Documenting tenebrionid diversity: progress on Blaps Fabricius (Coleoptera, Tenebrionidae, Tenebrioninae, Blaptini) systematics, with the description of five new species

With about 250 species, the genus Blaps Fabricius, 1775 is one of the most diverse genera of darkling beetles (Coleoptera: Tenebrionidae: Tenebrioninae: Blaptini: Blaptina). In this study, we provide new insights on the evolutionary relationships of Blaps species using a combined molecular and morphological dataset encompassing 69 distinct Blaps species and subspecies (105 specimens in total, all belonging to the subgenus Blaps ), four other representatives of the tribe Blaptini (from the subtribes Gnaptorina, Gnaptorinina and Prosodina) and 12 outgroup species. Five new species of Blaps are also described within the subgenus Blaps : B. effeminata sp. nov. from Libya, B. intermedia sp. nov. from Morocco, B. maldesi sp. nov. from Algeria, B. nitiduloides sp. nov. from Algeria and Tunisia and B. teocchii sp. nov. from Tunisia. The results of the phylogenetic analyses indicate that the genus Blaps is likely paraphyletic; the two highlighted clades are morphologically distinct and correspond to groups previously referred to as sections (I and II) within the subgenus Blaps . This suggests the need for more phylogenetic studies in order to clarify the status of the various genera and subgenera belonging to the tribe Blaptini.

All species of Blaps are flightless and well adapted to semi-arid and arid environments because of several specific behavioural and morphological adaptations (Condamine et al. 2011).Subspecies of Blaps are generally morphologically well differentiated and may constitute species of their own (Ardoin 1973;Soldati 1994;Soldati et al. 2009;Condamine et al. 2011), as underlined by recent molecular phylogenetic analyses (Condamine et al. 2013).Previous biogeographical analyses found that geological events played an important role in their evolutionary history (Condamine et al. 2013).Because of their flightless habit, species of Blaps have limited dispersal abilities; in the context of the Mediterranean Basin, both the topographical and environmental heterogeneity, and the geographic isolation of many mountainous areas made them very prone to allopatric speciation (vicariance).This complex biogeographic context parallels the convoluted history of the taxonomy of Blaptini where numerous taxa have been alternatively treated as species or subspecies.
In this study, we conduct phylogenetic analyses relying on the largest dataset ever assembled for the subgenus Blaps (105 specimens belonging to 69 distinct species and subspecies are included), which includes three species that were not sampled in the study of Condamine et al. (2013).In contrast with the 2013 study, this dataset also includes four representatives of other Blaptini genera (Gnaptor Brullé, 1832, Gnaptorina Reitter, 1887and Prosodes Eschscholtz, 1829), which belong to the subtribes Gnaptorina Medvedev, 2001 (genus Gnaptor), Gnaptorinina Medvedev, 2001 (genus Gnaptorina) and Prosodina Skopin, 1960 (genus Prosodes).To root the tree, we used 12 tenebrionid species as outgroups (Table 1).For this study, we assess the phylogenetic relationships of the subgenus Blaps, and we also include the description of five new species of Blaps from the Mediterranean Basin.
Table 1.Taxon sampling.For each species and subspecies, the avialability of morphological data (the 'Morpho.' column) and the number of sequenced specimens are indicated.Additional taxonomic information for species of Blaps is also provided (section and species group for members of section I).Holotypes of the new species were deposited in the MNHN, whereas paratypes were deposited in HNHM, MCG, MNHN, CMF, CS and CT.Photographs of specimens were taken using a Canon EOS 450D digital single-lens reflex camera fitted with an EF-S 60 mm macro lens.Photographs of male aedeagi were taken using EntoVision multifocus system.

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In the Material examined sections, literal quotes from labels are placed within quotation marks.

Morphological dataset
The morphological dataset corresponds to the one published in the study of Condamine et al. (2011, see Appendix).This dataset consists of 47 morphological characters that have been scored for 62 species and subspecies of Blaps plus two other members of the tribe Blaptini (Gnaptor spinimanus (Pallas, 1781) and Prosodes jakowlewi Semenov, 1894).It is based on the examination of more than 2500 specimens; all characters are discrete, and multistate characters were treated as unordered (the annotated dataset is deposited in MorphoBank, homology of phenotypes over the web, under project number 2341, available from http://dx.doi.org/10.7934/P2341;see also http://morphobank.org/index.php).

