Studies on the subgenus Agraphoderus Bates of Blennidus Motschulsky from Peru : the jelskii species – group ( Coleoptera , Carabidae , Pterostichini )

Studies on the subgenus Agraphoderus Bates of Blennidus Motschulsky from Peru: the jelskii species–group (Coleoptera, Carabidae, Pterostichini).— Four new species of Blennidus subgenus Agraphoderus are described from the Andes of Southern Peru: B. (A.) procerus n. sp., B. (A.) abramalagae n. sp., B. (A.) etontii n. sp. and B. (A.) straneoi n. sp. Together with B. (A.) jelskii (Tschitschérine, 1897), they form a very homogeneous group of probably closely related species (the jelskii group), which is distinguished from other members of the subgenus by the distinctive morphology of the aedeagus. A redescription of B. (A.) jelskii is given based on the lectotype and paralectotype designated by Straneo & Vereshagina (1991), supplementing Tschitschérine’s brief original description. The distribution pattern of the species presently included in the jelskii species–group is discussed, emphasizing distinctive traits of stenoendemic species inhabiting restricted geographical areas, and discussing their possible origin by allopatric speciation.


Introduction
The genus Blennidus Motschulsky, 1866 at present includes about 120 species (Lorenz, 2005a) distributed in the Andean region of South America, extending from North Colombia to Chile.Moret (1995) considered the genera/subgenera Agraphoderus Bates, 1891, Ogmopleura Tschitschérine, 1899, Sierrobius Straneo, 1951, Pachyabaris Straneo, 1953and Pseudocynthidia Straneo, 1953 (formerly separated based on the presence/absence of metathoracic wings as well as of a transverse sulcus on the abdominal sterna IV-VI) as synonyms of the senior name Blennidus.He regarded these characters were largely inconsistent, in particular the presence/absence of an abdominal transverse sulcus.This view-point was accepted by Lorenz (2005a).
In a later paper, Moret (2005) used three 'convenience subgenera' (Blennidus s. str., Sierrobius Straneo, 1951 andAgraphoderus Bates, 1891) and described a new subgenus (Jasinskiellus Moret, 2005).His aim was to keep the lineages of some well characterized species separate, even though these taxa lacked any phyletic value (Moret, in litteris 2011).Among these four subgenera, the subgenus Agraphoderus is the richest in species.It includes the micropterous species which usually display a transverse sulcus and/or puncture rows only laterally on abdominal sterna IV-VI, with a gap in the middle.They were formerly attributed to Ogmopleura Tschitschérine, 1899.We agree with Moret (1995) on the inconsistency of such characters, as many species in Agraphoderus display only very superficial and hardly visible lateral impressions or very tiny punctures.Moreover, high variability can be observed among species which are probably related based on the similarities of their external morphology and, above all, their genitalia.For these reasons, assessment of the phyletic relationships among species within the genus is urgently required.In the meantime, we prefer to retain these 'convenience taxa', as the species concerned in this article can all be included in the subgenus Agraphoderus sensu Moret (2005).
A total of 34 Agraphoderus species are currently recorded from Peru (Straneo, 1993;Allegro, 2010).They display a rather uniform 'harpaloid' habitus (Straneo, 1993) and share common and peculiar features such as the enlarged basal half of the aedeagus, just above the insertion of the parameres.Most species have a stumpy, convex body with short appendages; only a few species have a slender and/or depressed body and long appendages (Allegro, 2010).Their habitat and altitudinal distribution are also very similar; they mainly inhabit high altitude Andean grasslands at 3,300-4,800 m a.s.l.
During two stays on the mountains of the Cordillera Blanca, Peru, one in November-December 2005 and the other in June-July 2008, one of the authors (G.Allegro) had the opportunity to collect abundant material of Carabidae.Data concerning some of this material have already been published (Allegro et al., 2008;Allegro, 2010).He explored an area included in the Dept. of Ancash (Provinces of Huaráz, Asunción, Yungay and Fitzcarrald) where he collected abundant material of the Blennidus subgenus Agraphoderus.The other author (P.M. Giachino) is in possession of Pterostichine material collected in Peru by M. Etonti.This material includes several specimens that belong to the same genus and subgenus.A few specimens collected by M. Etonti were also deposited in the Mateu Collection at Museo Regionale di Scienze Naturali, Torino, Italia.The study of this material and its comparison with the type material of the Straneo Collection (at Museo Civico di Storia Naturale, Milano, Italia) and the type material described by Tschitschérine (Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia) allowed us to recognise groups of species that can be distinguished by their very homogeneous male genitalia morphology.The study also allowed us to recognise many undescribed species.These groups, which we will consider one at a time in separate articles, seem to be congruent from a geographical point of view, as they include species ranging in clearly delimited areas of the country.The same approach was adopted by Moret (1995), who grouped the Ecuadorian Blennidus into eight different species-groups.However, some Peruvian species cannot yet be placed in any defined species-group and only further surveys in the Andean countries will provide material and information for reliable phylogenetic analysis.
This work deals with the species belonging, in our opinion, to the jelskii species-group of the genus Blennidus, all of them localized in the Andes of Southern Peru.In addition to Blennidus (Agraphoderus) jelskii (Tschitschérine, 1897), this group includes four new species which are described, illustrated and discussed here.B. (A.) jelskii is redescribed, as the original description (Tschitschérine, 1897) is incomplete concerning some characters.The following acronyms were used for the type material: HT.Holotype; PT, PTT.Paratype(s); LT.Lectotype; PLT.Paralectotype.

