New species,new genera,and new records in Cerambycidae from Peru (Coleoptera)

Three new species and two new genera of longhorn beetles are described from Peru: Cosmoplatus polis n. sp. (Pteroplatini), Hemilocallia grishami n. gen., n. sp. (Calliini) and Arabela ramirezi n. gen., n. sp. (Hemilophini). Esmeralda laetifica Bates, 1869 (Prioninae), Stratone transversalis (Chevrolat, 1862) (Cerambycinae), Gymnocerina cratosomoides (Bates, 1862) (Lamiinae), and Trachysomus verrucosus (Olivier, 1795) (Lamiinae) are new records for Peru.     

The oldest Cenozoic passandrid (Coleoptera: Cucujoidea: Passandridae) from the Paleocene of Menat (France)

The new oldest representative of the subfamily Passandrinae (Passandridae) was found in materials from Menat (Puy-de-Dôme, France). It was preliminarily assigned to the genus Passandra Dalman in Schönherr, 1817, although the new species (?) Passandra plenaria sp. n., in contrast to other congeners, demonstrates an exposed labrum and an absence of preapical groove or impression on the hypopygidum. Some notes on fossil records and classification of the family Passandridae are made. The distinctiveness of the families Passandridae and Laemophloeidae in light of the new data on fossils is considered, and the attribution of Mesopassandra Jin, Ślipiński, Zhou and Pang, 2019 (type genus of Mesopassandrinae) to the family Passandridae is regarded as problematic.     

Molecules,morphology and minute hooded beetles: a phylogenetic study with implications for the evolution and classification of Corylophidae (Coleoptera: Cucujoidea)

Eight genes (nuclear: 18S, 28S, H3, CAD; mitochondrial: 12S, 16S, COI, COII) and morphology were used to infer the evolutionary history of Corylophidae, some of the smallest free‐living insects. The study included 36 corylophid exemplars, representing approximately 60% of the known generic diversity of the family and 16 cucujoid outgroup taxa. Multiple partitioning strategies, molecular datasets, combined datasets and different taxon sampling regimes using maximum likelihood and mixed‐model Bayesian inference were utilized to analyse these data. Most results were highly concordant across analyses. There was strong agreement across (i) partitioning strategies, (ii) maximum likelihood and Bayesian inference analyses of the molecular data, and (iii) Bayesian inference of the molecular data alone and Bayesian inference of the combined morphological and molecular data when all terminal taxa were included. When a strict taxon sampling protocol was employed so that only single generic exemplars were included, deep relationships were affected in the resulting phylogenetic hypotheses. Under such narrow sampling strategies, deep phylogenetic relationships were also sensitive to the choice of generic exemplars. Although it is often challenging to obtain single representatives for many taxa in higher‐level phylogenetic analyses, these results indicate the importance of using denser taxon sampling approaches even at the specific level for genera included in such studies. Molecular data alone support Anamorphinae (Endomychidae) strongly as the sister group of Corylophidae. In combined data analyses, Coccinellidae is recovered as the sister group to Corylophidae. In all analyses, Corylophidae and the subfamily Corylophinae are recovered as monophyletic. The monophyly of Periptyctinae was untested, as only a single species was included. All included corylophine tribes were recovered as monophyletic with the exception of Aenigmaticini; Aenigmaticum Matthews forms the sister group to Orthoperus Stephens and Stanus?lipiński et al. is recovered as the sister group of Sericoderus Stephens. Stanus tasmanicus?lipiński et al. is transferred to a new genus, Pseudostanus Robertson, ?lipiński & McHugh gen.n. incertae sedis. We propose a new tribe, Stanini Robertson, ?lipiński & McHugh trib.n. for Stanus bowesteadi?lipiński et al. and a new concept of Aenigmaticini sensu.n. to include only the nominate genus. Anatomical transitions associated with corylophid miniaturization are highlighted. Key phenotypic modifications and elevated rates of substitution in nuclear rRNA genes are evident in a subgroup of Corylophinae that includes the most diminutive species. Other taxonomic and evolutionary implications are discussed in light of the results.     

Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae

Abstract Phylogenetic relationships within the family Corylophidae were investigated. Twenty ingroup taxa and six outgroups were included in a cladistic analysis, based on 48 characters derived from adult and larval morphology. Phylogenetic analysis confirms that Corylophidae are monophyletic within the superfamily Cucujoidea and may be subdivided into two subfamilies: the Australian Periptycinae and the cosmopolitan Corylophinae containing 10 tribes: Foadiini trib.n. , Cleidostethini, Aenigmaticini, Parmulini, Sericoderini, Peltinodini, Orthoperini, Corylophini, Teplinini and Rypobiini. All currently recognized family‐group taxa are thoroughly diagnosed, and keys to their identification based on adults and larvae are provided. Two new genera and three species are described: Weirus gen.n ., containing only W. tozer sp . n . (Australia: Queensland), and Stanus gen.n. , with the two species S. bowesteadi sp.n . (New Zealand) and S. tasmanicus sp.n. (Tasmania). The larvae of Pakalukodes bimaculatus?lipiński et al. from Queensland and of Stanus bowesteadi sp.n. from New Zealand are described and illustrated for the first time.     

New genera and new species of leaf beetles (Coleoptera: Polyphaga: Chrysomelidae) from Lowermost Eocene French amber

Three new monotypical genera of leaf beetles are described: Aoriopsis eocenicus n. gen, n. sp. and Acolaspoides longipes n. gen., n. sp. from the subfamily Eumolpinae, and also Crepidocnema yantarica n. gen., n. sp. from the subfamily Alticinae. Short reviews of known fossil records of the mentioned subfamilies are given and position of the new genera discussed.     

New Genera and Species of ‘Giant’ Ostracods (Crustacea: Cyprididae) from Australia

Two new genera of giant ostracods Lacrimicypris n. gen. and Repandocypris n. gen. from Australia are described and eight new species:L. kumbar n. sp., R. austinensis n. sp., R. gleneagles n. sp., Mytilocypris coolcalalaya n. sp., Australocypris bennetti n. sp., A. beaumonti n. sp., A. mongerensis n. sp. and Caboncypris kondininensis n. sp. The number of known Australian genera and species of giant ostracods are now 6 and 21, respectively. Keys to genera and species are provided: all species can be distinguished using the hemipenis and male first leg. The usefulness of the bursa copulatrix as a species-level taxonomic character is highlighted. Most of the species described in this paper occur in salt lakes and existing ecological information on Western Australian species, as well as the distributions of all species, are summarized. Western Australia has a particularly rich halobiont fauna but current explanations relating richness of the halobiont fauna to the widespread occurrence of salt lakes appear incomplete. Australocypris bennetti is unusual in that it is frequently found at pH < 4. Its morphology differs slightly in acidic and alkaline waters.     

Report on the Polycystididae (Rhabdocoela, Kalyptorhynchia) from Australia, with the Description of Twelve New Species and Six New Genera

Twelve new species of Polycystididae Graff, 1905 are described from the Australian east coast. Nine of them could not be placed in any of the existing genera, and therefore six new genera are erected. Alchoides n. gen. differs from all other polycystidid genera by the presence of a bundle of glands that opens into the distal part of the male atrium (accessory vesicle type V). Two species are included in the genus, A. alchoides n. sp. and A. dittmannae n. sp., which differ from each other in the shape of the stylet. Ametochus gehrkei n. gen n. sp. can be distinguished from other polycystidids by the presence of a prostate vesicle type III associated with a prostate stylet type III, combined with the presence of an accessory stylet type III, while a prostate stylet type II is lacking. Typical for Arrawarria inexpectata n. gen n. sp. is the combined presence of an armed cirrus with a prostate vesicle type II connected to a prostate stylet type II. Two species are assigned to the new genus Duplexostylus n. gen.: D. rowei n. sp. and D. winsori n. sp. These two species have two unique features: a prostate vesicle type IV directly connected to a prostate stylet type III and an asymmetrical septum that surrounds the proximal part of the male atrium. They can be distinguished from each other by differences in the detailed construction of their stylets. Two new species are placed in the new genus Stradorhynchus n. gen.: S. caecus n. sp. and S. terminalis n. sp. They have unpaired gonads and a prostate vesicle type IV connected to a prostate stylet type IV, which combination of characters makes them unique within the Polycystididae. The two species differ from each other in the shape and size of the stylet. Triaustrorhynchus armatus n. gen., n. sp. is the first species of the Polycystididae to have three types of stylet in the male atrium: a prostate stylet type II, a prostate stylet type III and an accessory stylet type III. Three new species could be placed in existing genera: Cincturorhynchus monaculeus n. sp., Paraustrorhynchus caligatus n. sp. and Polycystis australis n. sp. All three differ from their congeners in the form and dimensions of their stylets. All newly described species are discussed, giving their similarities and differences with resembling species. Austrorhynchus hawaiiensis Karling, 1977 is reported for the first time in Australia, and the Australian population is compared with populations from other regions. The presence of Gyratrix hermaphroditus Ehrenberg, 1831, one new species of Paulodora Marcus, 1948 and a new genus of Typhlopolycystidinae Evdonin, 1977 is mentioned.     

Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera

Phytophagous ladybird beetles of the tribe Epilachnini are a cosmopolitan, species‐rich group of significant economic importance as pests of agricultural crops. The tribe is well characterized morphologically and clearly monophyletic, but very little is known about its internal phylogenetic relationships and their genus‐level taxonomy. In order to infer the evolutionary history of Epilachnini, test its monophyly and provide a phylogeny‐based classification, we assembled a comprehensive dataset, consisting of four DNA markers (18S and 28S rRNA and 16S, COI mtDNA) and a matrix of 104 morphological characters for 153 species of Epilachnini representing all previously recognised genera, ～11% of the known species, and 14 outgroup taxa. Molecular, morphological and combined datasets were analysed using maximum likelihood, parsimony and Bayesian inference. Bayes factors and Approximately Unbiased tests (AU) were used to compare alternative phylogenetic hypotheses of unconstrained and backbone‐constrained analysis. Only 14 of the 25 included genera were recovered monophyletic, as originally defined. Afidentula Kapur, Afidenta Dieke, Afissula Kapur, Epilachna Chevrolat, Henosepilachna Li Toxotoma Weise and Mada Mulsant are shown to be poly‐ or paraphyletic; Chnootriba Chevrolat, Subafissa Bielawski, Lalokia Szawaryn & Tomaszewska and Papuaepilachna Szawaryn & Tomaszewska form monophyletic groups within larger clades of genus level. All of these genera are redefined here. The two largest genera of Epilachnini, Epilachna Chevrolat and Henosepilachna Li were represented by multiple monophyletic clades, which we described as new genera: Chazeauiana Tomaszewska & Szawaryn gen.n. ; Diekeana Tomaszewska & Szawaryn gen.n .; Fuerschia Tomaszewska & Szawaryn gen.n. and Ryszardia Tomaszewska & Szawaryn gen.n . The following new synonyms are proposed: Afissa Dieke (=Afissula Kapur); Henosepilachna Li in Li & Cook (=Subafissa Bielawski); Papuaepilachna Szawaryn & Tomaszewska (=Lalokia Szawaryn & Tomaszewska). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:440E7FA4‐C859‐47E0‐8335‐30D478CBA8FA .     

Discheramocephalini,a new pantropical tribe of featherwing beetles (Coleoptera: Ptiliidae): description of new taxa and phylogenetic analysis

Abstract Two new genera and eight new species of featherwing beetles (Coleoptera: Ptiliidae) possessing a remarkable horizontal perforation of the mesoventral keel are described: Skidmorella vietnamensis sp.n. (Vietnam), S. memorabilis sp.n. (Indonesia), S. serrata sp.n. (Vietnam), Fenestellidium capensis gen. et sp.n. (South Africa, type species), F. kakamegaensis sp.n. (Kenya), Cissidium okuensis sp.n. (Cameroon), Dacrysoma usambarensis gen. et sp.n. (Tanzania, type species) and D. felis sp.n. (Madagascar). A phylogenetic analysis of 24 taxa and 37 parsimony‐informative characters supports the hypothesis of a single origin of the mesoventral perforation, thus uniting Discheramocephalus, Skidmorella, Africoptilium, Fenestellidium, Cissidium and Dacrysoma into a pantropically distributed clade, for which a new tribe Discheramocephalini (type genus Discheramocephalus) is proposed. Identification keys to Discheramocephalini genera and, in some cases, to species are provided. Each new species is illustrated with scanning electron microscopy images.     