Molecular dataset
The molecular dataset is based on that presented in the study of Condamine et al. (2013)

Phylogenetic analyses
Bayesian inference was used to infer phylogenetic relationships from a dataset combining the morphological and molecular matrices.This combined dataset encompasses 121 individuals (105 members of the genus Blaps, four other representatives of the Blaptini tribe and 12 outgroup species), of which nine are only included based on morphological characters.To improve the phylogenetic accuracy, we carried out partitioned analyses (Nylander et al. 2004).Partitions and substitution models for the molecular subset of the matrix were determined using PartitionFinder v. 1.1.1(Lanfear et al. 2012).
The Bayesian information criterion (BIC) was used for both partition and model selection (Ripplinger & Sullivan 2008).Based on the BIC results we used five partitions for the molecular subset (Table 2).
For the morphological data subset, we used an evolutionary model analogous to the JC69 model, except that it has a variable number of states (one-parameter Markov k-state model; Lewis 2001).Bayesian inference analyses were then carried out using MrBayes v. 3.2.6 (Ronquist et al. 2012).We conducted two independent runs with four MCMC (one cold and three incrementally heated) that ran for 50 million generations, with trees sampled every 5000 generations.A conservative burn-in of 25% was then applied after checking for stability on the log-likelihood curves and the split-frequencies of the runs.Support of nodes for MrBayes analyses was provided by clade posterior probabilities (PP) as directly estimated from the majority-rule consensus topology.Nodes supported by PP > 0.95 were considered strongly supported following Erixon et al. (2003).
Pronotum.Broadest in its middle, sides more strongly narrowed toward base than forward, not or barely sinuous before posterior angles and arcuately narrowed frontwards.Anterior angles rounded.Posterior angles blunt.External rim fine, complete on lateral margins, but obliterated in middle of front edge and base.Disc convex.Punctation fine, uniform.Carina of anterior foramen of pronotum thick and neckshaped below.Episternum of prosternum wrinkled near coxae and lighter toward margins.Prosternal apophysis oblique or steep and upright towards apex, S-shaped, just behind anterior coxae, then shortly denticulate in middle at apex.elytra.Ovoid, broadest around its middle, flat on disc.Upper surface leathery and covered with extremely fine and sparse punctures.Elytra costulate.Each elytron bearing 10 costae, alternate ones (3-5-7-9) slightly higher than others.Pseudopleural carina only visible on anterior half (in ♂♂ and ♀♀) from above, because of transversal convexity of elytra.Caudal extension at apex of elytra (mucro) measuring between 2.0 and 4.0 mm.Seen from above, right, parallel: deep suture flanked by two thick, Table 2. Results of PartitionFinder analyses, based on the BIC.transversely wrinkled ribs, which end in acute angular gap (Fig. 1E).In lateral view, mucro narrow, elongated, progressively narrowed to apex (Fig. 1F) and often bent upwards.Below, mucro grooveshaped, with external edges blurred and bottom transversally wrinkled.

Partitions Models
abdomen.Ventrites heavily wrinkled-punctate; anal ventrite finely and densely punctate, transversally impressed on disc, external rim complete and fine.
legs.Long and slender.Protibiae with internal face sinuous, external one straight in males; similar but less pronounced in females.Outer and posterior face of protibiae and posterior face of mesotibiae superficially grooved.Mesotibiae almost straight.Tarsi long, slender.
sexual dimorPHism.In males, no tuft of bristles between ventrites 1 and 2, and presence of light callosity near base of intercoxal process of ventrite 1.This callosity reduced to short transverse strip, with some longitudinal and oblique folds on front side, and coarse transversal wrinkles behind.In lateral view posterior declivity of elytra steeper in females.Mucro longer in males (3.5-4.0 mm) than in females (2.0-2.5 mm) (Fig. 1E, G).In males, rear edges of median and especially hind tibiae strongly denticulated.In males, metatibiae straight and flat on inner face; less pronounced in females.