We examined material of the genus
Locality labels of the material examined are quoted in their original form.
Drawings of the genitalia were made using a camera lucida connected to a Leica MZ 12.5 stereo microscope.The habitus drawings are by G. Allegro.

Taxonomy
B. (Agraphoderus) jelskii species-group Straneo (1993) considered B. (A.) jelskii easy to distinguish on account of the basally sinuate sides of the pronotum as he knew of no other Peruvian species with such a character.Probably for this reason, he labelled 'Ogmopleura jelskii' two specimens from Callanga (ex Staudinger) showing this feature, putting them in his collection together with a paralectotype of B. jelskii from the Solsky Collection (ZIRA).It should be noted that in the same paper Straneo wrongly attributed the specimens of the type series to the locality Lima, when they were undoubtedly from Puno (green label handwritten by Jelski specifying 'Puno Peru-Jelski 1870'), as correctly reported in Straneo & Vereshagina (1991).More detailed examination revealed that the specimens from Callanga belonged to an undescribed species with similar external morphology but a different aedeagus (although of similar structure).Moreover, in the material collected by Etonti, we were able to find three further species that had a uniform structure of male genitalia, all from a restricted area of the Cuzco Region.Callanga, the type locality of Blennidus (Agraphoderus) straneoi n. sp., probably refers not to the town of that name near Lima but to the ancient archaeological site in the Cuzco Dept., from where four of the five species of the jelskii group are recorded.This is confirmed by the presence of two further specimens from Callanga (ex Staudinger) in the type series of Blennidus (Agraphoderus) mesotibialis (Straneo, 1993) (in CSt), a species which is only recorded from the Cuzco area.This evidence led us to consider the five species we attribute to the jelskii species-group as probably strictly related, probably sharing a common ancestor and distributed only, according to present knowledge, in the Southern Andean districts of Peru.
As far as diagnostic characters are concerned, four species show sinuate (or nearly straight) sides at the base of pronotum (B.(A.) jelskii, B. (A.) straneoi n. sp., B. (A.) procerus n. sp. and B. (A.) abramalagae n. sp.).A fifth species, B. (A.) etontii n. sp., in spite of its similarity with the other species of the group on account of structure of male genitalia, displays rounded sides.The elytral striae are equally impressed in all these species.In some cases males possess a faint metallic lustre.Females are distinguished for their markedly dull elytra with strong polygonal microsculpture.The postangular seta of pronotum is always placed at the posterior angle.The mesotibiae of males are not distally swollen or provided with spines; only one species (B.(A.) etontii n. sp.) displays preapically swollen male metatibiae.The abdominal sterna IV-VI are usually smooth, showing at most very tiny punctures at the sides; only B. (A.) jelskii has a short transverse impression interrupted in the middle, together with a few punctures.Sternum VII has a pair of apical setae in males and two pairs in females.The character which defines this group is the distinct and very homogeneous morphology of the aedeagus: it is slender in lateral view, sometimes ventrally depressed, distally very thin and flat (not laterally sinuate), and bent downwards but turning up at apex; it also shows a triangular apical blade, that is more or less acutely pointed in the dorsal view.Moreover, the ostium is always large, covering almost the whole dorsal part, and very long, extending to the basal bulb.Such a long ostium is only known in a few other Peruvian Agraphoderus species, including Blennidus (Agraphoderus) aulacostigma (Tschitschérine, 1897), a taxon with uncertain affinities recorded from Puno.The left paramere is always in discoid in shape, while the right paramere is narrow, curved (almost straight only in B. (A.) jelskii) and apically spatulate.

