Hitchhikers at the dinner table: a revisionary study of a group of ant parasitoids (Hymenoptera: Eucharitidae) specializing in the use of extrafloral nectaries for host access

Extrafloral nectaries (EFNs) are nectar‐secreting glands found on plants independent of their flowers. EFNs are diverse in form, present on a wide variety of plants, and their secretions are known to recruit ants. However, much less information has been published on insects with known EFN associations other than ants. Two distinct species groups of Orasema Cameron (Hymenoptera: Eucharitidae) deposit their eggs close to the EFNs of their plant hosts. The simulatrix group comprises six species found in deserts and xeric shrublands of the southwestern United States and Mexico. This species group is revised, retaining O. aureoviridis, O. beameri and O. simulatrix as valid species, and describing O. cancellata sp.n. , O. difrancoae sp.n. and O. zahni sp.n. The wayqecha group is from Peru and Colombia and includes the newly described O. wayqecha sp.n. and O. quadrimaculata sp.n. Members of the simulatrix group oviposit near EFNs of Chilopsis linearis Cav. (Bignoniaceae), Prosopis glandulosa Torr. (Fabaceae), Prosopis velutina Wooton (Fabaceae) and Populus angustifolia James (Salicaceae), whereas Orasema wayqecha oviposit on leaves of two species in the family Primulaceae. Monophyly of the two species groups is proposed based on a molecular analysis of ribosomal (28S and 18S) and mitochondrial (cytochrome c oxidase subunit I) DNA, morphological features of the adults and planidia larvae, and their shared behavioural association with EFNs. Adults of both species groups have an expanded postgena that encloses the mouthparts, but are otherwise morphologically divergent. The planidia of both groups also share several features, including long cerci that may facilitate their movements within EFNs. Oviposition near EFNs is proposed as a means of increasing encounter rates of the first‐instar larvae with their myrmicine ant host; however, it remains unclear whether the planidia are transported directly by the foraging workers of their host Pheidole (Formicidae: Myrmicinae) or indirectly with the help of an intermediate host. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:EA112E3D-52BE-45D1-B03B-8764E8C94BE0 .     

Eucharitid Parasitism of the Neotropical Ant Ectatomma tuberculatum: Parasitoid Co‐occurrence,Seasonal Variation,and Multiparasitism1

Multiparasitism is reported for the first time in Eucharitidae (Hymenoptera: Chalcidoidea). Dilocantha lachaudii, Isomerala coronata, and Kapala sp. were found simultaneously attacking the same population of Ectatomma tuberculatum (Olivier) (Hymenoptera: Formicidae: Ectatomminae) in southern Mexico. Adults of D. lachaudii and I. coronata completed development on the same individual host on three occasions. The genus Kapala is reported for the first time as a parasitoid of E. tuberculatum, and I. coronata is reported in Mexico for the first time.     

A new extant family of primitive moths from Kangaroo Island,Australia, and its significance for understanding early Lepidoptera evolution

We report the first discovery since the 1970s of a new extant family (Aenigmatineidae fam.n. ) of homoneurous moths, based on the small Aenigmatinea glatzella sp.n . from Kangaroo Island off southern Australia. It exhibits a combination of extraordinary anatomical characters, and, unlike most homoneurous moths, its larva is a conifer‐feeder (stem mining in Callitris, Cupressaceae). While the adult's mouthparts are strongly regressed, evidence from other morphological characters and from a Bayesian analysis of 25 genetic loci convincingly places the taxon among Glossata (‘tongue moths’). An unexpected tongue moth clade including Acanthopteroctetidae and Neopseustidae, suggested with low support in recent molecular analyses, remarkably becomes strongly supported when Aenigmatinea is included in the molecular analysis; the new taxon becomes subordinated in that clade (as sister group to Neopseustidae) and the clade itself appears as the sister group of all Heteroneura, representing the vast majority of all Lepidoptera. Including Aenigmatinea into the analysis thereby strengthens the surprising indication of non‐monophyly of Myoglossata, and the new phylogeny requires an additional number of ad hoc assumptions of convergence/character reversals in early Lepidoptera evolution. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:44393B52‐1889‐431A‐AB08‐6BBCF8F946B8 .     