New Genera and Higher Taxa of Septate Gregarines (Protozoa,Apicomplexa)*

SYNOPSIS The following new taxa of septate gregarines are established: METAMERIDAE fam. nov. includes genera Metamera Duke, Deutomera Bhatia & Setna, and Cognettia gen. n., the last with the type and only species C. legeri (Cognetti de Martiis); family GREGARINIDAE Labbé includes Degiustia gen. n. with the type and only species D. hyallela (Batten & DeGiusti); Cirrigregarina gen. n., with the type species C. spissa (Henry) and another species, C. kamenote (Hoshide); and Molluskocystis gen. n., with the type and only species M. pterotracheae (Stuart). Family HIRMOCYSTIDAE (Grassé) includes Arachnocystis gen. n., with the type species A. arachnoidea (Devdhar & Gourishankar) and 3 other species A. nitida (Geus), A. oribatidae (Geus), A. scutovertexi (Erhardová). Family ACTINOCEPHALIDAE (Léger), subfamily Actinocephalinae (Léger), includes Chilogregarina gen. n., with the type species C. striata (Léger & Duboscq) and 3 other species C. brasiliensis (Pinto), C. dujardini (Schneider), and C. stella (Léger).     

New species and new data on Protrinemuridae and Nicoletiidae (Zygentoma) from Eastern Asia and Pacific islands

Four new genera and eleven new species of Zygentoma thysanurans (families Protrinemuridae and Nicoletiidae) are described and some faunistic novelties reported from Oriental and Australian Regions, viz.: Protrinemura leclerci n. sp., from northern Thailand, Protrinemurella allacrotelsoides n. gen. n. sp., from southern Thailand, and Protrinemuroides celebensis n. gen. n. sp., from the Celebes islands (Protrinemuridae), Lepidospora (L.) digitata n. sp., from northern Thailand, L. (L.) deharvengi n. sp., from the Celebes, and Pseudobrinckina anempodiata n. gen. n. sp., from northern Thailand (Nicoletiidae: Coletiniinae), Gastrotheus (G.) papuanus n. sp., from Papua-New Guinea (Nicoletiidae: Atelurinae), and Metrinura celebensis n. sp., from the Celebes, Trinemurodes anomalocoxa n. sp., from southern Thailand, T. bedosae n. sp., from northern Thailand, and Allotrinemurodes thai n.gen. n. sp., from northern Thailand (Nicoletiidae: Subnicoletiinae). Bharatatelura malabarica Mendes is reported for the first time off the Indian sub-continent (in Suva). Proatelura jacobsoni Silvestri is recorded in Macao (southern China) and in the Moluccas islands and notes are presented on its male sex. Gastrotheus (Lasiotheus) nanus (Escherich) is found for the very first time in Macao, in Cook islands and in Niue. Identification keys are provided to Protrinemuridae genera and to species of Trinemurodes, and modifications are suggested to previously presented keys to Nicoletiidae genera and to Lepidospora and Metrinura species.     

Ski trail effects on a beetle (Coleoptera: Carabidae, Elateridae) community in Vermont

Beetle diversity and species composition were studied using pitfall traps placed along three parallel transects that extended from the center of a downhill ski trail, 100 m into a large, unfragmented spruce-fir forest on Mount Mansfield State Park, VT, USA. Groups of beetles showed associations with three distinct habitats: ski trail openings (Agonum cupripenne, Amara spp., Poecilus lucublandus, Hypnoidus bicolor, and H. abbreviatus), edge (Pterostichus coracinus, Agonum retractum, Sphaeroderus canadensis, and Calathus ingratus), and forest (Pterostichus adstrictus, P. punctatissimus, P. brevicornis, Platynus decentis, Trechas apicalis, T. crassiscapus, and Eanus maculipennis). Consequently, carabid and elaterid species composition in the ski trail, edge, and 5 m into the unfragmented forest showed nearly complete turnover. Most beetles within the ski trail were colonizers from lower elevations; however, one species, H. bicolor, was a tundra species that appeared to have colonized lower elevations along the ski trail opening. Of "forest" beetles, only 2 of 540 individuals (< 0.5%) were collected in the ski trail opening. Thus, ski trails are strong barriers to dispersal for forest beetles, several of which are flightless or dimorphic and primarily short-winged. Consequently, fragmentation of mountain slopes by ski trails, especially those that run perpendicular to the climatic gradient, may put isolated populations at a greater risk of local extirpation.     

