Large carrion and burying beetles evolved from Staphylinidae (Coleoptera, Staphylinidae, Silphinae): a review of the evidence.

Large carrion beetles (Silphidae) are the focus of ongoing behavioral ecology, forensic, ecological, conservation, evolutionary, systematic, and other research, and were recently reclassified as a subfamily of Staphylinidae. Twenty-three analyses in 21 publications spanning the years 1927-2023 that are relevant to the question of the evolutionary origin and taxonomic classification of Silphidae are reviewed. Most of these analyses (20) found Silphidae nested inside Staphylinidae (an average of 4.38 branches deep), two found Silphidae in an ambiguous position, and one found Silphidae outside Staphylinidae, as sister to Hydrophilidae. There is strong evidence supporting the hypothesis that large carrion beetles evolved from within Staphylinidae and good justification for their classification as the subfamily Silphinae of the megadiverse, and apparently now monophyletic, Staphylinidae. Considerable uncertainty remains regarding the interrelationships and monophyly of many staphylinid subfamilies. Nonetheless, the subfamily Tachyporinae was found to be the sister of Silphinae in more analyses (7) than any other subfamily.

Kekveus brevisulcatus sp. nov., a new featherwing beetle from mid-Cretaceous amber of northern Myanmar (Coleoptera: Ptiliidae).

Ptiliidae (featherwing beetles) is a group of minute staphylinoid beetles with a scarce fossil record. Here a second member of the Mesozoic genus Kekveus Yamamoto et al. is reported from mid-Cretaceous Burmese amber, with detailed morphology obtained through confocal microscopy. Kekveus brevisulcatus Li, Yamamoto, Newton & Cai sp. nov. shares with K. jason Yamamoto et al. the unpaired medial pronotal fovea and narrowly separated transverse metacoxae, but can be separated from the latter based on its less elongate body, shorter pronotal foveae, and much weaker transverse depression on the head. Our phylogenetic analyses support the discheramocephalin affinity of Kekveus, although its relationship with other members of Discheramocephalini cannot be confidently resolved.

Revisiting the Raractocetus Fossils from Mesozoic and Cenozoic Amber Deposits (Coleoptera: Lymexylidae).

The fossils once assigned to Raractocetus Kurosawa from the Mesozoic and Cenozoic amber deposits differ from extant Raractocetus in the longer elytra, the more strongly projecting metacoxae, and the hind wing with vein 2A forked. Thus, these fossils should be removed from Raractocetus. Cretoquadratus engeli Chen from Kachin amber appears to be conspecific with R. fossilis Yamamoto. As a result, R. fossilis and R. extinctus Yamamoto from Kachin amber, R. balticus Yamamoto from Baltic amber, and R. sverlilo Nazarenko, Perkovsky & Yamamoto from Rovno amber are transferred to Cretoquadratus Chen, as C. fossilis (Yamamoto) comb. nov., C. extinctus (Yamamoto) comb. nov., C. balticus (Yamamoto) comb. nov., and C. sverlilo (Nazarenko, Perkovsky & Yamamoto) comb. nov., and C. engeli syn. nov. is suggested to be a junior synonym of C. fossilis.

Integrated phylogenomics and fossil data illuminate the evolution of beetles.

Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic-Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.

Dasycerine rove beetles: Cretaceous diversification, phylogeny and historical biogeography (Coleoptera: Staphylinidae: Dasycerinae).

Within the hyperdiverse beetle family Staphylinidae, Dasycerinae is one of the smallest and most cryptic subfamilies, comprising a sole extant genus characterized by a latridiid beetle-like body form. Little has been known about their early diversification, character evolution, phylogeny and historical biogeography because of limited fossil material and lack of a phylogeny integrating extant and extinct representatives. Here we report an unexpectedly diverse dasycerine fauna from the mid-Cretaceous of northern Myanmar, including a new genus and four new species. To reconstruct the early evolutionary history of Dasycerinae, we present a phylogenetic framework of the subfamily based on a dataset integrating all extant and extinct taxa using parsimony, maximum-likelihood and Bayesian methods. †Cedasyrus gen. n., characterized by distinct sexual dimorphism in antennal and elytral lengths, is recovered as the basal-most lineage, sister to the remaining two extinct genera and all living Dasycerus species. †Vetudasycerus is recovered as sister to †Protodasycerus + Dasycerus. Among all extinct taxa, †Protodasycerus bears distinctly longer elytra, and appears to represent a transitional form from †Vetudasycerus to Dasycerus. Phylogenetic inferences and ancestral distribution reconstruction support an "Out-of-Orient" model for Dasycerinae. Either the Bering- or North Atlantic Land Bridge may have served as a passageway for dasycerine dispersal between Eurasian and North American continents. An elevation-reconstruction analysis indicated that the ancestor of the extant Dasycerus probably lived at a high altitude and stayed at this elevation through the end of the Miocene. We propose that the extinction of dasycerine ancestors living on the Tethyan islands at low altitude was likely caused by sea-level rise and climatic warming during the Late Cretaceous. The high-altitude areas might have played the role of refugia that harboured subalpine derivatives which eventually gave rise to the extant Dasycerus.