Bionomics
Most known specimens were collected in antique ruins between April and October.

Distribution
Libya.This species appears to be endemic to the eastern part of Libya (formerly known as Cyrenaica).

Remarks
This species belongs to the Blaps gigas species group (sensu Condamine et al. 2011).It is morphologically close to Blaps polychresta Forskål, 1775 from Egypt, from which it differs by: (i) its stronger elytral costulation, especially on the disc; (ii) the absence of the abdominal hair tuft in males (between ventrites 1 and 2); (iii) in males, the presence of a very reduced abdominal callosity located near the base of the intercoxal process of ventrite 1. In
Pronotum.Broadest in its middle, sides distinctly sinuated before posterior angles and arcuatly narrowed frontwards.Anterior angles rounded.Posterior angles sub-right and blunt.External rim fine, complete on lateral margins, but obliterated in middle of front edge and base.Disc convex.Punctuation fine, uniform.Carina of pronotum anterior foramen thick and neck-shaped below.Episternum of prosternum shallowly wrinkled near coxae and obliterate toward margins.Prosternal apophysis vertically bent just behind anterior coxae, then very slightly denticulate in middle at apex.elytra.Ovoid, broadest around its middle, flat on disc.Upper surface leathery, covered with extremely fine and sparse punctures.Pseudopleural carina only visible from above because of tranversal convexity of elytra.Caudal extension at apex of elytra (mucro) measuring between 1.5 and 3.5 mm.Seen from above, mucro triangularly narrowed; deep suture flanked by two thick, coarsely transversely wrinkled ribs, converging toward apex, which ends in acute angular gap (Fig. 2E).In lateral view, mucro regularly narrowed up to apex, with lower face sub-horizontal (Fig. 2F); below, regularly but not deeply excavated, with sharp external edges and bottom transversally wrinkled.
legs.Mesotibiae curved.Inner face of pro-and mesotibiae superficially grooved.sexual dimorPHism.Yellow tuft of bristles between ventrites 1 and 2, and strong callosity between middle and front edge of intercoxal process of ventrite 1. Callosity directed backwards, bifid on top (when not eroded), with some longitudinal and oblique folds on front side, and coarse, transversal wrinkles hindwards.Pseudopleural carina of elytra only visible on anterior half in males, visible on anterior third in females.In lateral view, posterior declivity of elytra steep in males and subvertical in females.Mucro longer in males (2.0-3.5 mm) than in females (1.5-2.0 mm) (Fig. 2E, G).Male protibiae slightly flexuous; straight in females.In males, rear edges of median and especially hind tibiae strongly denticulate.Male metatibiae flexuous and shallowly emarginated on inner face; female metatibiae straight and simple.
aedeagus.Parameres open on sternal face (Fig. 2H), with apex acuminate.In lateral view (Fig. 2I), parameres thick and convex at base, then narrowed almost in straight line up to apex.

Bionomics
The examined material was collected between April and October.

Distribution
Morocco: Middle Atlas mountain range and further north, up to the surroundings of Taza and Aknoul.