A review of the morphologically diverse leafhopper subfamily Stegelytrinae (Hemiptera: Cicadellidae) with description of new taxa

Abstract. The morphologically diverse leafhopper subfamily Stegelytrinae is reviewed and its biology and biogeography are discussed as well as its classification, monophyly and relationship to Deltocephalinae s.l. In addition, the first stegelytrine is recorded east of Wallace's and Weber's lines, extending the distribution of the subfamily outside the Oriental and Palaearctic regions. A checklist and key to the genera of the subfamily are given. The following new genera and species are described: Honguchia rubrofascia Wei & Webb, gen.n & sp.n. from Sabah; H. superba Wei & Webb, sp.n. from Sumatra; Yaontogonia flavopicta Wei & Webb, gen.n & sp.n. from Brunei and Sabah; Y. darjeelingensis Wei & Webb, sp. n. from West Bengal; Sychentia breviata Wei & Webb, gen.n & sp.n. from Malaysia; Louangana stellata Wei & Webb, gen.n & sp.n. from Laos; Shangonia sarawakensis Wei & Webb, gen.n & sp.n. from Sarawak; Quiontugia fuscomaculata Wei & Zhang, gen.n. & sp.n. from Hainan; and Neophansia wallacei Wei & Webb, gen.n & sp.n. from New Guinea.     

Minute moss beetles in the Southern Hemisphere: Molecular phylogeny,historical biogeography and habitat shifts (Coleoptera: Hydraenidae)

Minute moss beetles (Hydraenidae) are one of the most speciose and widespread families of aquatic Coleoptera, with an estimated 4000 extant species, found in the majority of aquatic habitats from coastal rock pools to mountain streams and from the Arctic Circle to the Antarctic islands. Molecular phylogenetic works have improved our understanding of the evolutionary history of the megadiverse Hydraena, Limnebius and Ochthebius in recent years, but most genera in the family have not yet been included in any phylogenetic analyses, particularly most of those which are restricted to the Southern Hemisphere. Using a multimarker molecular matrix, sampling over 40% of described species richness and 75% of currently recognized genera, we infer a comprehensive molecular phylogeny of these predominantly Gondwanan Hydraenidae. Whilst the genera we focus on are morphologically diverse, and currently classified across all four hydraenid subfamilies, our phylogenetic analyses suggest that these Gondwanan genera may instead constitute a single clade. As a result of our findings, the African genus Oomtelecopon Perkins syn.n. is shown to nest within Coelometopon Janssens, the New Zealand Homalaena Ordish syn.n. and Podaena Ordish syn.n. are synonymised with Orchymontia Broun, and the South African Pterosthetops Perkins syn.n. is synonymised with Prosthetops Waterhouse, resulting in Pterosthetopini Perkins syn.n. being synonymised with Prosthetopini Perkins. Mesoceratops Bilton & Jäch gen.n. is erected to accommodate six former members of Mesoceration Janssens, which is shown to be polyphyletic. We propose the replacement name Orchymontia ordishi Jäch & Bilton nom.n. for Homalaena dilatata Ordish, 1984 (now a junior homonym); altogether 39 new combinations are proposed. Our Bayesian divergence times infer an origin for this ‘Gondwana group’ of genera in Africa plus Madagascar in the mid-Cretaceous and suggest that both vicariant and dispersal processes, together with extinctions, have shaped the biogeographic history of these beetles in the Southern Hemisphere during the Cretaceous, resulting in geographically conserved extant lineages. Finally, we reconstruct ancestral habitat shifts across our phylogeny, revealing numerous changes in habitat occupancy in these genera, including multiple origins of fully terrestrial, humicolous taxa in different regions.     

New mesozoic diving beetles (Coleoptera,Dytiscidae) from China

A new subfamily of predaceous diving beetles, Liadytiscinae subfam. nov., including two genera and four species, Liadytiscus gen. nov. (L. cretaceus sp. nov., L. longitibialis sp. nov., and L. latus sp. nov.) and Liadroporus gen. nov. (L. elegans sp. nov.), from the Late Tithonian-Berriasian (Huangbanjigou, Yixian Formation) of China is described. Two new genera, Mesoderus gen. nov. with two species, M. magnus sp. nov. and M. ventralis sp. nov., and Sinoporus gen. nov. with one species, S. lineatus sp. nov., are also described; their position in the system of Dytiscidae remains uncertain. Relationships of the taxa described with Recent and fossil taxa of the same rank and some presumable ecological features of the new taxa are discussed.     