Type material
In the original description Tschitschérine (1897) described this species based on three specimens (2 ♂♂ and 1 ♀).Curiously, Straneo & Vereshagina (1991) mention four specimens of the type series (3 ♂♂ and 1 ♀), designating a Lectotype (♂) and three Paralectotypes.Finally, Straneo (1993) affirms to have examined three specimens (LT ♂ and 2 PLT with sex not specified) of the type series, quoting a fourth specimen from the original description (?).
Type locality Puno and Lima, Peru (Tschitschérine 1897).The two examined specimens (LT ♂ and PLT ♀) are labelled 'Puno' by Jelski (handwritten green label), and we could not locate any material from Lima.According to Straneo & Vereshagina (1991), only the LT ♂ is labelled 'Puno', whilst three other type specimens are generically labelled 'Peru'; these authors mentioned the presence in CSt of two further specimens from Callanga, not belonging to the type series, which are described in this paper as a new species (B.(A.) straneoi n. sp.).Straneo (1993) subsequently, and rather surprisingly, attributed the LT and two specimens of the type series to the locality 'Lima', quoting a specimen labelled 'Puno' from the original description.In our opinion, assuming the Southern distribution of the species of the jelskii group in the Peruvian Andes and based on the labels by Jelski in the examined material, B.
Head moderately large, eyes convex, temples as long as 1/3 of eyes.Clypeus bisetose; labrum transverse, 6-setose.Frontal impressions short and superficial.Frons between eyes smooth and shiny, with sparse tiny punctures.Terminal labial Identification key for the species of the jelskii group of the genus Blennidus (Agraphoderus): Clave de identificación de las especies del grupo jelskii del género Blennidus (Agraphoderus): 1. Sides of pronotum sinuate at the base or nearly straight; male metatibiae not preapically swollen > 2 Sides of pronotum rounded over entire length; male metatibiae preapically swollen (sometimes scarcely evident) B. (Agraphoderus) etontii n. sp.Pronotum decidedly wider than long (width/ length = 1.33).Microsculpture evident only at sides, disk smooth and shiny, with regular tiny punctures on the whole surface.One basal impression on each side, superficial, linear and impunctate.Mid longitudinal line well impressed between anterior and posterior submarginal sulci, which are scarcely evident.Lateral margins narrowly bordered on overall length, rounded and markedly sinuate in basal 3 rd .Anterior and posterior margins not beaded.Front angles scarcely prominent; hind angles right (fig.7).Two lateral setae on each side, one at hind angles and one at about ¾ from base.Prosternal process glabrous, cuneate and not margined at apex.

Impressed
Elytra oval elongate (length from basal margin to apex/width = ♂ 1.49, ♀ 1.51), pear-shaped and moderately convex on disk.Shoulders obtuse, without denticles.Scutellar stria usually evident between striae 1 and 2. No setigerous punctures near base.Sides rounded and sinuate near apex; lateral border narrow.Usually three setae on each elytra, the 1 st at basal 5 th and in the 3 rd interval or on 3 rd stria, the following adjoining the 2 nd stria.Striae smooth or weakly punctate, superficial but evident to apex, all equally impressed but more impressed in males than in females.Intervals flat (♀) or hardly convex (♂); 2 nd interval wider than 1 st and as wide as 3 rd .
Metepisterna short, slightly longer than wide.Abdominal sterna IV-VI glabrous except for the pair of central setae; a short transverse impression together with sparse, hardly visible punctures are present at each side.
Legs stout.Mesotibiae crenulate at the external edge; male mesotibiae and metatibiae distally not swollen and without inner spines or denticles.Metatrochanters less than half length of femora.5 th tarsomeres with one pair of setae above and three pairs beneath.Male protarsomeres 1-3 triangular and strongly dilated.Metatarsomeres 1-4 externally furrowed.
Aedeagus (fig.15) slender (length 1.95 mm), with enlarged basal bulb and median lobe long, almost cylindrical and in lateral view regularly rounded, distally very thin and bent downwards, but turning up at apex; the apical blade is, in dorsal view, triangular and bluntly pointed (fig.20).Ostium in dorsal position, large and very long, extended to the basal bulb.Right paramere almost straight.