Checklist of Staphylinidae (Coleoptera) of El Salvador.

This paper is primarily a compilation of published data on the staphylinid beetle species reported for El Salvador. It is based on taxonomic and ecological literature, with inclusion of some new records from different entomological collections. Thirty eight genera and 96 species are reported in the list, including a first country record: Eulissus chalybaeus. Country-level distribution outside of El Salvador, locality references and topographic zones are included for each species. In the list, three endemic species are recorded: Seeversiella badia, S. minima and Stenus salvadorensis. It is remarkable that 14 genera (37%) and 52 species (54%) have only been reported at a country level in the literature, without a specific locality of collection, indicating the need for local inventories of this coleopteran family in the salvadoran territory.

New names for junior homonyms in Scydmaeninae (Coleoptera, Staphylinidae).

As in most species-rich groups, cases of homonymy are not uncommon among Scydmaeninae. Many primary and secondary junior homonyms were recently replaced by O'Keefe (2000), Davies (2004), Davies Vít (2004), Castellini (2010), Jaloszynski (2011), Asenjo (2016), and Newton (2017). One new primary homonym has been published since then, and changes in generic placement (Jaloszynski 2013, 2020a) have generated several new secondary homonyms. New names for those are proposed below.

Linking evolutionary mode to palaeoclimate change reveals rapid radiations of staphylinoid beetles in low-energy conditions.

Staphylinoidea (Insecta: Coleoptera) is one of the most species-rich groups in animals, but its huge diversity can hardly be explained by the popular hypothesis (co-radiation with angiosperms) that applies to phytophagous beetles. We estimated the evolutionary mode of staphylinoid beetles and investigated the relationship between the evolutionary mode and palaeoclimate change, and thus the factors underlying the current biodiversity pattern of staphylinoid beetles. Our results demonstrate that staphylinoid beetles originated at around the Triassic-Jurassic bound and the current higher level clades underwent rapid evolution (indicated by increased diversification rate and decreased body size disparity) in the Jurassic and in the Cenozoic, both with low-energy climate, and they evolved much slower during the Cretaceous with high-energy climate. Climate factors, especially low O2 and high CO2, promoted the diversification rate and among-clade body size disparification in the Jurassic. In the Cenozoic, however, climate factors had negative associations with diversification rate but little with body size disparification. Our present study does not support the explosion of staphylinoid beetles as a direct outcome of the Cretaceous Terrestrial Revolution (KTR). We suppose that occupying and diversifying in refuge niches associated with litter may elucidate rapid radiations of staphylinoid beetles in low-energy conditions.

Updated catalog and generic keys of the Leiodidae (Insecta: Coleoptera) of the Neotropical region ("Latin America": Mexico, the West Indies, and Central and South America).

This paper presents an updated catalog of all taxa of Leiodidae (s.lat.) reported from the Neotropical Region. Keys are presented for the identification of all subfamilies, tribes, and 62 described genera. Three undescribed genera are included in the keys. A total of 600 valid named species are listed, with type localities, type depositories, synonyms, distributions, and biologies where known, and some unnamed species as recorded in the literature. Many species remain to be described. In this work we formally establish no new synonyms and no new combinations although we may indicate the existence of these; but we add new records for described species, and we make spelling corrections of scientific names, when appropriate. A brief review of distribution patterns is given. The fauna has been derived partly from some Nearctic elements that have penetrated as far south as Bolivia. A few genera in the Neotropical element have penetrated the Nearctic Region as far north as the northern U.S.A. or southeastern Canada. Most of the Neotropical genera are autochthonous. In southern South America there is a diverse Neo-Austral fauna with clear "Transantarctic" relationships to Australia and New Zealand and weakly to southern Africa. Some genera variously occur on other continents.