Higher‐level phylogeny of longhorn beetles (Coleoptera: Chrysomeloidea) inferred from mitochondrial genomes

Cerambycidae (longhorn beetles) and related families in the superfamily Chrysomeloidea are important components of forest ecosystems and play a key role in nutrient cycling and pollination. Using full mitochondrial genomes and dense taxon sampling, the phylogeny of Chrysomeloidea with a focus on Cerambycidae and allied families was explored. We used 151 mitochondrial genomes (75 newly sequenced) covering all families and 29 subfamilies of Chrysomeloidea. Our results reveal that (i) Chrysomelidae (leaf beetles) are sister to all other chrysomeloid families; (ii) Cerambycidae sensu stricto (s. s.) is polyphyletic due to the inclusion of other families that split Cerambycidae into a ‘lamiine’ clade comprising Lepturinae sensu lato (s. l.) + (Lamiinae + Spondylidinae) and a ‘cerambycine’ clade comprising Dorcasominae + (Cerambycinae + Prioninae s. l.); (iii) the subfamilies within the two clades of Cerambycidae s. s. were monophyletic, except for the placement of Necydalinae nested in Lepturinae, and the placement of Parandrinae within Prioninae (now considered as tribes Necydalini and Parandrini, respectively); (iv) smaller families were grouped into two major clades: one composed of Disteniidae+Vesperidae and the other composed of Orsodacnidae + (Megalopodidae + Oxypeltidae); (v) relationships among the four major clades were poorly supported but were resolved as ((cerambycines + (Disteniidae + Vesperidae) + Orsodacnidae + (Megalopodidae + Oxypeltidae)) + lamiines. Divergence time analyses estimated that Chrysomeloidea originated ca. 154.1 Mya during the late Jurassic, and most subfamilies of Cerambycidae originated much earlier than subfamilies of Chrysomelidae. The diversification of families within Chrysomeloidea was largely coincident with the radiation of angiosperms during the Early Cretaceous.     

New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the Upper Mesozoic of north‐eastern China,with phylogeny of the family

Specimens of the velvety shore bugs (Hemiptera: Ochtheridae) occur in the Early Cretaceous Yixian Formation in north‐eastern China. We describe two new genera and three new species –Pristinochterus ovatus Yao, Zhang & Ren sp. nov., Floricaudus multilocellus Yao, Ren & Shih gen.n. et sp.n. and Angulochterus quatrimaculatus Yao, Zhang & Ren gen.n. et sp.n. – from this deposit. A cladistic analysis based on a combination of fossil and extant taxa, and their morphological characters, clarifies the phylogenetic status of the new fossils and allows the reconstruction of relationships within the family Ochteridae. Two main clades within Ochteridae are recognized from the cladistic analysis: Pristinochterus Yao Cai & Ren and Floricaudus Yao, Ren & Shih gen.n. form a monophyletic lineage; and Angulochterus Yao, Zhang & Ren gen.n. emerges as a sister group to all extant velvety shore bugs. A key to all fossil and extant genera of Ochteridae is provided.     

Mesozoic Trachypachidae (Insecta: Coleoptera) from China

Abstract: Four well‐preserved beetles, attributed to Eodromeinae within Trachypachidae, are described from the Mesozoic of Inner Mongolia, China. Eodromeus robustus sp. nov., E. daohugouensis sp. nov. and Unda chifengensis sp. nov. are from the Middle Jurassic of Daohugou. The diagnostic characters for the two genera are revised, and all species of these genera are keyed. A new genus and species, Sinodromeus liutiaogouensis gen. nov., sp. nov., is described from the Lower Cretaceous Yixian Formation of Liutiaogou. The current fossil records of Trachypachidae from China are reviewed, and only five species (four described here) can be convincingly attributed to this family. The morphological disparity of Mesozoic Eodromeinae suggests that they evolved a broad spectrum of locomotory lifestyles.