Distribution and habitat
The only known specimens of B. (A.) jelskii are the type series described by Tschitschérine (1997).According to the above considerations this material probably originates from the Puno area.Nothing is known of the environmental conditions of the sites inhabited by this species or its ecology.

Etymology
The specific epithet refers to the slender, oval elongate habitus.(A.) jelskii by the more superficial elytral striae, the elytra not pear-shaped and the median lobe of the aedeagus, which is more angulately inserted on the basal bulb, more slender at middle and ventrally depressed.
Head large, eyes moderately convex, sometimes more or less flattened in both ♂♂ and ♀♀; temples as long as 1/3 of eyes.Clypeus bisetose; labrum transverse, 6-setose.Frontal impressions short and superficial.Frons between eyes smooth and shiny, with sparse tiny punctures.Terminal labial palpomere with thin and sparse hairs; penultimate palpomere bisetose and with a short apical seta.Median tooth of mentum prominent and not excavate at apex.Antennae short, hardly reaching the base of pronotum, with antennomeres 4-10 only a little longer than wide.
Pronotum moderately transverse and very variable in shape (width/length = 1.18-1.33).Microsculpture evident only at sides, nearly absent on disk, with regular and very tiny punctures on the whole surface.In some specimens some lateral transverse wrinkles are evident.One basal impression on each side, superficial, linear and impunctate.Mid longitudinal line superficial, sometimes barely visible; submarginal sulci scarcely evident.Lateral margins narrowly bordered on overall length, weakly sinuate to nearly straight at basal 3 rd (figs 7 and 11-14).Anterior margin unbordered, the posterior bordered only at sides.Base sinuate to nearly straight at sides.Front angles very slightly prominent; hind angles from nearly right to obtuse (figs. 7, 11-14).Two lateral setae on each side, one at hind angles and one at about ¾ from base.Prosternal process glabrous, cuneate and not margined at apex.
Elytra slender, oval elongate (length from basal margin to apex/width = 1.50-1.58),narrow at base and subdepressed on disk (fig.32).Microsculpture markedly impressed in both sexes.Shoulders obtuse, without denticles.Scutellar stria usually evident between striae 1 and 2. No setigerous punctures near base.Sides rounded and scarcely sinuate near apex; lateral border narrow.Usually 3 setae on each elytra, the 1 st often at basal 5 th and in the 3 rd interval or on 3 rd stria (sometimes very close to 2 nd stria), the following adjoining the 2 nd stria.Striae smooth, superficial but evident to apex, all equally impressed, but more so in males than in females.Intervals flat (♀) or slightly convex (♂); 2 nd interval wider than 1 st and as wide as or narrower than 3 rd .
Metepisterna short, a little longer than wide.Abdominal sterna IV-VI glabrous except for the pair of central setae; no transverse impressions nor punctures are evident at sides.Legs stout.Mesotibiae crenulate at the external edge; male mesotibiae and metatibiae distally not swollen and without inner spines or denticles.Metatrochanters shorter than half femora.5 th tarsomeres with one pair of setae above and 2 pairs beneath.Male protarsomeres 1-3 triangular and strongly dilated.Metatarsomeres 1-4 externally not furrowed.
Aedeagus (fig.17) slender (length 1.85 mm), with median lobe sharply inserted on the large basal bulb, in lateral view slightly curved, distally very
Head moderately large, eyes convex in both sexes; temples as long as 1/2.5 of eyes.Clypeus bisetose, a little excavate at middle; labrum transverse, 6-setose.Frontal impressions superficial and barely visible.Frons between eyes smooth and shiny, with sparse tiny punctures.Terminal labial palpomere with thin, sparse hairs; penultimate palpomere bisetose and with a short apical seta.Median tooth of mentum prominent and excavate at apex.Antennae short, hardly reaching the base of pronotum, with antennomeres 4-10 only a little longer than wide.
Pronotum transverse (width from basal margin to apex/length = 1.30-1.35).Microsculpture evident only at sides, disk smooth and shiny.One basal impression on each side, superficial, linear and impunctate.Mid longitudinal line superficial, sometimes hardly visible, only impressed between the submarginal sulci, which are barely visible.Lateral margins narrowly bordered on overall length, sinuate at basal 3 rd .Anterior margin unbordered, the posterior bordered at sides; base markedly sinuate at sides.Front angles very scarcely prominent; hind angles nearly right or obtuse (fig.10).Two lateral setae on each side, one at hind angles and one at about ¾ from base.Prosternal process glabrous, cuneate and widely margined at apex.