Remarks
This species belongs to the Blaps emondi species group (sensu Condamine et al. 2011).Blaps intermedia sp.nov. is morphologically very close to B. debdouensis, and the aedeagus is the most reliable character to the two species: in B. intermedia sp.nov. the parameres are more extended laterally and appear inflated, whereas they are parallel and gutter-shaped in B. debdouensis.However, it is worth underlining that the aedeagus is often distorted by dehydration because of the very thin lateral tegument of the parameres.In the case of isolated females, the geographic distribution is a good criterion to distinguish females of Blaps intermedia sp.nov.from females of B. debdouensis.
Head.Clypeus arcuately emarginated, with lateral angles right and protruding frontwards, leaving clypeo-labial membrane partially visible.Clypeo-frontal suture reduced to thin, superficially engraved gloss line.Middle of frons (disc) with faint transverse depression.Mentum transverse, punctate and slightly impressed in middle of anterior edge.Gula microshagreened, matte.Labium deeply notched in middle of front edge.
Pronotum.Broadest in its middle, sides narrowed almost in straight line posteriorly, sinuated before posterior angles and arcuately narrowed toward front.Anterior angles rounded.Posterior angles obtuse and blunt.External rim fine, complete on lateral margins, but briefly obliterated in middle of front edge and base.Disc quite convex.Punctation fine, uniform.Carina of anterior foramen of pronotum thick and neck-shaped below.Episternum of prosternum wrinkled near coxae and obliterate toward sides.Prosternal apophysis vertically bent just behind anterior coxae, then flat and non-protruding posteriorly.elytra.Ovoid, broadest around its middle, flat on disc.Upper surface leathery and covered with extremely fine and sparse punctures.Pseudopleural carina only visible on anterior half from above because of transversal convexity of elytra.Caudal extension at apex of elytra (mucro) measuring less than 2.0 mm.Deep suture of mucro flanked by two thick, coarsely transversely wrinkled ribs, converging toward apex, apex ends in narrow apical gap (Fig. 3E).In lateral view, mucro regularly sloping up to apex (Fig. 3F).Below, mucro regularly excavated, with sharp external edges and bottom transversally wrinkled.
abdomen.Ventrites wrinkled-punctate; anal ventrite densely punctate, especially along external rim, external rim complete and quite thick.In one case (paratype), external rim of anal ventrite shortly interrupted in middle of apex.legs.Protibiae long and flexuous, especially in males.Mesotibiae curved.Inner face of pro-and mesotibiae widely grooved.Tarsi elongated; claws long.Hind tarsi approximately as long as one half of length of corresponding tibiae.sexual dimorPHism.Yellow tuft of bristles between ventrites 1 and 2, and a callosity in middle of intercoxal process of ventrite 1. Callosity directed backwards, with some oblique folds on front side.Elytra slightly larger than pronotum in males.In lateral view, posterior declivity of elytra steep in males and subvertical in females.Mucro longer in males (1.5-2.0 mm) than in females (0.5 mm) (Fig. 3E, G).Seem from above, mucro triangularly narrowed in males and acuminate in females.Rear edges of median and especially hind tibiae strongly denticulate in males.Male metatibiae flexuous and shallowly emarginated on inner face; female metatibiae straight and simple.aedeagus.Parameres bottleneck-shaped on sternal face (Fig. 3H), with apex acuminate.In lateral view (Fig. 3I), parameres thick and convex at base, then narrowed almost in a straight line up to apex.

Bionomics
The three known specimens were taken in the cedar forest of S'Gag (Algeria).

Distribution
Algeria.This species is currently known only from the type locality of S'Gag in the Aurès region.

Remarks
This species belongs to the Blaps emondi species group (sensu Condamine et al. 2011).It is morphologically most similar to B. teocchii sp.nov.from Tunisia.Blaps maldesi sp.nov. is most clearly separated from B. teocchii sp.nov.by having shorter antennae and tarsi, a wider pronotum and by the presence of a tuft of yellow bristles between abdominal ventrites 1 and 2. Until now, these two species have passed unnoticed in the collections because they superficially look like small specimens of other species of the emondi group, especially when the specimens are prepared in the old-fashioned way, with the legs tucked under the body.Compared to other species in the emondi group, these two species are characterized by the following combination of characters: slender legs; long and flexuous fore-tibiae, with inferior face distinctly grooved; tarsi long; and bottleneck-shaped parameres of aedeagus.
Head.Covered with fine punctation, more pronounced than on rest of upper body surface.Punctation becomes even denser on clypeus.Clypeus arcuately emarginated, with anterior angles right and protruding frontwards, leaving clypeo-labial membrane partially visible.Clypeo-frontal suture reduced to thin, brilliant line.In middle of frons, just behind eyes, two shallow, circular depressions, sometimes joined by short transverse depression.Mentum transverse, densely and coarsely punctate.Gula microshagreened, matte.Labium deeply notched in middle of front edge.
Pronotum.Usually weakly transverse, not wider than long, broadest at or just behind its middle.Disc slightly convex.Sides regularly arcuate, except at posterior angles.Posterior angles obtuse and blunt.Anterior angles rounded.External rim complete on lateral margins, but obliterated in middle of front edge and base.Punctation extremely fine, sometimes barely visible.Carina of anterior foramen of pronotum very thick and neck-shaped.Episternum of prosternum superficially wrinkled.Prosternal apophysis vertically bent just after anterior coxae, then flat and generally non-protruding posteriorly.elytra.Ovoid, broadest around its middle.Upper surface smooth and covered with extremely fine and sparse punctures; background integument shining.Pseudopleural carina just visible on anterior half from above because of transversal convexity of elytra.Posterior declivity of elytra steep in lateral view.Mucro at apex of elytra measuring between 2.0 and 5.0 mm.Seen from above, deep suture flanked by two thick, coarsely transversely striated ribs, converging in curve toward apex, apex ending in deep and narrow apical gap (Fig. 4E).In lateral view, mucro thick on ¾ of its length, then abruptly sloping toward apex (Fig. 4F).Basin-shaped below, with bottom coarsely striated transversely.abdomen.Ventrites wrinkled; anal ventrite punctate, with light depression on disc, external rim complete.legs.Quite short and stout.Protibiae slightly curved.Tarsi short.sexual dimorPHism.No tuft of bristles between ventrites 1 and 2, but strong callosity present in middle of intercoxal process of ventrite 1 in males.Mucro longer in males (2.5-5.0 mm) than in females (2.0-2.2 mm) (Fig. 4E, G).Male mesotibiae curved, slightly grooved on upper face.Male posterior tibiae flexuous and shallowly emarginated in middle of inner side, straight and simple in females.

