A new genus of Staphylinidae (Coleoptera) from the Lower Cretaceous: the first fossil rove beetles from the Southern Hemisphere

Systematic Entomology (2012), 37, 379–386

A new genus of Staphylinidae (Coleoptera) fromthe Lower Cretaceous: the first fossil rove beetlesfrom the Southern Hemisphere

A N D R E A S C H O M A N N and A L E X E Y S O L O D O V N I K O V

Department of Entomology, Zoological Museum, Natural History Museum of Denmark, Universitetsparken, Copenhagen,Denmark

Abstract. The fossil genus of rove beetles Apticax gen.n. with two new species,A. volans sp.n. and A. solidus sp.n., is described from the Nova Olinda Member ofthe Crato Formation in north-eastern Brazil (Aptian–Albian, dated as 125–99.6 Maold). Both species belong to the clade Staphylininae + Paederinae in the staphylininegroup of subfamilies and are the first fossil true Staphylinidae to be described from theentire Southern Hemisphere. Although they resemble Paederinae, the low number ofsatisfactorily preserved characters do not allow definite placement of either A. volansor A. solidus in any of the subfamilies of Staphylinidae.

Introduction

The Crato Formation in northeastern Brazil includes a seriesof laminated limestones up to 13 m thick known as theNova Olinda Member that contains well-preserved fossils ofanimals and plants (Martill et al., 2007). This formation isdated as late Aptian/early Albian (Batten, 2007; Martill et al.,2007; Heimhofer et al., 2010) in the Lower Cretaceous strata(Aptian–Albian period, 125–99.6 Ma, according to Gradsteinet al., 2004), with Nova Olinda being the oldest member (mostlikely late Aptian, according to Batten, 2007). Being depositedclose to the heart of the ancient supercontinent Gondwanaand containing many remarkably well-preserved fossils, theCrato Formation is one of the best windows into an ancientecosystem of that landmass (Martill et al., 2007). The location,relative old age, and good preservation make any rove beetlefossils from that formation (reviewed in Wolf-Schwenninger& Schawaller, 2007) an important discovery.

In Wolf-Schwenninger & Schawaller (2007) there are sixspecimens of Staphylinidae mentioned:

Specimen 1, Cratophyllina minuscula Martins-Neto, 2002,is described in a doctorate thesis, but not published, thus anomen nudum.

Correspondence: Andrea Schomann, Department of Entomology,Zoological Museum, Natural History Museum of Denmark, Uni-versitetsparken 15, 2100 Copenhagen, Denmark. E-mail: [email protected]

Specimen 2, Caririderma pilosa Martins-Neto, 1990, isvalidly described (Martins-Neto, 1990) and originally placedin Dermaptera.

Both were listed under Staphylinidae in Martins-Neto(2005). Having seen a drawing of C. minuscula by Martins-Neto (A. Newton and R.G. Martins-Neto, personal commu-nication) it can be said that this might not be a beetle,and Martins-Neto also stressed in the personal communica-tion that there are more than 11 antennal segments, whichwould be atypical for Coleoptera. The drawing of C. pilosain Martins-Neto (1990: 782, fig. 3) has been classified ascertainly not Dermaptera by Haas (2007), but, based on themorphology of head, thorax and abdomen, it is also very proba-bly not Coleoptera (the elytra, if ever present, are unfortunatelymissing).

Specimens 3 and 4, previously undescribed, are the subjectof this paper, Apticax gen.n. with two new species (Figs 1, 2),and are definitely Staphylinidae (discussion see below). Oneof these specimens, here described as Apticax volans gen.n.and sp.n. appears in the same book (Martill et al., 2007) onplate 13e, although it is less exposed from the matrix.

Specimens 5 and 6 are undescribed, but depicted in Grimaldi& Maisey (1990: 7, fig. 2E) and Grimaldi & Engel (2005:376, fig. 10.28; also depicted in Grimaldi & Maisey, 1990,fig. 2D, in lower quality) and also certainly Staphylinidae.Their rather compact habitus strikingly resembles Apticax,and they, too, are rather large rove beetles. The specimenin fig. 10.28 (Grimaldi & Engel, 2005) nevertheless seems tohave more affinities to the subfamily Staphylininae, whereasthe specimen in fig. 2E (Grimaldi & Maisey, 1990) could be

© 2012 The AuthorsSystematic Entomology © 2012 The Royal Entomological Society 379

380 A. Schomann and A. Solodovnikov

A B

C D

Fig. 1. Apticax volans gen.n. & sp.n. (A) Habitus; (B) elytra; (C) anterior part of head (obs. shapes of mandible and labrum); (D) fore and middlelegs (obs. dilated protarsus and spines at middle tibia).