mm
Elytra oval (length/width = 1.42-1.43),subdepressed on disk.Microsculpture more impressed in ♀♀.Shoulders obtuse, without denticles.Scutellar stria usually evident between striae 1 and 2. No setigerous punctures near base.Sides rounded and scarcely sinuate near apex; lateral keel moderately large.Usually 3 setigerous punctures on each elytron, the 1 st foveate at basal 5 th and in the 3 rd interval, the second just behind middle and adjoining the 2 nd stria; the third before apex and on the 2 nd stria.Striae smooth, superficial but evident to apex, all equally and weakly impressed in both sexes.Intervals flat in both sexes; 2 nd interval wider than 1 st and as wide as 3 rd .
Metepisterna longer than wide.Abdominal sterna IV-VI glabrous except for the pair of central setae; no transverse impressions or punctures evident at sides.Legs stout.Mesotibiae crenulate at the external edge; male mesotibiae distally not swollen and without inner spines or denticles.Metatrochanters shorter than half of femora.5 th tarsomeres with one pair of setae above and two or three pairs beneath.Male protarsomeres 1-3 triangular and strongly dilated.Metatarsomeres 1-4 not furrowed externally.
Aedeagus (fig.18) slender (length 2.12 mm), with median lobe sharply inserted on the large basal bulb, in lateral view slightly curved, ventrally depressed, distally thin and almost straight, slightly turning up at apex; in dorsal view, the median lobe is wide at base and the apical blade is triangular and bluntly pointed (fig.23).Ostium in dorsal position, large and very long, extended nearly to the basal bulb.Right paramere curved.
Distribution and habitat At present B. (A.) abramalagae n. sp. is known from the type locality, the Abra Malaga pass near Cuzco, and from Nevado Salcantay, not far from Abra Malaga, in Southern Peru.These sites (4,200-4,400 m a.s.l.) are characterized by Andean grassland.

Etymology
We are pleased to dedicate this species to Mirto Etonti, collector of the specimens forming the type series of this species.

