Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae

A new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses. Eulophidae was strongly supported as monophyletic only after exclusion of the enigmatic genus Trisecodes. The subfamilies Eulophinae, Entiinae (=Euderinae) and Tetrastichinae were consistently supported as monophyletic, but Entedoninae was monophyletic only in combined analyses. Six contiguous bases in the 3e′ subregion of the 28S D2 rDNA contributed to placement of nominal subgenus of Closterocerus outside Entedoninae. In all cases, Euderomphalini was excluded from Entiinae, and we suggest that it be retained in Entedoninae. Opheliminae n. stat. is raised from tribe to subfamily status. Trisecodes is removed from Entedoninae but retained as incertae sedis in Eulophidae until its family placement can be determined new placement . The genera Neochrysocharis stat. rev. and Asecodes stat. rev. are removed from synonymy with Closterocerus because strong molecular differences corroborate their morphological differences. Closterocerus (Achrysocharis) germanicus is transferred to the genus Chrysonotomyia n. comb. based on molecular and morphological characters.     

DISCRIMINATIVE POWER OF NUCLEAR rDNA SEQUENCES FOR THE DNA TAXONOMY OF THE DINOFLAGELLATE GENUS PERIDINIUM (DINOPHYCEAE)1

The genus Peridinium Ehrenb. comprises a group of highly diversified dinoflagellates. Their morphological taxonomy has been established over the last century. Here, we examined relationships within the genus Peridinium, including Peridinium bipes F. Stein sensu lato, based on a molecular phylogeny derived from nuclear rDNA sequences. Extensive rDNA analyses of nine selected Peridinium species showed that intraspecies genetic variation was considerably low, but interspecies genetic divergence was high (>1.5% dissimilarity in the nearly complete 18S sequence; >4.4% in the 28S rDNA D1/D2). The 18S and 28S rDNA Bayesian tree topologies showed that Peridinium species grouped according to their taxonomic positions and certain morphological characters (e.g., epithecal plate formula). Of these groups, the quinquecorne group (plate formula of 3′, 2a, 7″) diverged first, followed by the umbonatum group (4′, 2a, 7″) and polonicum group (4′, 1a, 7″). Peridinium species with a plate formula of 4′, 3a, 7″ diverged last. Thus, 18S and 28S rDNA D1/D2 sequences are informative about relationships among Peridinium species. Statistical analyses revealed that the 28S rDNA D1/D2 region had a significantly higher genetic divergence than the 18S rDNA region, suggesting that the former as DNA markers may be more suitable for sequence‐based delimitation of Peridinium. The rDNA sequences had sufficient discriminative power to separate P. bipes f. occultaum (Er. Lindem.) M. Lefèvre and P. bipes f. globosum Er. Lindem. into two distinct species, even though these taxa are morphologically only marginally discriminated by spines on antapical plates and the shape of red bodies during the generation of cysts. Our results suggest that 28S rDNA can be used for all Peridinium species to make species‐level taxonomic distinctions, allowing improved taxonomic classification of Peridinium.     

A global molecular phylogeny of 147 periwinkle species (Gastropoda,Littorininae)

Reid, D. G., Dyal, P. & Williams, S.T. (2012) A global molecular phylogeny of 147 periwinkle species (Gastropoda, Littorininae). —Zoologica Scripta, 41, 125–136. Complete species‐level molecular phylogenies have been published for several genera of Littorinidae (e.g. Echinolittorina, Littoraria). Here we add new sequence data from three genes (28S rRNA, 12S rRNA, cytochrome oxidase c subunit I) for single specimens of an additional 24 species, to make a data set of 147 (97%) of the 152 recognized species of the subfamily Littorininae. This three‐gene data set is analysed to produce a phylogenetic hypothesis for the subfamily, which includes the first complete species‐level phylogeny of the genus Peasiella and the first three‐gene phylogeny of all Littorina species. The non‐planktotrophic species of Littorina have previously been classified together (as subgenus Neritrema), implying a single origin of this developmental mode. Tests of this hypothesis with the new data are inconclusive, and resolution is not improved in a tree constructed from five genes (adding previously published sequences of 16S rRNA and cytochrome b). Using available fossils for calibration we generate a BEAST chronogram, which emphasizes that the radiation of Littorina is more recent than that of other littorinine genera. A database is provided, listing all known species of Littorininae, with their distributions, development, ecology and gene sequences, as a tool for future evolutionary studies of this model group.     

