INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

Fossil Cycadales of Argentina

A survey of Cycadalean taxa of Argentina (including Antarctica) is presented. The record of leaves represented byNilssonia, Pseudoctenis, Ctenis, Mesodescolea, Ticoa, Almargemia,Kurtziana, andZamia genera are summarized. Recent investigations made of cuticles with transmission and scanning electron microscopy are included. In stems, a preliminary study of two forms found in the Upper Cretaceous of Rio Negro Province are incorporated intoMichelilloa, Bororoa, andMenucoa. The fossil record shows some characters of leaf morphology, presence of inverse xylem and medullary bundles, two kinds of leaf traces, and monoxylic and polyxylic steles of systematic importance when compared with both extant and fossil cycads. Affinities of fossil taxa related to extant families are suggested through analysis of the above-mentioned characters. The presence of medullary vascular rings related to the emission of terminal reproductive strobili are recognized inBororoa andMenucoa. This character associated with polyxyly makes it possible to assign these genera to the Zamiaceae-Encephalartoideae sensu Stevenson, widening the paleogeographical distribution of this subfamily.     

EXTINCT TRANSITIONAL FAGACEAE FROM THE OLIGOCENE AND THEIR PHYLOGENETIC IMPLICATIONS

Fruits, catkins, and associated leaves of at least two extinct trigonobalanoid taxa have been discovered at an Oligocene fossil plant locality rich in fagaceous remains. These fossils exhibit a mosaic of fruit and pollen characters found in the two extant subfamilies Castaneoideae and Fagoideae of Fagaceae. Comparison with cladograms based on modern taxa suggests that these extinct taxa were similar to the ancestors of subfamily Fagoideae and may have been intermediate between Fagus and the modern trigonobalanoid genera. Pollen types isolated from the fossil staminate catkins provide unique character states that are transitional between modern pollen types in Fagaceae and are important in understanding the evolution of exine micromorphology within the family. This analysis provides a striking example of the use of character data from fossils to determine character-state adjacency prior to polarization of characters using outgroup comparison. Because of the mosaic nature of their character complexes, these fossils support monophyly in both the family Fagaceae and the subfamily Fagoideae. In addition, the occurrence of trigonobalanoid fossils in the Oligocene of North America has interesting biogeographic implications and provides insights into the nature of North American Fagaceae during the Tertiary.     

ON PENTAMPLEXUSSCHINDEWOLF, 1940 (ANTHOZOA,RUGOSA) AND ITS POSSIBLE RELATIVES AND ANALOGUES

Abstract: Three ampleximorphic taxa are revised and their most important characters are discussed in terms of possible or apparent relationships. Re‐interpretation of its early ontogeny allows the assignment of Pentamplexus Schindewolf, 1940 to the family Polycoeliidae de Fromentel, 1861. Stereolasma variabilis Vojnovsky‐Krieger, 1934 is established as the type species of Vojnovskytes gen. nov. It resembles the family Polycoeliidae in some characters and the Antiphyllidae Ilina, 1970 or the Laccophyllidae Grabau, 1928 in others. Thus, its family status is not established. Revision of the type material of Fasciculophyllum tripus Schindewolf, 1952 allows its inclusion within the new genus Silesamplus, probably related to the family Laccophyllidae Grabau, 1928 . Amplexoid morphology is further shown to be inadequate for the establishment of relationships on the family or subfamily level. Early ontogeny is most important in that respect, but biform vs normal morphology in the tabularium and free vs contratingent development of minor septa must also be considered, where appropriate.     

Evolution of Philopotinae,with a revision and phylogeny of the New World spider fly genus Philopota Wiedemann (Diptera,Acroceridae)

Philopotinae are hunchbacked spider flies represented by 63 fossil and extant species in 15 genera worldwide. Philopota Wiedemann, 1830, is the most species‐rich genus within the subfamily. Here, the evolution of Philopotinae is discussed, and a revision and phylogeny of Philopota based on adult morphology are presented. Nine of the 12 extant Philopotinae genera were included in our analysis, and 22 species were recognized in Philopota, of which 13 are described as new. Seven new synonymies are proposed. The phylogenetic analysis included 33 terminal taxa (22 ingroup and 11 outgroup species) and used 53 morphological characters, resulting in a single most parsimonious tree under equal weights. The monophyly of Philopota is recovered, and the Palaearctic genus Oligoneura is hypothesized as its sister‐group.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

A specimen of Curculioninae (Curculionidae,Coleoptera) from the Lower Cretaceous,Araripe Basin,north‐eastern Brazil