Differential diagnosis
Blennidus (Agraphoderus) etontii n. sp. is the only species in the jelskii group, as far as we know, with completely rounded sides of pronotum.Moreover, it is the only species with male metatibiae preapically swollen.It is distinguished from the other Peruvian Agraphoderus that have rounded sides of the pronotum by the morphology of the median lobe of aedeagus, which suggests its relationship with the species of the jelskii group.
Head moderately large, eyes convex in both sexes; temples as long as 1/2.75 of eyes.Clypeus  bisetose, slightly excavate at middle; labrum transverse, 6-setose.Frontal impressions superficial and barely visible.Frons between eyes smooth and shiny, with sparse tiny punctures.Terminal labial palpomere with sparse, thin hairs; penultimate palpomere bisetose and with a short apical seta.Median tooth of mentum prominent and excavate at apex.Antennae short, hardly reaching the base of pronotum, with antennomeres 4-10 only a little longer than wide.
Pronotum transverse (width/length = ♂ 1.30, ♀ 1.34).Microsculpture evident only at sides, disk smooth and shiny.One basal impression on each side, superficial, linear and impunctate.Mid longitudinal line superficial, only impressed between the submarginal sulci, which are barely visible.Lateral margins rounded and narrowly bordered on overall length.Anterior margin unbordered, the posterior bordered at sides; base sinuate at sides.Front angles very scarcely prominent; hind angles obtuse (fig.9).Two lateral setae on each side, one at hind angles and one about ¾ from base.Prosternal process glabrous, cuneate and widely margined at apex.
Elytra oval elongate (length from basal margin to apex/width =1.47-1.52),moderately convex.Microsculpture more impressed in ♀♀.Shoulders obtuse, without denticles.Scutellar stria superficial and scarcely evident between striae 1 and 2. No setigerous punctures near base.Sides rounded and scarcely sinuate near apex; lateral keel narrow.Usually three setae on each elytron (the HT with four setae on left elytron), the 1 st at basal 4 th and in the 3 rd interval or on 3 rd stria, the following adjoining the 2 nd stria.Striae smooth, superficial but evident to apex, all equally and weakly impressed in both sexes.Intervals flat in both sexes; 2 nd interval wider than 1 st and as wide as 3 rd .
Metepisterna longer than wide.Abdominal sterna IV-VI glabrous except for the pair of central setae; only a few tiny punctures are hardly visible at sides.Legs stout.Mesotibiae crenulate at the external edge; male mesotibiae distally not swollen and without inner spines or denticles; male metatibiae preapically moderately swollen.Metatrochanters shorter than half femora.5 th tarsomeres with one pair of setae above and 2 pairs beneath.Male protarsomeres 1-3 triangular and strongly dilated.Metatarsomeres 1-4 externally not furrowed.
Aedeagus (fig.19) slender (length 2.10 mm), with median lobe sharply inserted on the large basal bulb, in lateral view hardly curved, ventrally depressed, distally thin and slightly bent downwards, turning up at apex; in dorsal view, the median lobe is wide at base and the apical blade is triangular and acutely pointed (fig.24).Ostium in dorsal position, large and very long, extended nearly to the basal bulb.Right paramere curved.

Etymology
We dedicate this species to the author of the most important study concerning the Peruvian Agraphoderus, Stefano L. Straneo, renowned specialist of world Pterostichinae.
Head moderately large, eyes convex in both sexes; temples as long as 1/3 of eyes.Clypeus bisetose, moderately concave at middle; labrum transverse, 6-setose.Frontal impressions superficial and directed inward.Frons between eyes smooth and shiny, with sparse very tiny punctures.Terminal labial palpomere with thin and sparse hairs; penultimate palpomere bisetose and with a short apical seta.Median tooth of mentum prominent and excavate at apex.Antennae short, hardly reaching the base of pronotum, with antennomeres 4-10 only a little longer than wide.
Pronotum transverse (width/length = ♂ 1.25, ♀ 1.35).Microsculpture more evident at sides, where some transverse wrinkles are evident; disk smooth.One basal impression on each side, superficial, linear and impunctate.Mid longitudinal line superficial, only impressed between the submarginal sulci, which are scarcely evident.Lateral margins narrowly bordered on overall length, nearly linear (♂) or weakly sinuate (♀) at basal 3 rd .Anterior margin unbordered, the posterior bordered at sides; base sinuate at sides.Front angles scarcely prominent; hind angles obtuse (fig.8).Two lateral setae on each side, one at hind angles and one at about 4/5 from base.Prosternal process glabrous, cuneate and not margined at apex.
Elytra oval (length from basal margin to apex/width = 1.45 in both sexes), moderately convex on disk with marked apical declivity (fig.31).Microsculpture more impressed in ♀.Shoulders obtuse, without denticles.Scutellar stria usually evident between striae 1 and 2. No setigerous punctures near base.Sides rounded and scarcely sinuate near apex; lateral keel moderately large.Usually three setae on each elytra, the 1 st at basal 5 th and in the 3 rd interval or on 3 rd stria, the following adjoining the 2 nd stria.Striae smooth or weakly punctuate (♀), superficial but evident to apex, all equally and slightly impressed in both sexes.Intervals flat in both sexes; 2 nd interval wider than 1 st and wider than 3 rd .Metepisterna longer than wide.Abdominal sterna IV-VI glabrous except for the pair of central setae; no transverse impressions or punctures are evident at sides.Legs stout.Mesotibiae crenulate at the external edge; male mesotibiae distally not swollen and without inner spines or denticles.Metatrochanters shorter than half femora.5 th tarsomeres with one pair of setae above and two pairs beneath.Male protarsomeres 1-3 triangular and strongly dilated.
Aedeagus (fig.16) slender (length 2.0 mm), with median lobe roundly inserted on the large basal bulb, in lateral view hardly curved at middle, thin and ventrally depressed, distally tapering and slightly bent downwards, turning up at apex; in dorsal view, the median lobe is wide at base and the apical blade is triangular and moderately sharply pointed (fig.21).Ostium in dorsal position, large and very long, extended nearly to the basal bulb.Right paramere curved.The apparently separate distribution of the Blennidus (Agraphoderus) species of the jelskii group, each one inhabiting a specific Andean valley or district, suggests that these flightless high-altitude carabids constitute a complex of stenoendemic species living in restricted geographic areas and in narrow altitudinal ranges.They probably originated by allopatric speciation due to the isolating effects of Pleistocene climatic cycles combined with the effects of orographic barriers, in particular when a warmer climate shifted the montane biota upwards during the interglacials, causing its fragmentation and isolation (Simpson-Vuilleumier, 1971).The same differentiation processes occurred to other high-altitude Andean carabid genera such as Trechisibus and Oxytrechus (Allegro et al., 2008;Moret, 2005).According to our preliminary observations carried out on the Blennidus (Agraphoderus) populations inhabiting the Cordillera Blanca, many other species of this genus must be considered as stenoendemics.The data of Straneo (1993) referring to species covering wide geographical ranges need careful reexamination, as they could refer to specimens wrongly attributed to a single species (Allegro & Giachino, unpublished).