Bionomics
The examined material was collected between April and September.

Distribution
Algeria and Tunisia.This species is currently only known from Tunisia and the Constantine region of Algeria.

Remarks
This species belongs to the Blaps emondi species group (sensu Condamine et al. 2011).It is morphologically related to B. emondi Solier, 1848 andB. binominata Escalera, 1914.In B. binominata the characteristic mucro is very broad and parallel, with a strong apical gap at a right angle.The male of B. emondi exhibits a tuft of yellow bristles between ventrites 1 and 2, and the abdominal callosity is located near the front edge of the intercoxal process of ventrite 1.It is also worth highlighting that Blaps nitiduloides sp.nov. is not found in sympatry with either B. binominata or B. emondi; the latter occurs in central and western Algeria, north of the 'Hauts Plateaux', and northeastern Morocco, whereas the distribution of B. binominata extends from the Oran region to the northeast of Morocco and the Spanish enclave of Melilla.

Etymology
This species is named in honour of Pierre Teocchi, cerambycid specialist, to whom one of us (L.Soldati) owes having become an entomologist.
Head.Clypeus arcuately emarginated, with lateral angles right and protruding frontwards, leaving clypeolabial membrane partially visible.Clypeo-frontal suture reduced to thin gloss line.Middle of frons (disc) with faint transverse depression.Mentum transverse, coarsely punctate and slightly impressed in middle of anterior edge.Gula microshagreened, matte.Labium deeply notched in middle of front edge.
Pronotum.Broadest in its middle, with sides narrowed almost in straight line posteriorly and arcuately towards front.Anterior angles rounded.Posterior angles obtuse and blunt.External rim fine, complete on lateral margins but obliterated in middle of front edge and base.Disc slightly convex.Punctation fine, uniform, as on head.Carina of anterior foramen of pronotum thick and neck-shaped below.Episternum elytra.Ovoid, broadest around its middle, depressed on disc.Upper surface smooth and covered with very fine and sparse punctures; background integument semi-gloss.Caudal extension at apex of elytra (mucro) measuring between 1.0 and 2.5 mm.Seen from above, mucro almost subparallel; deep suture flanked by two thick, coarsely transversely wrinkled ribs, slightly converging toward apex, apex ending in narrow apical gap (Fig. 5E).In lateral view, mucro regularly sloping up to apex (Fig. 5F).Seen from below, mucro regularly excavated, with sharp external edges and bottom transversally wrinkled.
legs.Protibiae long and flexuous.Mesotibiae curved.Inner face of pro-and mesotibiae widely grooved.Tarsi elongated and stout; claws long.Hind tarsi as long as ¾ of length of corresponding tibiae.sexual dimorPHism.No yellow tuft of bristles between ventrites 1 and 2, but strong callosity present between middle and anterior ridge of intercoxal process of ventrite 1 and directed backwards, with some oblique folds on front side.Elytra slightly larger than pronotum in males.Pseudopleural carina only visible on anterior half from above in males, or on anterior third in females, because of transversal convexity of elytra.In lateral view, posterior declivity of elytra steeper in females.Mucro longer in males (2.5 mm) than in females (1.0 mm) (Fig. 5E, G).Rear edges of median and especially hind tibiae strongly denticulate in males.Male metatibiae flexuous and shallowly emarginated on inner face at middle third; straight and simple in females.
aedeagus.Parameres bottleneck-shaped on sternal face (Fig. 5H), with apex acuminate.In lateral view (Fig. 5I), parameres thick and convex at base, then narrowed almost in straight line up to apex.