A B

C D

Fig. 2. Apticax solidus sp.n. (A) Habitus; (B) head from dorso-lateral (obs. maxillary palpus); (C) middle leg (obs. spines on tibia); (D) pronotumfrom lateral (obs. hypomeron and well-developed postcoxal process, arrow).

© 2012 The AuthorsSystematic Entomology © 2012 The Royal Entomological Society, Systematic Entomology, 37, 379–386

A new genus of Staphylinidae (Coleoptera) from the Lower Cretaceous 381

something close to the subfamily Osoriinae (see discussion inthe comparative section for Apticax gen.n.).

The two specimens of Apticax superficially look likeeither a paederine or some rather unusual staphylinine (thesubfamily sister to Paederinae), or members of a hithertounknown lineage that would be phylogenetically at leastclose to the paederine–staphylinine stem group, with anintermediate character combination. They would thus either bethe oldest Paederinae so far described (according to EDNA,2011; the oldest known Paederinae were from Baltic Amber,54.8–23.8 Ma, Pasnik & Kubisz, 2002), or a remarkablefind to illuminate the evolution of the staphylinine group(sensu Lawrence & Newton, 1982, 1995; Thayer, 2005;Grebennikov & Newton, 2009). These specimens are thefirst fossil representatives of Staphylinidae from the formersupercontinent Gondwana (the Caribbean plate being ofdifferent tectonic origin; see, e.g., Meschede & Frisch, 2002;James et al., 2009) and the entire Southern Hemisphere to bedescribed.

As there are not enough characters visible for a rigorousphylogenetic analysis, we refrained from the formal assignmentof Apticax to a subfamily within Staphylinidae. It is left asincertae sedis in the clade Staphylininae + Paederinae (afterGrebennikov & Newton, 2009) of the staphylinine group.Such placement seems plausible when characters observablein Apticax are compared to the full set of characters optimizedon the most parsimonious tree topology in the latest accounton the phylogeny of the staphylinine group of subfamilies(Grebennikov & Newton, 2009, fig. 11; comparison in Fig. 3here, Apticax is not placed in the tree). With an imaginaryplacement of Apticax in the clade Staphylininae + Paederinaeon the tree diagram of Grebennikov & Newton (2009),characters with states conflicting with those in Apticax occuron more basal branches, but these conflicts are resolvedmore distally towards the clade Staphylinae + Paederinae(‘transitional conflicts’). By similar imaginary experiments, theplacement of Apticax in any other clade can be ruled out as anyof these alternative positions causes ‘permanent conflicts’, i.e.character conflicts not being resolved in any of the terminals(Fig. 3; further explanation in the respective figure caption).Oxyporinae, Steninae and Neooctavius in Euaesthetinae haveone supporting character each and no additional conflict onthe respective branch, but also here are unresolved conflictson more basal branches. Apticax can also be excluded fromall three subfamilies because of their respective habitus (formof head, head–pronotum size relationship), the very prominentoxyporine mandibles, and stenine and euaesthetine form ofantennae, maxillary palps etc. being completely different fromthose of Apticax.

Material and methods

The fossil specimens are deposited at the Staatliches Museumfur Naturkunde Stuttgart, Germany.

The specimens were examined using a Leica MZ APO dis-secting microscope; the measurements were taken, employing

a Leica ocular micrometer, as follows: HL: head length fromapical margin of clypeus to weak furrow demarcating begin-ning of neck in A. solidus, or to anterior margin of pronotumin A. volans (in which the pronotum covers the neck regionwhere such furrow is expected); PL: length of pronotum alongmid-line; EL: length of elytra from humerus to most distalapical margin; EW: maximal width of one elytron; TBL: totalbody length from apical margin of clypeus to abdominal apex(approximate position).

Due to a diagonal distortion of both specimens, especiallyof their head and thorax, some of the usual measurementswere impossible to carry out. Insufficient preservation and onlypartial exposure from the matrix is also the reason why severalcharacters could not be observed. Photographs were takenwith a Leica DFC 420 camera attached to a Leica MZ16Amicroscope (lighting by Leica CLS 150XE) with the help ofLeica Application Suite (Leica Microsystems, 2003–2007) andAutomontage Pro (Synoptics Ltd, 1997–2004).