Metaphycus parthenolecanii sp. n. (Hymenoptera: Encyrtidae), a parasitoid of Parthenolecanium corni (Hemiptera: Coccidae) in Iran

Metaphycus parthenolecanii Japoshvili sp. n. (Hymenoptera, Chalcidoidea, Encyrtidae), a parasitoid of the European fruit lecanium Parthenolecanium corni (Bouché, 1844) (Hemiptera, Coccoidea, Coccidae) in Iran, is described and illustrated.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:3B846DA1-7E97-442B-B4AF-B5C727A72521     

A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure

An unusual Cretaceous trap jaw ant is described from Burmese amber dated to the Late Cretaceous. Linguamyrmex vladi gen.n. sp.n. is distinguished by an unusual suite of morphological characters indicating specialized predatory behaviour and an adaptive strategy no longer found among modern ant lineages. The clypeus, highly modified as in other closely related haidomyrmecine hell ants, is equipped with a paddle‐like projection similar to Ceratomyrmex. X‐ray imaging reveals that this clypeal paddle is reinforced, most probably with sequestered metals. Presumably this fortified clypeal structure was utilized in tandem with scythe‐like mandibles to pin and potentially puncture soft‐bodied prey. This unique taxon, which stresses the diversity of stem‐group ants, is discussed in the context of modern and other Cretaceous trap jaw ant species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:40D636A3‐4D88‐470A‐BC5B‐85ABFD1A49E2 .     

Out of the Neotropics: newly discovered relictual species sheds light on the biogeographical history of spider ants (Leptomyrmex,Dolichoderinae, Formicidae)

Spider ants of the genus Leptomyrmex Mayr (Hymenoptera: Formicidae: Dolichoderinae) are conspicuous species of Australasian rainforests, with putative fossil relatives in the Neotropics and Europe. There is longstanding debate over the biogeographical history of the genus, with the Palaearctic and Neotropical regions proposed as alternate centres of origin. We propose a resolution of this debate with the recent discovery and analysis of an extant species from central Brazil, L. relictus sp.n. , which we describe from workers, males and brood. We sequence ten nuclear genes in the new species and in several Australian Leptomyrmex species, and append these data to a 54‐taxon, 10‐gene data matrix previously generated for the subfamily Dolichoderinae. We conduct phylogenetic and divergence dating analyses, and re‐evaluate the fossil record of the group. We recover Leptomyrmex relictus sp.n. as a member of the Leptomyrmex clade with high support. It is sister to the Australasian species, and the genus Leptomyrmex is, in turn, sister to a pair of Neotropical genera, Forelius and Dorymyrmex. We infer a Neotropical origin for the genus and estimate a mid‐Eocene (46 Ma, 95% CI 56 to 36 Ma) origin for the crown genus and an Oligocene origin for the Australasian clade (29 Ma, 95% CI 40 to 19 Ma). We confirm placement of the Dominican amber species ?L. neotropicus Baroni Urbani in the genus but reject a close relationship with the Palaearctic fossil taxa ?Leptomyrmula Emery and ?Usomyrma Dlussky, Radchenko & Dubovikoff, considering them incertae sedis in the subfamily (Dolichoderinae). In contrast to the mesophilic preferences of the Australasian species of Leptomyrmex, the new Brazilian species inhabits cerrado (dry savannah). Our results support a Neotropical origin for spider ants with dispersal to Australia. Rafting on west‐bound currents and/or a historical diversity imbalance between Australia and South America are proposed as alternate hypotheses to explain a pattern of biased E–W mid‐Tertiary dispersal for ants with austral distributions. This pattern is suggested by our results in conjunction with observations of other ant clades. Overall, our findings highlight the value of integrated taxonomy, critical interpretation of morphology, and a comparative phylogenetic framework when conducting palaeontological and biogeographical studies of insect species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:6E9E6617‐6E53‐40B8‐82C7‐67F89A83C553 .     

