaszab, Croscherichia bedeli Bleuse, Dr Kaszab det. 1957): 4 exx. (HNHM). MOROCCO: Figuig, 29-V-1991, G. Chavanon leg.: 2 exx. (JLR); Figuig, 21-V-1993, G. Chavanon leg.: 3 exx. (JLR); Figuig, erg à Aristida pungens, 10-V-1997, G. Chavanon leg.: 15 exx. (JLR); Figuig, 14-V-1999, G. Chavanon leg.: 9 exx. (JLR). Croscherichia delarouzei (Reiche, 1865): SYRIA: Syrien, Kafa, Reitter (Coll. Reitter, Croscherichia delarouzei Reich., Dr Kaszab det. 1957): 1 ex. (HNHM). Croscherichia fulgurita (Reiche, 1866): ALGERIA: Aïn-Sefra (Oran), L. Bleuse (Coll. Reitter, Mylabris fulgurita Reiche, det. Dr Kaszab, Croscherichia fulgurita Reiche, Dr Kaszab det. 1957): 1 ex. (HNHM). MOROCCO: Figuig, 14-V-1999, G. Chavanon leg.: 1 ex. (JLR). Croscherichia gilvipes (Chevrolat, 1840): EGYPT: Le Caire Hénon (Slg. R. Oberthür, Coll. E. Martin, Eing. Nr. 4 1956, Croscherichia angulata Klug, Dr Kaszab det. 1959): 1 ex. (HNHM). MOROCCO: Bouârfa, 20-V-1993, G. Chavanon leg.: 3 exx. (JLR); Bouârfa, 23-V-1993, G. Chavanon leg.: 5 exx. (JLR); Figuig, erg à Aristida pungens, 10-V-1997, G. Chavanon leg.: 1 ex. (JLR); Route Bouàrfa-Figuig, 32.47915/-1.74761, 1170 m, 24-V-2015, A. François, M. García-París & J.L. Ruiz leg.: 2 exx. (
The morphological study was carried out on dry-mounted specimens using stereomicroscopy. Male specimens were rehydrated prior to the extraction of their genital structures, which were subsequently mounted on cardboard with dimethylhydantoin formaldehyde resin and pinned adjacent to their respective specimen. Measurements were taken using a micrometre coupled to one of the eyepieces, and camera lucida drawings were made of the structures. The ethanol-preserved paratypes were not measured to prevent possible tissue deterioration. Total specimen length was measured along the dorsal side from the anterior end of the labrum (with the head extended) to the elytral apex. Maximum width was measured as the width between the outer edges of the elytra at approximately three-fourths of the elytral length, also in dorsal vision. Photographs of live and recently dry-mounted specimens were taken with a digital camera. The terminology suggested by
In the present study, the evolutionary species concept (as modified by
For practical purposes and to separate groups of morphologically similar species of Croscherichia, the taxonomic scheme in the species key of
Holotype
: 1 male (dry-preserved) (Fig.
Total length
: 9.1 mm. Maximum width: 2.85 mm. General appearance elongated, stylized (Fig.
Head
(Fig.
Antennae
(Fig.
Pronotum
(Fig.
Scutellum hemi-elliptic, rounded along its posterior margin, with black and shiny integument, very densely and finely punctured, with a setation that completely covers the surface, similar to that of the pronotum, giving it a silvery appearance.
Elytra
(Fig.
Mesopleurae
without a careniform fold or rim along the anterior margin; setation dense and directed backwards, covering most of the mesopleural surface except along the central zones contiguous to the mesosternum, which are smooth and almost hairless. Mesosternum (Fig.
Ventral region of the body with black integument, shiny, finely microreticulated, with fine and very dense punctures, subconfluent, hidden under the setation; setation very dense, evenly distributed and longer than that of the pronotum, giving the body a silver appearance except along the narrow mid-longitudinal band of the metasternum, which is smooth and hairless. Last abdominal ventrite with a deep and wide V-shaped central notch in its posterior margin, with a setation much less dense than the rest of the abdominal ventrites.