Abstract: A specimen of Curculionidae (Curculioninae) is described as Arariperhinus monnei gen. et sp. nov. The specimen is preserved on a laminated limestone sample of the Crato Formation (Santana Group), Lower Cretaceous (Aptian–Albian), and was collected from a quarry near Nova Olinda, Chapada do Araripe, State of Ceará, Brazil. The genus is placed in the subfamily Curculioninae because of its strongly convex body and relatively slender rostrum, but mainly by its rounded eyes and lack of a prosternal sulcus and tibial spurs. The very prominent eyes in lateral view, a cylindrical rostrum and a straight posterior margin of ventrite II are strong indications that this fossil belongs to the tribe Anthonomini. However, the claws, which would resolve the exact placement of this fossil, are poorly preserved. Arariperhinus monnei gen. et sp. nov. is distinguishable by the combination of several characters and the first record of the family Curculionidae in the Santana Group; it is the oldest record of a member of the subfamily Curculioninae.     

Phylogeny of the bee family Megachilidae (Hymenoptera: Apoidea) based on adult morphology

Phylogenetic relationships within the bee family Megachilidae are poorly understood. The monophyly of the subfamily Fideliinae is questionable, the relationships among the tribes and subtribes in the subfamily Megachilinae are unknown, and some extant genera cannot be placed with certainty at the tribal level. Using a cladistic analysis of adult external morphological characters, we explore the relationships of the eight tribes and two subtribes currently recognised in Megachilidae. Our dataset included 80% of the extant generic‐level diversity, representatives of all fossil taxa, and was analysed using parsimony. We employed 200 characters and selected 7 outgroups and 72 ingroup species of 60 genera, plus 7 species of 4 extinct genera from Baltic amber. Our analysis shows that Fideliinae and the tribes Anthidiini and Osmiini of Megachilinae are paraphyletic; it supports the monophyly of Megachilinae, including the extinct taxa, and the sister group relationship of Lithurgini to the remaining megachilines. The Sub‐Saharan genus Aspidosmia, a rare group with a mixture of osmiine and anthidiine features, is herein removed from Anthidiini and placed in its own tribe, Aspidosmiini, new tribe . Protolithurgini is the sister of Lithurgini, both placed herein in the subfamily Lithurginae; the other extinct taxa, Glyptapina and Ctenoplectrellina, are more basally related among Megachilinae than Osmiini, near Aspidosmia, and are herein treated at the tribal level. Noteriades, a genus presently in the Osmiini, is herein transferred to the Megachilini. Thus, we recognise four subfamilies (Fideliinae, Pararhophitinae, Lithurginae and Megachilinae) and nine tribes in Megachilidae. We briefly discuss the evolutionary history and biogeography of the family, present alternative classifications, and provide a revised key to the extant tribes of Megachilinae.     

Early Cretaceous origin of pollen‐feeding beetles (Insecta: Coleoptera: Oedemeridae)

The taxonomic position of a new pollen‐feeding fossil beetle from Spanish amber (late Albian, 105 Ma) is analysed. A phylogenetic analysis allows me to accommodate Darwinylus marcosi gen. et sp. nov. in the Polyphaga: Oedemeridae within current limits for the family, which clearly belongs in the subfamily Oedemerinae. It corresponds to the oldest definitive record for the family. Some autapomorphies, mainly in antennae, are observable in the fossil compared with extant members of the family. A discussion about these problematic characters and the evolution of the family is proposed.     

Phylogeny of the tribe Abrotrichini (Cricetidae,Sigmodontinae): integrating morphological and molecular evidence into a new classification

The tribe Abrotrichini (five genera and 14 living species) is a small clade within the speciose subfamily Sigmodontinae (Rodentia, Cricetidae), representing one of the extant successful radiations of mammals at southern high latitudes of the Neotropics. Its distribution is mostly Andean, reaching its greatest diversity in southern Argentina and Chile. We evaluate the phylogenetic relationships within this tribe through parsimony and Bayesian approaches based on 99 morphological characters (including 19 integumental characters, 38 skull characters, 31 dental characters, three postcranial skeletal characters, seven from the male accessory glands and phallus and one from the digestive system) and six molecular markers (one mitochondrial and five nuclear). We include representatives of all, except one, of the currently recognized species of living Abrotrichini plus one fossil form. Based on total evidence, we recovered a primary division between the genus Abrothrix and a group including the long‐clawed Abrotrichini, Chelemys, Geoxus, Notiomys and Pearsonomys. Both clades are recognized and named here as subtribes. The large degree of morphological variation observed within Abrothrix suggests that species in the genus fall into four groups, which we recognize as subgenera. In addition, the two known species of Chelemys do not form a monophyletic group, and Geoxus was recovered as paraphyletic with respect to Pearsonomys. To reconcile classification and phylogenetics, we describe a new genus for Chelemys macronyx and include Pearsonomys as a junior synonym of Geoxus. Our results highlight the importance of both morphology and molecules in resolving the phylogenetic relationships within this tribe. Based on biogeographical analyses, we hypothesize that Abrotrichini originated in south‐western South America by vicariance and then diversified mostly by successive dispersal events.     