Two new species of Torymidae (Hymenoptera: Chalcidoidea) reared from flower heads of Echinops orientalis Trautvetter in south-eastern Turkey

Two new species of torymids, Pseudotorymus hasanberkayi sp. n. and Microdontomerus kahramanmarasensis sp. n. (Hymenoptera: Torymidae), were reared from flower heads of Echinops orientalis Trautvetter (Asteraceae) collected in Kahramanmara? province, south-eastern Turkey. The new species are described and their diagnostic characters are illustrated.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:848B4E9D-AE8A-4EF8-8072-1F28BE21ADA1     

Phylogeny,divergence times and biogeography of window flies (Scenopinidae) and the therevoid clade (Diptera: Asiloidea)

The evolution of the ‘therevoid’ clade, with an emphasis on window flies (Scenopinidae), is presented by combining DNA sequence data with morphological characters for living and fossil species. The therevoid clade represents a group of four families (Apsilocephalidae, Evocoidae, Scenopinidae and Therevidae) of lower brachyceran Diptera in the superfamily Asiloidea. A comprehensive phylogenetic analysis using parsimony and likelihood methods was undertaken using extensive taxon sampling from all families and subfamilies, and compared with outgroup taxa sampled from the related families Asilidae, Mydidae, Apioceridae and Empididae. Fifty‐nine morphological characters (adult, larval and pupal) were combined with 6.4 kb of DNA sequences for two ribosomal genes (16S and 18S ribosomal DNA) and three protein‐encoding genes [cytochrome oxidase I (COI), triose phosphate isomerase (TPI) and the CPSase region of carbamoyl‐phosphate synthase‐aspartate transcarbamoylase‐dihydroorotase (CAD)]. Results from combined analyses of morphological and molecular data for 78 taxa representing all families of the therevoid clade are presented. Specific hypotheses of the relationship between respective families and subfamilies were tested statistically using four‐cluster likelihood mapping. The therevoid clade is a well‐supported monophyletic group within Asiloidea, with Evocoidae sister to Apsilocephalidae and Therevidae sister to Scenopinidae. Temporal and zoogeographical aspects of therevoid clade evolution were investigated using Bayesian divergence time estimates and Lagrange ancestral range scenarios. The effect of inclusion of fossils as terminal taxa on phylogenetic and divergence time estimation was investigated, with morphological scoring for fossil representatives included in the analyses rather than used simply as minimum age constraints. In each analysis there was either improvement in estimation, or only marginal and localized loss in tree resolution, and with younger estimates of divergence time across the tree. The historical biogeography of the therevoid clade was examined with multiple trans‐Antarctic vicariance events between Australasia and South America evident during the Late Cretaceous to early Palaeogene. Scenopininae is newly subdivided into two tribes, Metatrichini trib.n. and Scenopinini Fallén stat.r. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4974EBF8‐3117‐4189‐B6DE‐7D5BF9B23E53 .     

Mutualism between armoured scale insects and ants: new species and observations on a unique trophobiosis (Hemiptera: Diaspididae; Hymenoptera: Formicidae: Melissotarsus Emery)

The association between African armoured scale insects (Hemiptera: Coccoidea: Diaspididae) and ants belonging to Melissotarsus Emery (Hymenoptera: Formicidae: Myrmicinae) is the only trophobiosis known in which ants do not receive honeydew or nectar in exchange for protection and other services. Food reward for the ants in this mutualism remains unknown, despite repeated suggestions that diaspidids are consumed by the associated ants, thus serving as ‘domestic cattle’. We describe new observations on interactions between Melissotarsus emeryi Santschi and the diaspidid Morganella conspicua (Brain) from South Africa. Worker ants exhibited previously undescribed tending behaviours, most notably a ‘squeezing and licking’ performed on an adult female diaspidid and ‘culling’, in which a worker removed an adult female armoured scale from the host plant. These could represent the gathering of secretory products and the cultivation of an individual for consumption, respectively. An ant exclusion study over 12 days of isolation showed that adult female diaspidids and second‐instar nymphs secreted no wax or exudates that attending ants would ordinarily collect. Workers of M. emeryi did not defend their nest against invading colonies of Crematogaster and other unidentified ants: we hypothesize that the primary mode of defence is maintenance of isolation within galleries. We describe three new ant‐associated diaspidid species: Affirmaspis cederbergensis Schneider sp.n. from South Africa, Diaspis doumtsopi Schneider sp.n. from Cameroon, and Melissoaspis incola Schneider sp.n. from Madagascar. Melissoaspis formicaria (Ben‐Dov) comb.n. is transferred from Morganella (Brain). Diagnostic characteristics for Melissoaspis Ben‐Dov are revised, and additional taxonomic information defining this genus allows ease of identification. An updated identification key to the species of ant‐associated diaspidids is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:632C1122-954A-4DE6-B946-8E6AB6A4D27F .     