Systematic palaeontology

Apticax volans gen. & sp.n. and A. solidus sp.n. share severalstructural features that are usually considered important in rovebeetle systematics. At the same time, they differ so significantlyin detail that, if they were from a recent fauna with more mate-rial available and the possibility of a better genus definitionand delineation, they would probably be assigned to differentgenera. Contrary to recent specimens, many characters impor-tant for the genus definition in Staphylinidae cannot be seen inthese fossils due to their incomplete preservation. Therefore,we assign them to the same genus for now, even though thismight eventually become untenable. If additional material isdiscovered, it will be easier to split a vaguely defined genusinto two or more better characterized genera, than to deal withtwo or more poorly defined genera to begin with.

Family: Staphylinidae LatreilleSubfamily incertae sedis

Apticax gen.n.

Type speciesApticax volans sp.n., by present designation

Taxonomic placement and comparisonSince we are unable to perform a numerical parsimony-

based phylogenetic analysis due to shortage of well-preservedcharacters in Apticax, our hypothesis of its taxonomic place-ment within Staphylinidae is explored and justified by thecomparison of characters observable in Apticax (Table 1) withthe optimized characters in the preferred tree topology of thelatest relevant phylogeny by Grebennikov & Newton (2009;further below abbreviated as G&N, with relevant characterand character states numbers, also fully explained in theTable 1 here). Following this approach Apticax is assignedto the staphylinine group based on: shape of anterior margin



Staphylinine group (incl. subtrees A and B)

permanent conflict

transitional conflict

agreement

194

0

196

2 [1]

192

1 [0]

194

1 [0] 107

0

150

1

186

0 [1]

141

3

196

3 [1] 192

1 [0]

194

1 [0]

105

0 [1]

107

0

141

3

186

0 [1]

196

0 [1]

196

2 [1]

196

1

151

1

106

1

150

1

151

0 [1] 106

0 [1]

196

1105

1

196

3 [1]

186

1

196

2 [1] 196

2 [1]186

0 [1]

192

1 [0]

150

1

106

1

107

1 [0]

151

1

186

0 [1]

196

0 [1]

107

2 [0]

196

2 [1]

151

1

141

1 [3]

Fig. 3. Phylogeny of the staphylinine group after Grebennikov & Newton (2009: fig. 11), with character mapping reduced to characters observablein Apticax gen.n. (not included as terminal taxon in the tree); character and character states numbers (also listed in Table 1) are given as in theoriginal, states for Apticax in square brackets. Colour-coded circles (see legend) give the degree of congruence between the character states observedin Apticax and those in the original (Grebennikov & Newton, 2009) character map. ‘Permanent conflict’ indicates that Apticax has a characterstate different from the state on the branch in question and more distally, i.e. with all terminals in the clade subtended by the branch; ‘transitionalconflict’ means that Apticax has a different state from the optimisation on the given branch, but it has the same state as on some of the terminalswithin the given clade.

of labrum (G&N 107-0); short elytra with six abdominalterga exposed (G&N 141-3); and general habitus (not codedin G&N) resembling recent members of Staphylininae andPaederinae. Within the staphylinine-group, Apticax best fitsthe clade consisting of Paederinae and Staphylininae, withtwo synapomorphic characters: bilobed labrum (G&N 106-1);presence of vein MP3 on the wing (G&N 150-1); and againtheir habitus. Because of the concealed antennal insertions, thepresence of a postcoxal process in the pronotal hypomeron(visible only in A. solidus), and the form of the head, Apti-cax seems to have closer ties to Paederinae (Apticax has asclerotized, smooth anterior margin of the labrum, G&N 107-0,

whereas a sclerotized, denticulate or serrate anterior marginof the labrum, G&N 107-1, is given for Paederinae, althoughstate 0 is also not exceptional in that subfamily). In habitus,it resembles some recent members of the tribe Paederini andshares with them the small, conical fourth maxillary palpomere.

The habitus, shape of head, very vague neck constriction, aswell as the structure of labrum and mandibles (visible only inA. volans) are character states that are similar to those observedin the recent paederine genera Dicax Fauvel, Macrodicax Leaand Baryopsis Fairmaire and Germain, all preliminarily placedin Cryptobiina by Newton (1985) as defined in Herman (1981).The new genus differs from all three in the even weaker



Table 1. Characters and states (0–3) used by Grebennikov & Newton (2009) and observable in Apticax gen.n.