New Data, Old Story: Molecular Data Illuminate the Tribal Relationships among Rove Beetles of the Subfamily Staphylininae (Coleoptera: Staphylinidae).

The megadiverse subfamily Staphylininae traditionally belonged to the best-defined rove beetle taxa, but the advent of molecular phylogenetics in the last decade has brought turbulent changes to the group's classification. Here, we reevaluate the internal relationships among the tribes of Staphylininae by implementing tree inference methods that suppress common sources of systematic error. In congruence with morphological data, and in contrast to some previous phylogenetic studies, we unambiguously recover Staphylininae and Paederinae as monophyletic in the traditional sense. We show that the recently proposed subfamily Platyprosopinae (Arrowinus and Platyprosopus) is a phylogenetic artefact and reinstate Arrowinus as a member of Arrowinini stat. res. and Platyprosopus as a member of Platyprosopini stat. res. We show that several recent changes to the internal classification of the subfamily are phylogenetically unjustified and systematically unnecessary. We, therefore, reestablish Platyprosopini, Staphylinini, and Xantholinini as tribes within Staphylininae (all stat. res.) and recognize Coomaniini as a tribe (stat. nov.) rather than subfamily. Consequently, the traditional ranks of the subtribes Acylophorina, Afroquediina, Amblyopinina, Antimerina, †Baltognathina, Cyrtoquediina, Erichsoniina, Hyptiomina, Indoquediina, Quediina, and Tanygnathinina are restored (all stat. res.). We review the current classification of Staphylininae and discuss sources of incongruence in multigene phylogenies.

Congruence of morphological and molecular phylogenies of the rove beetle subfamily Staphylininae (Coleoptera: Staphylinidae).

Staphylininae is the third largest subfamily of the enormous family Staphylinidae. Monophyly of Staphylininae and its sister relationship to the subfamily Paederinae have been broadly accepted according to both conventional morphology- and molecular-based phylogenies until the last three years. Recent molecular phylogenies rejected monophyly of Staphylininae and regarded Paederinae as a clade within it. This paper re-evaluates the recent molecular work, aiming to clarify the relationship between Staphylininae and Paederinae and resolve intertribal relationships within Staphylininae. Based on a new six-gene data set (5707 bp) for 92 taxa including Oxyporinae (outgroup), representatives of Paederinae, and members of all extant tribes of Staphylininae from published DNA data in GenBank, we generated a well-resolved phylogeny of Staphylininae with all deep nodes (intertribal relationships) strongly supported, and reassert the hypothesis that Staphylininae is monophyletic and indeed the sister group to Paederinae using both Bayesian and maximum likelihood inference. Additionally, our study is a case-study to show that both outgroup selection and completeness of nucleotide data can influence the outcome of a molecular phylogeny. With an increasing number of staphylinid fossils being discovered, the robust phylogeny of Staphylininae inferred by our research will provide a good framework for understanding the early evolution of this group.

Basal polyphagan beetles in mid-Cretaceous amber from Myanmar: biogeographic implications and long-term morphological stasis.

The origin and early evolutionary history of polyphagan beetles have been largely based on evidence from the derived and diverse 'core Polyphaga', whereas little is known about the species-poor basal polyphagan lineages, which include Scirtoidea (Clambidae, Decliniidae, Eucinetidae, and Scirtidae) and Derodontidae. Here, we report two new species Acalyptomerus thayerae sp. nov. and Sphaerothorax uenoi sp. nov., both belonging to extant genera of Clambidae, from mid-Cretaceous Burmese amber. Acalyptomerus thayerae has a close affinity to A. herbertfranzi, a species currently occurring in Mesoamerica and northern South America. Sphaerothorax uenoi is closely related to extant species of Sphaerothorax, which are usually collected in forests of Nothofagus of Australia, Chile, and New Zealand. The discovery of two Cretaceous species from northern Myanmar indicates that both genera had lengthy evolutionary histories, originated at least by the earliest Cenomanian, and were probably more widespread than at present. Remarkable morphological similarities between fossil and living species suggest that both genera changed little over long periods of geological time. The long-term persistence of similar mesic microhabitats such as leaf litter may account for the 99 Myr morphological stasis in Acalyptomerus and Sphaerothorax. Additionally, the extinct staphylinoid family Ptismidae is proposed as a new synonym of Clambidae, and its only included species Ptisma zasukhae is placed as incertae sedis within Clambidae.

Lepicerus larva still unknown: a correction (Coleoptera: Lepiceridae, Phalacridae).