Distribution and habitat
Based on these considerations, it is very likely that new surveys in unexplored areas of Andes together with research into other entomological collections will bring to light many more unknown taxa.
(A.) jelskii probably occurs only in the Puno area.Differential diagnosis Among the Peruvian Agraphoderus, only four species display sinuate (or nearly straight) sides at the base of pronotum: B. (A.) jelskii, B. (A.) abramalagae n. sp., B. (A.) straneoi n. sp. and B. (A.) procerus n. sp.In comparison with the other species, B. (A.) jelskii is distinguished by more deeply impressed elytral striae, more convex intervals, especially in ♂♂, and distinctly pear-shaped elytra.Moreover, the median lobe of the aedeagus of B. (A.) jelskii is nearly cylindrical, regularly curved and not ventrally depressed.
) straneoi n. sp. is one of the four Peruvian Agraphoderus species with sinuate (or nearly straight) sides at the base of pronotum.Respect to B. (A.) abramalagae n. sp. and B. (A.) jelskii, the sides are more weakly sinuate; moreover, B. (A.) straneoi n. sp. is distinguished from B. (A.) jelskii also by slightly impressed elytral striae and nearly flat intervals in both sexes (more impressed striae and more convex intervals in B. (A.) jelskii ♂♂) as well as not pear-shaped elytra.It differs from B. (A.) procerus n. sp. by the oval, less slender habitus, by the more convex elytra, with marked apical declivity, and by a larger aedeagus as well (B.(A.) procerus n. sp.displays a smaller aedeagus even in the largest specimens).
Blennidus from the following Museums and private Collections: MRST.
(Tschitschérine, 1897)s) species from the Southern Peruvian Andes: B. (A.) mesotibialis(Straneo, 1993)from an area between Puno and Cuzco, B. (A.) aulacostigma (Tschitschérine, 1897) and B. (A.) jelskii(Tschitschérine, 1897)from Puno.In this paper we raise this number to seven, as four new species from the Cuzco Dept.are described, all of them with probable affinities with B. (A.) jelskii.The affinities of B. (A.) mesotibialis and B. (A.) aulacostigma remain unclear.Unfortunately, almost nothing is known about the Bolivian Blennidus species, which could be southern relatives of the species belonging to the jelskii group.The Blennidus species belonging to the jelskii group are probably related to one another based on the uniform morphology of male genitalia, which are characterized in lateral view by a thin apex, distally tapering and not sinuate, and in dorsal view by a triangular apical blade as well as a large dorsal ostium.Inside this group, B. (A.) procerus n. sp., B. (A.) abramalagae n. sp., B. (A.) etontii n. sp. and B. (A.) straneoi n. sp., all distributed in the Cuzco Dept., are morphologically more similar to each other than to B. (A.) jelskii, which is also more isolated from a geographical point of view (Puno).