G&N no. Character 0 1 2 3

105 Labrum, shape from above Subquadrate tomoderately transverse(up to 1.5×)

Strongly transverse(2× and above)

— —

106 Labrum, whether bilobed Not bilobed Bilobed — —107 Anterior margin of labrum Sclerotized, smooth Sclerotized, denticulate

or serrateWith transparent apical

membrane—

141 Elytral length relative toabdomen

Entire (abdomencompletely concealed)

Slightly truncate(1–2 terga exposed)

Strongly truncate(3–5 terga exposed)

Short(6–7 tergaexposed)

150 Wing venation, vein MP3 Absent Present — —151 Wing venation, vein MP4 Absent Present [poorly visible] — —186 Basolateral ridges of

abdominal tergites III–V(in Solierius IV–V)

Present Absent — —

192 Pairs of parasclerites ofabdominal segment III

Two One Absent —

194 Pairs of parasclerites ofabdominal segmentsIV–VI

Two One Absent —

196 Pairs of parasclerites ofabdominal segmentsIV–VI

Two, longitudinallyseparated

Two, transverselyor obliquelyseparated

One Absent

Character state of Apticax in boldface.

neck constriction and significantly smaller fourth maxillarypalpomere; from Baryopsis and Macrodicax in the shape of thepronotum, longer elytra, straighter and less spherical shape ofthe head, and having mandibles more robust than in Baryopsisbut slenderer than in Macrodicax ; from Dicax and Baryopsisin the stronger tibial spines and their occurrence also on thehind legs, and the lateral placement of the eyes; from Baryopsisin the shape of the labrum.

From all the previously described paederine fossil species(two Cryptobium, Scudder, 1890; one Achenium, Oustalet,1874; 20 Lathrobium, Giebel, 1856; Heer, 1862; Scudder,1876, 1890, 1900; Wickham, 1913a; Abdullah & Abdullah,1968; Zhang, 1989; Zhang et al., 1994; Pasnik & Kubisz,2002; two Lithocharis, Heer, 1856; Scudder, 1900; oneMiolithocharis, Wickham, 1913b; one Medon, Zhang, 1989;one Sunius, Wickham, 1913a; one Paederus, von Heyden &von Heyden, 1866; from North America, Europe and China;oldest 54.8–23.8 Ma, youngest c. 0.1 Ma), Apticax differs inhabitus as well as in morphological details. All listed fossilPaederinae are roughly at least 50 Ma younger than Apticax,and thus they are much more modern in their appearance, beingmore slender in habitus, and relatively unproblematic to assignto extant subtribes or even genera. Contrary to Apticax, all ofthem are from the Northern Hemisphere and none of themresemble Dicax, Macrodicax or Baryopsis, which are generaendemic to the Southern Hemisphere.

The two undescribed fossils from the Crato Formation(Grimaldi & Maisey, 1990: 7, fig. 2E) and Grimaldi & Engel(2005: 376, fig. 10.28) do have a certain resemblance toApticax, especially in their size and robust habitus, but dif-fer at least in some important details. The pronotal shape of

the specimen on fig. 10.28 in Grimaldi & Engel (2005) issimilar to that of Apticax, but in contrast to Apticax it hasa much more ovate head with a clear neck constriction. Theantennae are somewhat moniliform and thus similar to thoseof A. volans, but seem to be inserted closer to each other, asobserved in Staphylininae. The second specimen (Grimaldi &Maisey, 1990: fig. 2E) is even more robust than Apticax. Theantennae are slender, like those of A. solidus, and seem to beinserted laterally, possibly concealed, also as in both species ofApticax. The head is obviously practically without neck con-striction like that of Apticax, but at the same time remarkablyshort and wide, although not as wide as the pronotum, so thatit is not so much like that of staphylinine group representa-tives, but rather like that of Osoriinae and related subfamilies.Contrary to Apticax, the pronotum seems to be more roundedlaterally and is narrowed anteriorly.

The new genus also somewhat resembles Arrowinini andPlatyprosopini, two extant tribes of Staphylininae. However, itdiffers from both of them in concealed antennal insertions,size and shape of the apical maxillary palpomere, and instructure of the pronotum with distinct front angles and well-developed hypomeron with postcoxal process. Additionally,unlike Platyprosopini, Apticax does not have either a fullydeveloped neck constriction or a nuchal ridge, and it has alarger distance between the antennal insertions and a differentshape of the labrum. Contrary to Arrowinini it has noinfraorbital ridges.