Distribution
Tunisia.So far known only from the type locality of Zaghouan in Tunisia.

Remarks
This species belongs to the Blaps emondi species group (sensu Condamine et al. 2011).It is morphologically very similar to B. maldesi sp.nov.from Tunisia (see above for more information).

Phylogenetics analyses
Overall, the phylogenetic analyses resulted in a relatively well-supported tree (70.8% and 43.3% of nodes supported by PP > 0.50 and 0.95, respectively; see Fig. 6).Nodal support was usually low (PP < 0.50) for the terminal parts of the tree encompassing taxa for which only morphological information was available (but see also Blaps magica Erichson, 1841 andB. antennalis Allard, 1880).
The monophyly of the tribe Blaptini was recovered with strong support (PP = 0.96).Of particular interest is the fact that the genus Blaps is not monophyletic in our phylogenetic reconstruction, due to the placement of the four other members of the Blaptini tribe.The latter are recovered in a weakly supported clade (PP = 0.49) sister to a large, well-supported clade (PP = 0.95) that encompasses all sampled representatives of the subgenus Blaps belonging to section I.For their part, all sampled members of the subgenus Blaps belonging to section II are recovered in a weakly supported clade (PP = 0.31), which Fig. 6.Bayesian maximum consensus tree resulting from the analysis of the combined molecular and morphological dataset carried out with MrBayes.Support of nodes is indicated using empty circles (for PP > 0.50) and filled circles for PP > 0.95).Species groups for Blaps belonging to section I are highlighted using transparent coloured frames.Paraphyletic species groups are highlighted using bracketed names; orange arrows are also used to highlight the placement of the five new species.For illustrative purposes, the habitus of some adults are figured on the right side of the figure; on the left upper side, the habitus of the five new species are also presented (all photographs were taken by L. Soldati).
is sister to all other sampled Blaptini.Within section I of subgenus Blaps, half of the species groups (considering only groups for which we have more than one sampled representative) are recovered monophyletic, namely the species groups alternans, cordicollis, gigas, magica and peyerimhoffi.All species and subspecies for which several representatives are included are recovered monophyletic, with the noticeable exceptions of Blaps approximans Seidlitz, 1893 (due