Higher‐level molecular phylogeny of jumping plant lice (Hemiptera: Sternorrhyncha: Psylloidea)

Jumping plant lice (Hemiptera: Psylloidea) are known for a few deleterious pest species worldwide, yet the phylogeny of the group has been poorly understood until very recently. Here, we reconstruct the higher‐level phylogeny for the superfamily Psylloidea based on multilocus DNA sequences, three mitochondrial (COI‐tRNA^leu‐COII, 12S, 16S) and five nuclear (18S, 28S D2, 28S D3, 28S D6–7a, 28S D9–10) gene fragments, using maximum likelihood and Bayesian inference phylogenetic frameworks. Our results are largely congruent with the recent phylogenomic study and partly support prior classification of Psylloidea based mainly on morphology, with the following major exceptions: the family Calophyidae is revealed as polyphyletic, Aphalaridae as paraphyletic with respect to most other taxa of Psylloidea, and Liviidae as paraphyletic with respect to Calophyinae, Psyllidae and Triozidae. Our phylogenetic hypothesis identifies Phacopteronidae and the genus Cecidopsylla Kieffer as the very basal taxa within extant Psylloidea. Sister‐group relationships of Rhinocolinae with Togepsyllinae and of Pachypsyllinae with Homotomidae are also suggested. We present specific discussions for each group of interest recovered in our phylogenetic analysis. One nomenclatorial change is proposed: Spanioneura longicauda (Konovalova) comb.n. , from Psylla Geoffroy.     

Reevaluation of Blapimorpha and Opatrinae: addressing a major phylogeny‐classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology‐based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny‐classification gap in Tenebrionidae. Instead of a holistic approach (family‐level phylogeny), this study uses a bottom‐up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non‐overlapping gene regions: carbamoyl‐phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ～4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non‐monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ～50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher‐level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ～30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily.     

Molecular phylogeny,diagnostics, and diversity of plant‐parasitic nematodes of the genus Hemicycliophora (Nematoda: Hemicycliophoridae)

The genus Hemicycliophora (Nematoda: Hemicycliophoridae) contains 132 valid species of plant‐parasitic nematodes, collectively known as ‘sheath nematodes’. Hemicycliophora spp. are characterized morphologically by a long stylet with rounded basal knobs and a cuticular sheath, present in juvenile and adult stages. Populations of 20 valid and 14 putative species of Hemicycliophora and Loofia from several countries were characterized morphologically using light (LM) and scanning electron microscopy (SEM) and molecularly using the D2‐D3 segments of 28S rRNA and internal transcribed spacer (ITS) rRNA gene sequences. LM and SEM observations provided new details on the morphology of these species. PCR‐restriction fragment length polymorphisms (PCR‐RFLPs) of the D2‐D3 of 28S rDNA were proposed for identification of the species. Phylogenetic relationships within populations of 36 species of the genus Hemicycliophora using 102 D2‐D3 of 28S rDNA and 97 ITS rRNA gene sequences as inferred from Bayesian analysis are reconstructed and discussed. Ancestral state reconstructions of diagnostic characters (body and stylet length, number of body annuli, shape of vulval lip and tail), using maximum parsimony and Bayesian inference, revealed that none of the traits are individually reliable characters for classifying the studied sheath nematode. The Shimodaira–Hasegawa test rejected the validity of the genus Loofia. This is the most complete phylogenetic analysis of Hemicycliophora species conducted so far. © 2014 The Linnean Society of London     

The phylogeny and limits of Elateridae (Insecta,Coleoptera): is there a common tendency of click beetles to soft‐bodiedness and neoteny?

Kundrata, R. & Bocak, L. (2011). The phylogeny and limits of Elateridae (Insecta, Coleoptera): is there a common tendency of click beetles to soft‐bodiedness and neoteny? —Zoologica Scripta, 40, 364–378. Phylogenetic relationships in Elateroidea were investigated using partial 18S and 28S rDNA and rrnl and cox1 mtDNA sequences with special interest in the phylogeny of Elateridae and the position of soft‐bodied lineages Drilidae and Omalisidae that had been classified as families in the cantharoid lineage of Elateroidea until recently. Females in these groups are neotenic and almost completely larviform (Drilidae) or brachypterous (Omalisidae). The newly sequenced individuals of Elateridae, Drilidae, Omalisidae and Eucnemidae were merged with previously published datasets and analysed matrices include either 155 taxa with the complete representation of fragments or 210 taxa when some fragments were missing. The main feature of inferred phylogenetic trees was the monophyly of Phengodidae + Rhagophthalmidae + Omalisidae + Elateridae + Drilidae with Omalisidae regularly occupying a basal node in the group; Drilidae were embedded as a terminal lineage in the elaterid subfamily Agrypninae and soft‐bodied Cebrioninae were a part of Elaterinae. The soft‐bodied males and incompletely metamorphosed females originated at least three times within the wider Elateridae clade. Their atypical morphology has been considered as a result of long evolutionary history and they were given an inappropriately high rank in the previous classifications. The frequent origins of these modifications seem to be connected with modifications of the hormonal regulation of the metamorphosis. The superficial similarity with other soft‐bodied lineages, such as Cantharidae, Lycidae, Lampyridae, Phengodidae and Rhagophthalmidae is supposed to be a result of homoplasious modifications of the ancestral elateroid morphology due to the incomplete metamorphosis. The results of phylogenetic analyses are translated in the formal taxonomic classification. Most Drilidae are placed in Elateridae as a tribe Drilini in Agrypninae, whilst Pseudeuanoma and Euanoma are transferred from Drilidae to Omalisidae. The subfamily Cebrioninae is placed in Elaterinae as tribes Cebrionini and Aplastini. Oxynopterini, Pityobiini and Semiotini are lowered from the subfamily rank to tribes and classified in Denticollinae.     