The larva of Lepicerus inaequalis Motschulsky was described by Lawrence et al. (2013) based on several early instars and one late instar collected among wet leaves and debris near Gamboa, Panama. The identification was based on collection of an adult Lepicerus in a similar habitat nearby and a combination of characters found in other myxophagan immatures but not known in other beetle larvae. We accepted this identification with some reservations, but a misinterpretation of one feature in the original description plus an unexpected new source of evidence makes it likely that our identification was incorrect. The one misinterpreted feature in this larva was the retraction of the ventral mouthparts. Reexamining the larvae, it was found that a pair of tendons extending mesally from the hypostomal rods were misinterpreted as maxillary bases. The mouthparts are definitely protracted, which is expected with long hypostomal rods. The new source of evidence is somewhat complicated, as explained below.

Checklist and taxonomic changes for Central and South American Philonthina (Coleoptera: Staphylinidae).

A checklist of all described species of Philonthina, a subtribe of the staphylinid tribe Staphylinini, known to occur in Central and South America (CASA) is presented. Included for each species, and for synonyms known from CASA, is a reference to the original description, type locality and type depository, and for each species the known distribution within and outside CASA. Type material was sought in the main European and American collections where it is deposited (BMNH, MNHUB, IRSNB and FMNH) and is summarized for all indigenous CASA species, with lectotypes designated for 16 names and confirmation of holotypes and prior designation of lectotypes when necessary. Based on recent phylogenetic work in Philonthina and our revision of types of CASA species of Philonthus Stephens, 1829 and Belonuchus Nordmann, 1837, some taxonomic changes are proposed. Thirty-one species of Philonthus are transferred to Belonuchus (16), Gabrius Stephens 1829 (14), and Bisnius Stephens 1829 (one) resulting in the following new combinations: B. abnormalis (Sharp 1885), B. celatus (Sharp 1885), B. corticalis (Sharp 1885), B. extremus (Sharp 1885), B. infimus (Sharp 1885), B. iteratus (Sharp 1887), B. latecinctus (Sharp 1885), B. lucilius (Sharp 1885), B. muticus (Sharp 1876), B. optatus (Sharp 1885), B. platypterus (Sharp 1885), B. rufiventris (Sharp 1887), B. rufocaudus (Sharp 1885), B. rufopygus (Sharp 1885), B. serraticornis (Sharp 1876), B. supernus (Herman 2001), G. approximans (Sharp 1885), G. armatipes (Sharp 1885), G. atricolor (Sharp 1885), G. championi (Sharp 1885), G. dampfi (Bernhauer 1929), G. elegans (Sharp 1885), G. forsterianus (Scheerpeltz 1960), G. misellus (Sharp 1885), G. nugax (Sharp 1885), G. ovaticeps (Sharp 1885), G. peruvianus (Bernhauer 1916), G. planulatus (Sharp 1885), G. rusticus (Sharp 1885), G. serpens (Sharp 1885) and Bi. subaeneipennis (Bernhauer 1916). Endeius nitidipennis Solier 1849 is transferred to Gabrius, resulting in the following new combination, G. nitidipennis (Solier 1849). Leptopeltus carchiensis Chani-Posse Asenjo 2013 is proposed as junior synonym of Philonthus divisus Sharp 1891, which is transferred to Leptopeltus Bernhauer 1906 resulting in a new combination: Leptopeltus divisus (Sharp 1891). Belonuchus penetrans Silvestri 1946 is transferred to Pridonius Blackwelder 1952 as a new combination. Lectotypes are designated for Atopocentrum mirabile Bernhauer 1906, Philonthus armatipes Sharp 1885, Ph. atricolor Sharp 1885, Ph. championi Sharp 1885, Ph. misellus Sharp 1885, Ph. planulatus Sharp 1885, Ph. rusticus Sharp 1885, Ph. serpens Sharp 1885, Ph. abnormalis Sharp 1885, Ph. celatus Sharp 1885, Ph. infimus Sharp 1885, Ph. latecinctus Sharp 1885, Ph. muticus Sharp 1876, Ph. platypterus Sharp 1885, Ph. rufocaudus Sharp 1885 and Ph. rufopygus Sharp 1885. Of the 543 currently known species of Philonthina reported from CASA, at least 14 are believed to be adventive from elsewhere, 56 may occur naturally elsewhere, and 473 (87%) are evidently endemic to this region. Of the 31 genera represented by these described species, 20 (65%) are endemic to CASA. One genus, Gabronthus Tottenham 1955, is adventive. However, the actual philonthine fauna of CASA will undoubtedly be much larger, and the generic composition highly modified, when the fauna is fully explored and studied within a phylogenetical framework.