金缕梅科的种子形态学及其系统学评价

本文利用光学显微镜及扫描电镜在属级水平上观察了金缕梅科全部6亚科中的21属41种1变种植物的种子特征,结果表明：该科的种子形状、颜色、大小及种脐特征在各亚科中有差异(除了双花木亚科与金缕梅亚科的大多数属是一致的)可作为区分亚科的特征之一。根据种皮的特征,可划分为5类：(1)种皮平滑或仅在边缘部分具条纹状纹饰(双花木亚科及金缕梅亚科中大部分属);(2)种皮具条纹状纹饰,条纹有波浪加厚或升高(马蹄荷亚科);(3)种皮具瘤状纹饰或具条纹和瘤状相间排列的纹饰(红花荷亚科);(4)种皮具条一网状纹饰(壳菜果亚科);(5)种皮具网状纹饰(枫香亚科)。种皮纹饰对亚科的划分具有一定意义。此外,还详细观察了该科的种皮结构,确认金缕梅科的种皮由外种皮及内种皮构成,而外种皮又进一步划分为外层、中层和内层,种皮各层的细胞层数对该科的分类价值不大。研究结果还表明：枫香亚科的种子特征明显不同于该科的其它成员。关于枫香亚科的系统位置还需深入研究。     

Review of the Late Eocene Encyrtidae (Hymenoptera,Chalcidoidea) with a description of the first fossil genus with filum spinosum

Glaesus gibsoni gen. et sp. nov. and Eocencnemus gedanicus sp. nov. are described from materials stored in the amber collection of the University of Gdańsk. Eocencyrtus zerovae Simutnik, 2001 is for the first time recorded in the Baltic amber. The encyrtids recorded in Eocene ambers are reviewed, and a key to the known genera is given. The fossil Encyrtidae cannot be placed with confidence in any of the established subfamilies. Their comparative morphology is analyzed. One of the diagnostic characters of the subfamily Encyrtinae, the presence of the filum spinosum, is recorded for the first time for fossil encyrtids (in G. gibsoni).     

Pollen morphology of the Thymelaeaceae in relation to its taxonomy

 Pollen morphology of the four subfamilies of Thymelaeaceae sensu Domke (1934) was examined using LM, SEM and TEM. The variation of the exine architecture allows to distinguish four pollen types and three subtypes. Distribution of pollen types proves to be widely correlated with the occurrence of characters from wood anatomy and flower morphology. If pollen types are connected with subfamilies sensu Domke (1934), Gonystyloideae, Thymelaeoideae and Synandrodaphnoideae are stenopalynous, whereas Aquilarioideae are eurypalynous. Based on pollen morphology as well as on other characters it is evident that Aquilarioideae are not monophyletic. Its genera Aquilaria and Gyrinops are more closely related to Thymelaeoideae and Synandrodaphnoideae than to other genera of Aquilarioideae sensu Domke (1934). The remaining genera of Aquilarioideae Deltaria, Solmsia and Lethedon are most similar to Gonystyloideae and therefore are included in this subfamily, as it was suggested by Airy Shaw (1979). Furthermore, palynological and other characters favour the transfer of Octolepis from Aquilarioideae to Gonystyloideae. The re-circumscribed Aquilarioideae together with Thymelaeoideae and Synandrodaphnoideae are shown to form a monophyletic group. Received August 8, 2001 Accepted December 7, 2001     