Phylogeny of a new supertribe Cephenniitae with generic review of Eutheiini and description of a new tribe Marcepaniini (Coleoptera: Staphylinidae: Scydmaeninae)

Genera of Eutheiini are reviewed and Eutheimorphus is removed from this tribe of ant‐like stone beetles (Scydmaeninae) and transferred to Cephenniini. A monogeneric Marcepaniini trib.n. is described to accommodate Marcepania gen.n. from Malaysia, with five species: M. semengohensis sp.n. (the type species of Marcepania), M. tuberculata sp.n. , M. seramaensis sp.n. , M. minutissima sp.n. and M. elongata sp.n. A phylogenetic analysis of all genera of Cephenniini, Eutheiini and Marcepaniini based on adult morphological characters resulted in recovering a well‐supported monophyletic clade Eutheiini + (Marcepaniini + Cephenniini) and these tribes are included in Cephenniitae stat.n. (Eutheiini and Cephenniini are therefore removed from Scydmaenitae). Only a weak support for monophyly of Eutheiini was found, but morphological characters allow for maintaining this presumably relic group as a separate tribe. Previously proposed monophyletic groups within Cephenniini were recovered as such, but after inclusion of Eutheimorphus, a sister taxon to the ‘Cephennomicrus group’, the latter lineage gained weak statistical support. The evolutionary history of Cephenniitae is discussed, with focus on known northern hemisphere fossils classified in Scydmaenitae and Hapsomelitae, but possibly closely allied to Cephenniitae. Establishing the supertribe Cephenniitae is the first step toward a profound reclassification of Scydmaeninae on a robust phylogenetic basis. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:B0E1B12D-9587-4C4F-A908-A12A0C424A8C .     

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n. , has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796‐6E70‐4D81‐BB34‐0FEEA765DC25 .     

A new species of Copidosoma (Hymenoptera: Encyrtidae) from Iran

Copidosoma isfahan Japoshvili sp. n. (Hymenoptera: Chalcidoidea: Encyrtidae), a parasitoid of Altenia mersinella (Staudinger, 1879) (Lepidoptera: Gelechiidae) on pistachio (Pistacia vera L.) is described and illustrated from Iran.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:F1BCFE50-AD05-40BB-A2AC-2AF86956FE5F     

Integrative taxonomy reveals phenotypic plasticity in the freshwater mussel Contradens contradens (Bivalvia: Unionidae) in Thailand,with a description of a new species

The freshwater mussel Contradens contradens (Lea, 1838) occurs in most types of freshwater habitats throughout Thailand. The species shows extensive variation in shell morphology, which has led to the recognition of six different subspecies. In this study, the validity of these six subspecies plus one unknown species was assessed using an integrative taxonomic approach. Geometric morphometric analyses revealed significant differences in shell shapes among these six nominal morphological subspecies, although a considerable degree of overlap was detected in some groups. In contrast, the phylogenetic tree obtained from the concatenated data of mitochondrial COI and nuclear H3 gene sequences and molecular species delimitation analyses revealed only three supported clades. These clades are proposed herein as three distinct species, and strongly corresponded to the biogeographically disjunct drainage systems in Thailand. They consisted of the (i) C. contradens clade found in the Chao Phraya Basin and other rivers that drain into the Gulf of Thailand, (ii) C. crossei clade that is restricted to the Middle Mekong Basin, and (iii) a clade containing only the newly discovered species from Huai Luang River in the north-east of Thailand, which is described herein as Contradens rolfbrandti Jeratthitikul & Panha, sp. nov. Speciation among these congeners was probably caused by the restriction of gene flow due to the past geomorphology of the river systems. The intraspecific variation in the shell shape detected here does not reflect the evolution of the mussel, but rather is evidence of phenotypic plasticity.http://zoobank.org/urn:lsid:zoobank.org:pub:649B6093-E1DD-4FD8-8185-A4696C43AD36; http://zoobank.org/urn:lsid:zoobank.org:act:777DDE86-8397-4FF3-AA78-0BE0F34611F4     