Redescription and new records of Ontholestes brasilianus Bernhauer, 1906 (Coleoptera: Staphylinidae: Staphylininae).

The genus Ontholestes Ganglbauer includes 35 species distributed mainly in Eurasia, with a few additional species in Africa and North and South America (Herman, 2001; Yang Zhou, 2012; Smetana Shavrin, 2013; Rougemont, 2016). According to Asenjo et al. (2013), the South American record of the Palearctic species Ontholestes murinus (Linnaeus, 1758) for Brazil made by J. Guérin (1953) seems doubtful. Ontholestes murinus was recorded for the first time outside the Palaearctic region by Smetana (1981), from Newfoundland, in Canada, as an adventive species (e.g., Downie and Arnett, 1996; Brunke et al., 2011), but its occurrence in Brazil remains to be confirmed; if the Guérin (1953) record was based on a mistaken identification or mislabeled specimen, this would reduce the number of species distributed in this region from two to one. With respect to O. brasilianus Bernhauer, although it has been confirmed for Peru, Brazil and Argentina (Herman, 2001; Asenjo et al., 2013; Newton, 2015; Newton Caron, 2015), no specific localities of occurrence have been reported since its description in 1906. Thus, to solve problems of misidentification with Neotropical species of this genus, in this study we redescribe Ontholestes brasilianus and provide the first illustrations of the beetle including its aedeagus and a short key for South American species. Additionally, new records from South America are listed here.

Taxonomy of 'Euconnus complex'. Part XV. Disentangling the Euconnus-Sciacharis knot (Coleoptera, Staphylinidae, Scydmaeninae).

Many species currently placed in Sciacharis Broun were once included in Euconnus Thomson, and their taxonomic history is complicated due to diffuse and overlapping diagnoses of both genera applied by previous authors, numerous virtually unjustified transfers of species between them, and hundreds of species not assigned to any subgenus. Based on comparative morphological studies of respective type species, we clarify the status of Allomaoria Franz, Austroconophron Franz, Magellanoconnus Franz, Neuraphoconnus Franz and Valdivioconnus Franz. All of them, except for Neuraphoconnus, were originally described as subgenera of Euconnus, later transferred to Sciacharis. Austroconophron is removed from synonymy with Sciacharis s. str. and resurrected as subgenus of Euconnus (resulting in Euconnus (Austroconophron Franz, status rest.); Allomaoria is retained as a junior synonym of Sciacharis s. str.; Magellanoconnus is removed from Sciacharis and restituted as genus (resulting in Magellanoconnus Franz, status rest.); Anthicimimus Franz, previously removed from subgenus of Sciacharis and elevated to genus rank, is reduced to subgenus of Magellanoconnus (resulting in Magellanoconnus (Anthicimimus stat. rev.); Neuraphoconnus is removed from synonymy with Magellanoconnus and resurrected as genus (resulting in Neuraphoconnus Franz, status rest.); and Valdivioconnus is transferred from subgenus of Sciacharis to subgenus of Microscydmus Saulcy & Croissandeau (resulting in Microscydmus (Valdivioconnus Franz, status rev.). The only Australian species of Magellanoconnus is transferred to Kangarooconnus Jaloszynski, gen. n. (resulting in Kangarooconnus carinifrons (Franz), comb. n.). A complete checklist of 302 species and subspecies previously or currently placed in Sciacharis (including 102 restituted and 50 new combinations) is given. Three replacement names are proposed for secondary homonyms resulted from new combinations: Euconnus (Austroconophron) caledonensides Newton, nom. n. (for Euconnus caledonensis Franz, 1986, not Euconnus caledonensis Franz, 1979); Magellanoconnus (s. str.) castrianus Jaloszynski & Newton, nom. n. (for Magellanoconnus castrii (Franz, 1967: 636), not Magellanoconnus castrii (Franz, 1967: 614)); and Sciacharis (Sciacharis) bryantides Newton, nom. n. (for Euconnus bryanti Franz, 1975, not Euconnus bryanti Lhoste, 1939). Lectotypes are designated (des. Jaloszynski) for Euconnus tindoui Franz, 1971, Phagonophana lanosa Broun, 1885, and Neuraphoconnus caledonicus Franz, 1971.