Legs
thin and narrow, with reddish-orange femora and tibiae, somewhat obscured distal ends, brownish-red trochanters that are slightly orange at the ends, black coxae that turn brownish red towards the apex; tarsi, dark brown almost black, except for the first protarsomere and the basal half of the first meso- and metatarsomeres, which are reddish brown; relatively short protarsi (length excluding claws = 1.39 mm), with protarsomere I short and wide, subconical in dorsal view (length, l = 0.30 mm; maximum width, w = 0.22 mm; l/w = 1.36), protarsomeres II (l = 0.22 mm; w = 0.16 mm; l/w = 1.37), III (l = 0.20 mm; w = 0.16 mm; l/w = 1.25) and IV (l = 0.18 mm; w = 0.12 mm; l/w = 1.5) subequal to protarsomere I, although gradually becoming smaller; protarsomere V (l = 0.5 mm; w = 0.14 mm; l/w = 3.57) subcylindrical, narrow and elongated; mesotarsi similar to but slightly more elongated (l = 1.58 mm) than protarsi and with slightly narrower tarsomeres; metatarsi longer and narrower than mesotarsi (l = 2.02 mm), with narrower tarsomeres. Leg punctures very dense and thin, slightly smaller than those on the ventral region of the body; setation whitish yellow, dense and thinner and shorter than that of the ventral region, almost lying on the surface, with the highest density on the tibiae and the inferior side of the femora; internal side of the anterior tibiae with a band of whitish hairs shorter and slightly denser than on the rest of the protibiae, hairs on the distal half of the outer carina slightly more erect and denser than the rest, uniform in size, without longer hairs. External apical end of the protibiae terminates in a small or narrow digitiform expansion almost covered entirely by setation. Tarsi with tarsomeres I to V bearing a small hirsute brush on the underside; on tarsomere V, the brush only occupies the proximal half; hairs on the protarsomeres are exclusively white, while those on the meso- and metatarsomeres are a mixture of white and dark brown. Apical spines of pro- and mesotibiae very small and narrow, subequal, with a blunt tip; apical internal spine of the metatibiae similar to that of the pro- and mesotibiae (length = 0.21 mm), the external one slightly longer and thicker (length = 0.22 mm), subcylindrical, obliquely truncated at the apex, weakly spatulate but slightly widened distally (Fig.
Aedeagus
(Fig.
Aedeagus of Croscherichia armass sp. nov. A tegmen, lateral view B tegmen, dorsal view C median lobe, lateral view D spiculum gastrale. Note the narrow and elongate parameres and the middle lobe with two evident teeth in the ventral region that are subequal and clearly separated (type isoharpagae sensu
Female: Similar to the male but differing in the following features: protibiae with a small sharp tooth at the external apical end, with long and fine whitish semi-erect hairs along the outer edge, standing out from the short and lying setation of the surface; external side of the first four protarsomeres with long erect hairs, directed forward, similar to those on the outer edge of the protibiae, which also stand out from the short lying setation; last abdominal ventrite with a posterior margin that is not notched in the middle. Valvifer and stylus as in Figure
Not very marked but present in the following characters: total length, 7.3 to 10.1 mm, mean 8.6 mm (N = 19), males 7.6 to 9.1 mm, mean 8.2 mm (N = 5), females 7.3 to 10.1 mm, mean 8.7 mm (N = 14); maximum width, 2.2 to 3.7 mm, mean 2.8 mm (N = 19), males 2.2 to 2.8 mm, mean 2.6 mm (N = 5); females 2.3 to 3.7 mm, mean 2.9 mm (N = 14); frontal red spot with little variation in diameter and intensity; chromatic pattern of the antennae variable, particularly in the amplitude of the brown coloration: between antennomeres I and V, both inclusive: 26.3% (N = 19) (antennomere I presents a very dark basal region), between antennomeres II and V: 52.6% (sometimes the apical third of antennomere V is black), between II and IV: 5.3%, and between II and III: 15.8%, including the holotype; antennomeres VI–XI always black, although some specimens present a narrow basal brown ring on antennomeres VI and XI; punctures and setation of the head slightly variable in density, especially in the posterior area of the frons and in the vertex, with larger individuals showing greater density; smooth and glabrous pronotal areas that vary to a slight extent; elytral design generally constant, although there is a certain variation in the size of the spots of the anterior and posterior series (in the latter, the two rounded spots are joined in some specimens) and in the thickness of the sinuous median band, which, in some specimens, is interrupted in the middle; the external apical spine of metatibiae in larger specimens is visibly widened in the distal half. No variation is observed in the aedeagus of the studied males.