Description of a new species of deep-water crab of the genus Homolodromia A. Milne-Edwards, 1880 from the northern Indian Ocean (Crustacea: Decapoda: Brachyura: Homolodromiidae)

ABSTRACT

Homolodromia rajeevani, a new species of deep-water homolodromiid sponge crab, is described from the northern Indian Ocean (Arabian Sea, depth 957 m, and Bay of Bengal, 645 m), and is the first record of the genus from the area. This species resembles the western Indian Ocean species, namely, Homolodromia bouvieri Doflein, 1904, in having 2 terminal spines on the propodi of the last two pereopods, but can be easily distinguished from the latter species by the inflated carapace, simple long setae on carapace and appendages, slender pseudo-rostral spines separated by a U-shaped base, and a slender arched dactylus of cheliped with maximum elevation at proximal part which bears broadly circular depressions with sparse setae. The most diagnostic character is the higher number of spines on the occlusal surfaces of propodal thumbs and dactyli of the pseudochela of the last two pereopods as compared to H. bouvieri. A key for the identification of the species under the genus Homolodromia is also provided.

http://zoobank.org/urn:lsid:zoobank.org:pub:48894F49-5124-4723-9FF2-3D30FB536DA5     

A new species of the genus Haba Semenov, 1954 (Hymenoptera: Chrysididae) from Iran,with a key to species

Haba persica Strumia & Fallahzadeh sp. n. (Hymenoptera: Chrysididae: Elampini), from mountains in southwestern Iran, is described and illustrated.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:9D537878-6215-4874-8970-7F7DE8F57422     

A new species of ant-loving cricket Myrmecophilus Berthold, 1827 from Cyprus (Orthoptera: Myrmecophilidae)

A new species of ant-loving cricket, Myrmecophilus cyprius sp. n., collected in the Republic of Cyprus from the nests of Messor structor (Latreille, 1798) (Hymenoptera: Formicidae), is described and illustrated. The species belongs to the subgenus Myrmophilina Silvestri, 1912.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:B588546E-FC72-4996-A0C2-583BCE46905E     

An integrative approach to species discrimination in the Anagrus atomus group sensu stricto (Hymenoptera: Mymaridae), with a description of a new species

Egg parasitoids of the genus Anagrus Haliday (Hymenoptera: Mymaridae) are natural enemies of many pests around the world. We used an integrative approach to characterize some species belonging to the Anagrus atomus group, using specimens reared from leafhoppers infesting some Lamiaceae. Starting from morphological identifications based on available keys, we carried out a multi-locus genetic characterization using phylogenetic and species delimitation analyses, and integrated it with biological and morphometric evidence. This approach revealed the existence of a new species, A. nepetellae sp. nov., here described along with its phenological traits. The approach also allowed us to characterize the expected taxonomic stability of other putative species within the group. We propose two new species groups called the atomus group sensu stricto and the vilis group, with the goal of streamlining the taxonomy of the atomus group. We also present a key for the identification of females of the European species of the atomus group sensu stricto. Some specimens show heterozygosity in 28S-D2 sequences, suggesting hybridization between A. atomus and A. nepetellae, which, if it is common between other species as well, could partly explain the taxonomic problems in the genus Anagrus. DNA sequencing of specimens reared by the same biofactory at different times suggests involuntary contamination leading to the displacement of a laboratory strain of A. atomus by A. nepetellae, which may have resulted from undetected partial reproductive compatibility.http://zoobank.org/urn:lsid:zoobank.org:pub:DFCD7654-5AA5-4E4F-AE48-24377BAACEFE