Response to "Evidence from amber for the origins of termitophily".

In a recent Current Biology paper [1], we reported the oldest, morphologically specialized, and obligate termitophiles, Cretotrichopsenius burmiticus (Figure 1, left), from mid-Cretaceous Burmese amber, about 99 million years old. Cretotrichopsenius, belonging to the obligately termitophilous rove beetle tribe Trichopseniini, display the protective horseshoe-crab-shaped body typical of many extant termitophiles. However, the termitophilous lifestyle of Cretotrichopsenius is being questioned by Yamamoto et al.[2] based on their representation of the termitophile-related features and premature and presumptive phylogenetic placement of Cretotrichopsenius within Trichopseniini. We stand by our interpretation that Cretotrichopsenius are obligate termitophiles, and Mesosymbion[3], a member of the largely free-living Mesoporini, are not necessarily termitophilous.

Show me your tenent setae and I tell you who you are - Telling the story of a neglected character complex with phylogenetic signals using Leiodidae (Coleoptera) as a case study.

The tarsal setae in 97 species of Leiodidae and eight outgroups were examined using SEM imaging and dissections. Modified adhesive setae present in males are referred to as "male tenent setae" (MTS). In most cases, dilated tarsomeres were associated with MTS, which were always present on the protarsi and sometimes the mesotarsi. MTS are reported for the first time on the mesotarsi of Leptodirini and on the metatarsi in two genera of Sogdini. Contrary to reports in the literature, the reduction in the number of the MTS bearing mesotarsomeres is considered a derived condition. Both sexes of Leptinus (Platypsyllinae) have modified setae (referred to as tenent setae in the literature), probably related to their specialised association with mammals, and a patch of MTS was recognized for the first time among those modified setae among males. Four main types of MTS are recognised: (1) a plesiomorphic discoidal type that has a shaft with a round cross-section and maintains a similar diameter throughout its length until forming the expanded discoidal terminal plate; (2) a minidiscoidal type, similar to discoidal but with a relatively small terminal plate, found in Cholevinae; (3) a conical type, present in Leiodinae (excluding Estadiini) where the shaft increases in diameter until forming the terminal plate; and (4) a spatulate type, where an even wider terminal plate has a lateral projection, derived from the conical form and synapomorphic for the leiodine tribes Pseudoliodini, Scotocryptini, and possibly Agathidiini.

Early Evolution of Specialized Termitophily in Cretaceous Rove Beetles.

Termitophiles, symbionts that live in termite nests, include a wide range of morphologically and behaviorally specialized organisms. Complex adaptive mechanisms permit these animals to integrate into societies and to exploit their controlled physical conditions and plentiful resources, as well as to garner protection inside termite nests. An understanding of the early evolution of termitophily is challenging owing to a scarcity of fossil termitophiles, with all known reliable records occurring from the Miocene (approximately 19 million years ago [mya]) [1-6], and an equivocal termitophile belonging to the largely free-living Mesoporini from the mid-Cretaceous [7]. Here we report the oldest, morphologically specialized, and obligate termitophiles from mid-Cretaceous Burmese amber (99 mya). Cretotrichopsenius burmiticus gen. et sp. nov. belongs to Trichopseniini, a group of distinctive termitophilous aleocharine rove beetles, all of which possess specialized swollen or horseshoe-crab-shaped body plans. Cretotrichopsenius display the protective horseshoe-crab-shaped body form typical of many modern termitophiles, with concealed head and antennae and strong posteriorly directed abdominal setae. Cretotrichopsenius represent the earliest definitive termitophiles, shedding light on host associations in the early evolution of termite societies. The fossil reveals that ancient termite societies were quickly invaded by beetles and by multiple independent lineages of social parasites over the subsequent eons.

A catalog of the types of Staphylinidae (Insecta, Coleoptera) deposited in the Museo Argentino de Ciencias Naturales, Buenos Aires (MACN).

The type specimens (all current categories) of Staphylinidae deposited in this Museum are listed; names are recorded, most of them represented by name-bearing types (primary types). The specific and subspecific names are alphabetically ordered in a single list, followed by the generic names (and subgeneric ones, if they were stated) spelled as they were published; later combinations and/ or current binomina are mentioned insofar these are known to the authors. Two lists are added: 2. Specimens labeled as types of names not found in the literature and probably never published, or published as nomina nuda; and 3. Specimens labeled as types, but not originally designated as such. An appendix provides a systematically arranged list of all names discussed, with indication of where they are discussed in the text.