CHARACTER DIAGNOSIS, FOSSILS AND THE ORIGIN OF TETRAPODS

I. The traditional view of the origin of tetrapod vertebrates is that they are descendants of fossil osteolepiform fish, of which Eusthenopteron is best known. In recent years both that conclusion and the methodology by which it has been reached have been challenged by practitioners of cladistic analysis. Particularly a recent review by Rosen et al. (1981) claims that Dipnoi (lungfish) are the sister-group of the Tetrapoda, that Osteolepiformes is a non-taxon and that Eusthenopteron is more distant from tetrapods than are Dipnoi, coelacanths and probably the fossil Porolepiformes. We attempt to refute all these concludions by use of the same cladistic technique. 2. We accept that all the above-mentioned groups, together with some less well-known taxa, can be united as Sarcopterygii by means of shared derived (apomorph) characters. We also agree that Porolepiformes and Actinistia (coelacanths) can be characterized as valid taxa. The primitive and enigmatic fossil fish Powichthys is accepted as representing the plesiomorph sister-group of true porolepiforms. 3. Only two apomorph features, the course of the jaw adductor muscles and the position of incurrent and excurrent nostrils, appear to unite all the fish, living and fossil, currently regarded as Dipnoi. The characteristic tooth plates and the presence of petrodentine both exclude important primitive fossil forms. 4. Contrary to the opinion of Rosen et al., Osteolepiformes can be characterized — by the arrangement of bones forming the cheek plate, the presence of basal scutes to the fins and by the unjointed radials of the median fins. However, if these are true autapomorphies they exclude any osteolepiform from direct tetrapod ancestry. 5. Tetrapoda is a monophyletic group characterized by ten or more autapomorphies, including the bones of the cheek plate, a stapes and fenestra ovalis, and a series of characters of the appendicular skeleton. 6. Tetrapods have a true choana (internal nostril). We accept that the posterior (excurrent) nostril of Dipnoi is the homologue of the tetrapod choana. However, we assert that the posterior nostril of all bony fish is the homologue of the choana. This assertion would be refuted if any fish showed separate posterior nostril and choana. We reject the claim that this ‘three nostril condition’ occurred in porolepiforms and osteolepiforms. The evidence for a choana in porolepiforms is inadequate. Osteolepiforms had a true choana, characterized as in tetrapods by its relationship to the bones of the palate, but no third nostril. Dipnoans are not choanate. 7. Following cladistic practice, the relationship of the extant taxa is established first. Dipnoi are thus shown to be the living sister-group of tetrapods, but only on ‘soft anatomy’ characters unavailable in fossils. Coelacanths are the living sister-group of the taxon so formed. 8. The relationship of the fossil taxa to the extant sarcopterygians is then considered. The synapomorphy scheme proposed by Rosen et al. is discussed at length. Virtually all the characters they use to exclude close relationship of Eusthenopteron (and hence all osteolepiforms) to tetrapods, in favour of coelacanths and dipnoans, are invalid. 9. A series of synapomorphies uniting osteolepiforms and tetrapods is proposed, including a true choana (hence the taxon Choanata), the histology of the teeth, and a number of characters of the humerus. The recently discovered fossil Youngolepis, which lacks a choana, represents the sister-group of the Choanata, and is not uniquely close to Powichthys. The latter, as a porolepiform (s.l.) is a member of the sister-group to Choanata plus Youngolepis. 10. Our cladistic analysis suggests that all the extinct taxa considered are more closely related to tetrapods than are the Dipnoi. Moreover fossil evidence suggests that Dipnoi, considered as an extant taxon, may not even be the living sister-group of Tetrapoda. Early fossil dipnoans appear to have been marine fish without specific adaptations for air breathing. If so the apparent synapomorphies of Dipnoi and Tetrapoda may be homoplastic — the insistence on grouping extant taxa first would then have yielded an invalid inference.     

POLLEN MORPHOLOGY AND PLANT TAXONOMY OF WHITE OAKS IN EASTERN NORTH AMERICA

An evaluation of possible approaches to fossil oak pollen identification utilized scanning electron microscopy to examine exine-surface features of 171 collections, representing 16 Quercus subgenus Lepidobalanus species and varieties of eastern North America. Twenty qualitative pollen morphological characters were defined and tabulated for each of 217 pollen grains. The data were subjected to cluster analysis and cluster diagrams were compared with published white oak taxonomy. Pollen morphology and plant taxonomy compared well in series of the subgenus Lepidobalanus due primarily to consistency of character presence and absence within species and varieties. Pollen morphology of white oaks appears to reflect plant systematics above the species level. Use of routine SEM analysis to identify series of white oaks among fossil pollen grains likely will yield valid results.     