Apticax volans shares striate elytra with Sulcelytrinusantiquus Tikhomirova, a late Jurassic fossil speciesfrom Karatau (Kazakhstan) with presumed Staphylininae–Paederinae affinities (Tikhomirova, 1968). But both species



strongly differ in shape of head and, unlike Apticax, thathas six visible abdominal segments (segments III–VIII),S. antiquus has seven visible segments because of the well-developed and free second abdominal segment (see fig. 15 inTikhomirova, 1968).

From Tunicopterus sigara Tikhomirova, another poorlypreserved late Jurassic fossil species from Karatau(Kazakhstan) with presumed Staphylininae–Paederinaeaffinities (Tikhomirova, 1968), Apticax significantly differs inhabitus. Apticax has larger body size, more elongate head andpronotum but shorter elytra, and it lacks rugose microsculptureon the body.

Comparison of Apticax with the Early Cretaceous genusMesostaphylinus Zhang is not easy. It is described from theLaiyang Formation, China and represented by two species,M. laiyangensis Zhang and M. fraternus Zhang, Wang & Xu,with clear Staphylininae or Paederinae affinities. Descriptionsof these seemingly well-preserved specimens are poor (Zhang,1988; Zhang et al., 1992), whereas the respective type materialis unavailable for study. However, as can be judged from theillustrations in Zhang (1988) and Zhang et al. (1992), unlikeM. laiyangensis, Apticax has a much more robust ‘bulldozer-like’ body shape, and unlike M. fraternus, which has sevenvisible abdominal segments because of the well-developed andfree second segment (see fig. 2 in Zhang et al., 1992), Apticaxhas only six visible abdominal segments (segments III–VIII).

Finally, Apticax is rather different in habitus fromMegolisthaerus chinensis Solodovnikov & Yue, a recentlydescribed relatively large Early Cretaceous rove beetlefrom Yixian Formation from China with unclear subfamilyaffiliation (Yue et al., 2010). Additionally Apticax differs fromMegolisthaerus in the shorter, more robust mandibles, muchshorter apical segment of maxillary palps, and in the absenceof baso-lateral ridges on the abdominal tergites.

DiagnosisThe combination of the extremely vague neck constric-

tion, pronotum embracing the neck by its anterior margin,concealed antennal insertions, small conical fourth maxillarypalpomere, obtusely crescent-shaped, bilobate labrum, and avery sturdy habitus make this genus unique in the cladeStaphylininae + Paederinae.

DescriptionBody elongate, about parallel-sided, moderately convex,

robust (Figs 1A, 2A).Head about as wide as pronotum, without or almost without

neck constriction, with only very flat furrow demarcatingbeginning of neck (best visible in A. solidus), dorsoventrallyflattened and with straight or almost straight sides; eyessuboval, medium large to large, positioned laterally in anteriorthird of head (seen from laterally, best visible in A. solidus);antennal insertions situated far apart from each other, con-cealed, first antennomere slightly longer than antennomeres IIand III together, at least 2.5 times as long as wide, distalnotch dorsal/dorsolateral; third maxillary palpomere distally

slightly expanded, straight, c. 2–2.5 times as long as wide,fourth palpomere small to very small, asymmetric, conical(best visible in A. solidus, Fig. 2B); labrum bilobate, obtuselycrescent-shaped (visible only in A. volans, Fig. 1C); mandiblesrobust with at least one subapical tooth (visible only inA. volans, Fig. 1C).

Pronotum about as long as wide, tapered to base, withdistinct anterior and posterior angles, anterior margin tightlyencloses posterior part of head; pronotal hypomeron wide,also in posterior part, with well-developed postcoxal pro-cess (not completely exposed; visible only in A. solidus,Fig. 2D, arrow).

Elytra about as long as wide, parallel sided, with distincthumeral angles, striate or smooth with small irregular punc-tuation. Wings present and well developed (visible only inA. volans, Fig. 1A, D, but well-developed humeri of elytraof A. solidus, Fig. 2A, indicate possibility of well-developedwings also in latter species), with veins MP3 and probablyalso MP4 (very weakly visible) present.

Legs relatively elongate; femora strong; at least middleand hind tibiae with strong spines (Figs 1D, 2C; hind tibiavisible only in A. volans); tarsi five-segmented, protarsi withtarsomeres I–IV dilated (visible only in A. volans, Fig. 1D).

Abdomen subparallel, with two pairs of paratergites presenton segments III–VII and traces of longer setation on moreposterior segments.