Discussion
The moderate support for our phylogenetic reconstruction can likely be accounted for by the amount of missing data in our dataset, especially given the fact that no molecular data was available for nine taxa.The low rate of PCR success results from the fact that we mostly processed specimens collected more than 10 years ago (PCR failed for more than 50 specimens).Because numerous species of interest are distributed in areas that are now hardly accessible, we think that future phylogenetic studies on species of Blaps (especially for the species that are distributed in Northern Africa) will require the intensive use of museomics approaches (see, e.g., Tin et al. 2014;Kanda et al. 2015).
Though the inferred basal phylogenetic relationships within sampled representatives of the tribe Blaptini are not well supported overall (for instance, the sampled Blaps from section II are recovered monophyletic, but with weak support), the inferred paraphyly of the genus is likely not artefactual.Indeed, its possible paraphyletic status does not come as a surprise given the taxonomic history of the tribe Blaptini, where numerous species were originally described in the genus Blaps, including species that are now assigned to genera belonging to other subtribes (Löbl et al. 2008).As a result, the subtribe Blaptina is possibly also paraphyletic, as it encompasses two clades (one for the sampled representatives of section I and one for the sampled representatives of section II of the subgenus Blaps) that are separated by representatives of three distinct subtribes of Blaptini.Obviously, because two of the three basal nodes are weakly supported, these results should be taken cautiously and highlight the need for more molecular data and a denser sampling for the tribe Blaptini.Since the subtribe Blaptina currently consists of 11 genera, our results emphasize the need for phylogenetic studies with a more comprehensive sampling at the genus level in order to properly redefine the higher systematics of the tribe Blaptini.
Within section I of subgenus Blaps, our results highlight the paraphyletic nature of several species groups.In contrast with the results of Condamine et al. (2013), in this study the alternans group is recovered monophyletic, while the propheta group is recovered paraphyletic due to the placement of individuals of B. tripolitanica Karsch, 1881.This paraphyly is likely artefactual, and it is only moderately supported in our analyses (PP = 0.63 and 0.72 for the nodes leading to the other representatives of the propheta group).The discrepancies between the two studies probably result from the inclusion of additional taxa and molecular data, in combination with the use of more thorough partitioning strategies.In our study, the lusitanica species group is also recovered paraphyletic as a result of the inclusion of a representative of B. tichyi Martínez Fernández, 2010.With reference to the latter, we think that the position of Blaps tichyi (sister to B. murati Peyerimhoff, 1943) is also artefactual because of the lack of character overlap between the two taxa (no molecular data is available for B. murati and, conversely, no morphological data is available for B. tichyi).Regarding the remaining three paraphyletic species groups (bifurcata, emondi and nitens), our results stress the need for a reappraisal of the boundaries of several species groups.For instance, the inclusion of Blaps superstitiosa Erichson, 1841 within the nitens group is to be considered as being based on the results of the molecular analyses, despite the fact that this species does not share the unique synapomorphy of the group (i.e., a straight posterior tibia).Similarly, the inclusion of Blaps maroccana Seidlitz, 1893 andB. pinguis Allard, 1880 as members of the emondi group might be warranted, but will require redefining the combination of character states that describe/delineate the clade.
At the species level our phylogenetic framework allows us to discuss the placement of the five newly described species, four of which belong to the emondi group.In the case of Blaps intermedia sp.nov., the results of the phylogenetic analyses support our view of considering this taxon as a distinct species within the emondi group, not directly related to B. debdouensis.The placement of Blaps nitiduloides sp.nov.(clustered with B. pinguis) came as a surprise, because this species is morphologically most similar to B. binominata and B. emondi; we cannot exclude that this position is artefactual, especially given the fact that it is only weakly supported (PP = 0.79).Regarding Blaps maldesi sp.nov.and B. teocchii sp.nov., both species are clustered together as sister to all remaining members of the emondi group (plus the only representative of the pinguis group); this placement is not surprising given that both species share unique combinations of character states (see above).Within the gigas group, Blaps effeminata sp.nov. is recovered as sister to a clade encompassing B. doderoi and B. wiedemannii, while being morphologically closer to B. polychresta; that being said, the corresponding node is only weakly supported (PP = 0.69).
Our results also provide further evidence to support the hypothesis that most subspecies of Blaps correspond to distinct valid species; this is especially the case for the subspecies that do not cluster together in the phylogenetic tree, such as B. nefrauensis nefrauensis Seidlitz, 1893 andB. nefrauensis vespertina Koch, 1937, or for the subspecies of B. nitens, which do not constitute a monophyletic group due to the placement of individuals of B. sulcifera and B. superstitiosa.To advance on this issue, more morphological and molecular studies, as well as the sequencing of additional specimens, are definitely required.

Table 1 .
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to the placement of an individual of B. haberti Peyerimhoff, 1943) and B. nitens nitens Laporte, 1840 (due to the placement of an individual of B. nitens requieni Solier, 1848).Regarding the placement of the new species, it is worth underlining that although Blaps intermedia sp.nov. is morphologically very similar to B. debdouensis, it is more closely related to B. appendiculata, while B. debdouensis is more closely related to B. emondi.