Towards a phylogeny of the flesh flies (Diptera: Sarcophagidae): morphology and phylogenetic implications of the acrophallus in the subfamily Sarcophaginae

The morphology of the acrophallus, the distal portion of the male phallus carrying the phallotreme, was studied in 72 exemplar species representing 56 genera and subgenera of the flesh fly subfamily Sarcophaginae. For 42 of those species, scanning electron microscopy was used to clarify the phallic morphology. Terms used to describe the male genitalia were updated based on new interpretations of homology. Male genitalic characters, combined with other morphological characters of adult males and females and of larvae, were used to construct a phylogeny. The monophyly of the subfamily was supported, and some generic‐level sister‐group relationships proposed in the literature, but without previous cladistic analyses, were also supported. The genus Blaesoxipha Loew, as currently recognized, was not monophyletic in our analysis. The genus Helicobia Coquillett is synonymized with Sarcophaga Meigen syn. nov. and treated as a subgenus of the latter. The Sarcophaga subgenera Neobellieria Blanchard and Mehria Enderlein were not monophyletic. Many of the clades in the analysis were supported primarily or exclusively by male genitalic character states, highlighting the importance of the male genitalia as a source of morphological characters for sarcophagine phylogeny. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 740–778.     

Australia's oldest Anseriform fossil: a quadrate from the Early Eocene Tingamarra Fauna

Abstract: A partial quadrate (essentially the otic part) from the nonmarine, earliest Eocene (54.6 Ma) Tingamarra Local Fauna in Queensland, Australia, has been identified as the oldest Australian anseriform fossil. The Tingamarra quadrate shows a combination of plesiomorphic anseriform characters with a unique synapomorphic character complex of the Anhimidae (screamers), which today are endemic to South America. In concert with the basal position of the Anhimidae among the crown‐group anseriforms, this set of characters suggests a stem group of the Anhimidae, raising a possibility of the Transantarctic migration of stem anhimids to South America. The quadrate morphology supports palaeognathous rather than recently claimed anhimid relationships of the Dromornithidae and identifies Sylviornis as an anseriform rather than a galliform.     

On the phylogenetic position and systematics of extant and fossil Aclopinae (Coleoptera: Scarabaeidae)

The Aclopinae is a small subfamily within the family Scarabaeidae. It currently comprises five extant genera with 28 species, and eight fossil genera with 25 species. The systematic position of Aclopinae within the family Scarabaeidae is uncertain, particularly because representative species of Aclopinae have been absent in previous phylogenetic studies. Here we performed phylogenetic analyses using morphological and molecular data to investigate the phylogenetic position of fossil and extant Aclopinae. For this objective, we expanded and revised a former morphological data matrix (composed of 68 characters) including all extant genera of Aclopinae. We complemented our morphological investigations with a molecular phylogenetic analysis based on four genes of several extant taxa of Aclopinae and a wide sample of diverse Scarabaeoidea. Our phylogenetic analyses show that all the type species of the fossil genera formerly included within Aclopinae do not belong within the extant Aclopinae clade and support both the exclusion of those fossil taxa and the monophyly of the extant genera of Aclopinae: Aclopus Erichson, Desertaclopus Ocampo & Mondaca, Gracilaclopus Ocampo & Mondaca, Neophanaeognatha Allsopp and Phanaeognatha Hope. Our results also show that the fossil taxa Prophaenognatha robusta Bai et al. and Ceafornotensis archratiras Woolley are closely related to Ochodaeidae, while the remaining type species of fossils formerly included in Aclopinae (Cretaclopus longipes (Ponomarenko), Holcorobeus vittatus Nikritin, Juraclopus rodhendorfi Nikolajev, Mesaclopus mongolicus (Nikolajev), and Mongolrobeus zherikhini Nikolajev) belong to a distinct lineage closely related to Diphyllostomatidae. Based on these results, the subfamily Aclopinae appears monophyletic and sister to the ‘pleurostict’ lineage. Consequently, we propose the following changes to the current classification of the fossil taxa: Holcorobeus monreali (Gómez‐Pallerola) belongs to Carabidae (incertae sedis) as proposed by the original author, and we place Ceafornotensis Woolley, Cretaclopus Nikolajev, Holcorobeus Nikritin, Juraclopus Nikolajev, Mesaclopus Nikolajev, Mongolrobeus Nikolajev and Prophaenognatha Bai et al. in Scarabaeoidea (incertae sedis). Furthermore, we provide an identification key to, and diagnoses of, the genera, illustrations of diagnostic characters and checklists of their included species. The evolutionary perspective presented provides new insights into the evolution of the pleurostict condition in Scarabaeoidea and the biogeography of this group, which is now regarded as Gondwanan, probably evolving during the Cretaceous and not from the upper Jurassic as previously assumed.