EtymologyThe masculine generic epithet is derived from a fusion of

‘Aptian’, the stage of the geochronological scale the two fossilsare most probably from, and ‘Dicax ’, one of the genera whichthis genus resembles the most.

Apticax volans sp.n.(Fig. 1)

Type materialHolotype: SMNS66452, Crato Formation, Nova Olinda

Member, Brazil.

DiagnosisDistinguished from A. solidus sp.n. by its slightly more

gracile habitus, smaller size, significantly shorter anten-nomeres, and striate elytra.

DescriptionMeasurements: HL: 1.2 mm; PL: 1.2 mm; EL: 1.45 mm;

EW: 0.75 mm; TBL: c. 6.5–7 mm (abdomen poorlypreserved).

Head probably as long as wide, loosely punctuated withfine and larger punctures, with two pairs of larger puncturesbetween antennal insertions situated close to each otherwith mesal pair set slightly posterior to lateral pair, eyeslateral; antennomeres slender to moniliform (Fig. 1A), first



antennomere c. 2 times as long as wide, slightly bulbous, distalnotch dorso-lateral, rather deep (Fig. 1C), antennomere IIc. 1.5 times as long as wide, antennomere III c. 2 times aslong as wide, antennomeres IV–VI of about equal length andslightly longer than wide, antennomeres VII–XI about as longas wide or even slightly transverse, antennomere XI seemsto be distally rounded or truncate; third maxillary palpomerec. 2 times as long as wide, fourth palpomere very small andasymmetrically conical, apparently shorter than wide.

Pronotum about as wide as head, fine punctuation not visible,but with several larger punctures, which might be interpretedas one series along either side of the middle and more irregularpunctuation laterally, slightly tapered at least in posterior half,possibly already from anterior margin.

Elytra about as long as wide, with punctuation arrangedin eight distinct rows of punctures on each elytron (Fig. 1B),wings present and well developed (Fig. 1A, D).

Abdomen with long strong setae from at least tergite VII.

EtymologyThe Latin verb in present participle ‘volans’ (flying) was

chosen as species epithet because of the well-preservedexposed wing and elytron, as if the specimen was caught inflight.

Apticax solidus sp.n.(Fig. 2)

Type materialHolotype: SMNS66453, Crato Formation, Nova Olinda

Member, Brazil.

DiagnosisDistinguished from A. volans sp.n. by its larger size and

more robust habitus, as well as by irregularly punctured (non-striate) elytra, and by significantly longer antennomeres.

DescriptionMeasurements: HL: 2.1 mm; PL: 2.25 mm; EL: 2.55 mm;

EW: c. 1 mm; TBL: c. 11.3 mm (abdomen not fully exposed).Head probably as wide as long, irregularly medium densely

punctuated; antennomeres slender (Fig. 2A), first antennomerec. 3 times as long as wide, distal notch dorsal, not very deep,antennomeres II–IX of about equal length and c. 2 times aslong as wide, antennomere X about 1.5 times as long as wide,XI c. 2 times as long as wide and anteriorly tapered; thirdmaxillary palpomere c. 2.5 times as long as wide, fourthpalpomere small and asymmetrically conical, about as longas wide (Fig. 2B).

Pronotum slightly wider than head, tapered basad at leastalong posterior half of its length, punctuation irregular andof about same strength and density as on head, although afew stronger punctures suggest one loose series of these alongeither side of middle.

Elytra slightly longer than wide, slightly narrower thanpronotum; with irregular punctuation seemingly weaker thanthat on head and pronotum; surface rather smooth; epipleuralridge appears to be present (but only vaguely and fragmentaryvisible).

Abdomen slightly narrower than elytra; tergites slightlydenser punctuated than head and pronotum, at least fromtergite VI with rather long, strong setae.

EtymologyThe Latin adjective ‘solidus’ (strong, compact, solid) was

chosen as species epithet due to the stout and massiveappearance of the new species.

Acknowledgements

We would like to thank the Staatliches Museum fur NaturkundeStuttgart, Germany, for the loan of the two fossil specimensdescribed here, and especially Karin Wolf-Schwenninger forbringing the material to our attention. We are very gratefulto Alfred Newton for the generous provision of his valuablepersonal communication with the unfortunately deceasedR. G. Martins-Neto.

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Accepted 15 November 2011


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A new genus of Staphylinidae (Coleoptera) from the Lower Cretaceous: the first fossil rove beetles from the Southern Hemisphere