The coloration of live specimens varies markedly with that of the preserved specimens (Fig.
The word “armass” refers to the Tamazight (Bereber) name of the plant on which C. armass is usually found (Atriplex halimus).
Croscherichia armass is only known from three nearby localities (separated by a maximum distance of about 35 km) in the region of Missour (Boulemane Province, Fès-Boulemane Administrative Region), east of the Eastern Middle Atlas in the valley or middle section of the Muluya River (Oued Moulouya) (Fig.
The region is dominated by calcareous sedimentary materials: Quaternary alluvial plains with superficial soils. The area is within the mesomediterranean bioclimatic zone (
Live C. armass specimens can be found in flight and actively feeding on flowers of the Chenopodiaceae A. halimus roughly between the hours of 10:00 and 16:00 (UTC). The coloration of C. armass blends in with the inflorescences of A. halimus, which can make locating specimens difficult.
Croscherichia armass has a very particular combination of morphological characters that distinguishes it from all other species of the genus. It has a relatively small size (mean = 8.3 mm, range: 7.3–10.1 mm); a black head, pronotum, and ventral region; reddish-orange legs with darkened tarsi; black antennae except for the proximal-most three to four antennomeres, which have a dark reddish-brown colour; and a dense and silvery body setation lying over most of the body integument. The broad head has a red spot on the frons and a dense and relatively thick setation on the integument. The outer margins of the mandibles are straight, pointed, and protrude from the labrum (at approximately one third of its length). The antennae are relatively short and black except for antennomeres II or III to V (rarely including antennomere I and the base of VI), which are dark or reddish brown. The pronotum is as broad as it is long and is neither elongated or narrowed at the anterior margin. It is characterized by a weak depression in its anterior third, a marked central longitudinal depression, and two well-defined smooth and glabrous areas lateral to the mid-line. The pronotal setation is silvery, thick, and dense against the integument, masking it for most part. The mesosternum has an angulate anterior margin and is covered with setation lying along its lateral branches. The external metatibial spur is short, subcylindrical, and not spatulate; it is also slightly widened at its distal end and truncated obliquely at the apex. The leg claws have upper and lower lobes of equal length that are markedly curved. The last abdominal sternum of the male has a deep cleft in its posterior margin. The elytral design of C. armass is unique within Croscherichia: tricolored, with a small prehumeral black spot, a narrow black zigzagging central band, and two rows of black rounded spots. The anterior row is comprised of three spots and the posterior one of two spots that neither touch the suture nor the external margin of the elytron.
The general appearance of C. armass resembles that of some North African species of Ammabris, particularly Ammabris boghariensis (Raffray, 1873) and, to a lesser extent, A. avilai (Ruiz & García-París, 2008). The three species are similar in the following features: elytral coloration, mandible morphology (straight and pointed outer margin), the presence of a red spot on the frons, pronotal sculpture (a central longitudinal groove with smooth areas lateral to it), and the overall silvery setation on the integument (see
According to the identification key of
Croscherichia armass can be further grouped with species having claws with upper and lower lobes of similar length, which includes C. paykulli, from the Maghreb (from Morocco to Libya); C. delarouzei, from Palestine, Israel, Jordan, and Lebanon; two subspecies of C. sanguinolenta, C. s. sanguinolenta, widely distributed throughout arid and semi-arid zones of North Africa, Senegal, and the Near East to Iran, and C. s. arabica Bologna & Coco, 1991, only known from Saudi Arabia; and C. gilvipes, distributed throughout arid regions of North Africa and the Near East (Israel, Jordan, and Syria) (
Croscherichia sonyae, assigned with reservations to Croscherichia (
Prior to the present study, nine Croscherichia species (i.e., C. bedeli, C. fulgurita, C. gilvipes, C. litigiosa, C. mozabita, C. paykulli, C. sanguinolenta, C. tigrinipennis, and C. wartmanni) were known for Morocco. None of these species are endemic to the country and, with the exception of C. paykulli (widely distributed throughout Morocco), all are restricted to arid or semi-arid zones in the south or the east (
With the recent discoveries of new blister beetle species endemic to Morocco (
To Grégoire Liénart (