Simenoside A,a New Triterpenoid Saponin from Gypsophila simonii Hub.‐Mor.

Saponins are amphiphilic glycoconjugates which give soap‐like foams in H₂O. A new triterpenoid saponin, simenoside A ( 1 ), based on gypsogenin aglycone, and the known saponin 2 were isolated from Gypsophila simonii Hub.‐Mor. The structure of the new saponin was elucidated as 3‐O‐β‐D ‐galactopyranosyl‐(1→2)‐[β‐D ‐xylopyranosyl‐(1→3)]‐β‐D ‐glucuronopyranosylgypsogenin 28‐O‐β‐D ‐glucopyranosyl‐(1→3)‐[β‐D ‐glucopyranosyl‐(1→2)‐β‐D ‐xylopyranosyl‐(1→4)]‐α‐L ‐rhamnopyranosyl‐(1→2)‐β‐D ‐fucopyranosyl ester on the basis of extensive spectral analyses and chemical evidence. Saponins 1 and 2 were isolated from G. simonii for the first time.     

Inhibition of NO Production by Grindelia argentina and Isolation of Three New Cytotoxic Saponins

A bioassay‐guided phytochemical analysis of the ethanolic extract of Grindelia argentina Deble & Oliveira ‐Deble (Asteraceae) allowed the isolation of a known flavone, hispidulin, and three new oleanane‐type saponins, 3‐O‐β‐D ‐xylopyranosyl‐(1→3)‐β‐D ‐glucopyranosyl‐2β,3β,16α,23‐tetrahydroxyolean‐12‐en‐28‐oic acid 28‐O‐β‐D ‐xylopyranosyl‐(1→2)‐β‐D ‐apiofuranosyl‐(1→3)‐β‐D ‐xylopyranosyl‐(1→3)‐α‐L ‐rhamnopyranosyl‐(1→2)‐α‐L ‐arabinopyranosyl ester ( 2 ), 3‐O‐β‐D ‐glucopyranosyl‐2β,3β,23‐trihydroxyolean‐12‐en‐28‐oic acid 28‐O‐β‐D ‐xylopyranosyl‐(1→2)‐β‐D ‐apiofuranosyl‐(1→3)‐β‐D ‐xylopyranosyl‐(1→3)‐α‐L ‐rhamnopyranosyl‐(1→2)‐α‐L ‐arabinopyranosyl ester, ( 3 ) and 3‐O‐β‐D ‐xylopyranosyl‐(1→3)‐β‐D ‐glucopyranosyl‐2β,3β,23‐trihydroxyolean‐12‐en‐28‐oic acid 28‐O‐β‐D ‐xylopyranosyl‐(1→2)‐β‐D ‐apiofuranosyl‐(1→3)‐β‐D ‐xylopyranosyl‐(1→3)‐α‐L ‐rhamnopyranosyl‐(1→2)‐α‐L ‐arabinopyranosyl ester ( 4 ), named grindeliosides A–C, respectively. Their structures were determined by extensive 1D‐ and 2D‐NMR experiments along with mass spectrometry and chemical evidence. The isolated compounds were evaluated for their inhibitory activities against LPS/IFN‐γ‐induced NO production in RAW 264.7 macrophages and for their cytotoxic activities against the human leukemic cell line CCRF‐CEM and MRC‐5 lung fibroblasts. Hispidulin markedly reduced LPS/IFN‐γ‐induced NO production (IC₅₀ 51.4 μM ), while grindeliosides A–C were found to be cytotoxic, with grindelioside C being the most active against both CCRF‐CEM (IC₅₀ 4.2±0.1 μM ) and MRC‐5 (IC₅₀ 4.5±0.1 μM ) cell lines.     

Characterization of Gambierdiscus lapillus sp. nov. (Gonyaulacales,Dinophyceae): a new toxic dinoflagellate from the Great Barrier Reef (Australia)

Gambierdiscus is a genus of benthic dinoflagellates found worldwide. Some species produce neurotoxins (maitotoxins and ciguatoxins) that bioaccumulate and cause ciguatera fish poisoning (CFP), a potentially fatal food‐borne illness that is common worldwide in tropical regions. The investigation of toxigenic species of Gambierdiscus in CFP endemic regions in Australia is necessary as a first step to determine which species of Gambierdiscus are related to CFP cases occurring in this region. In this study, we characterized five strains of Gambierdiscus collected from Heron Island, Australia, a region in which ciguatera is endemic. Clonal cultures were assessed using (i) light microscopy; (ii) scanning electron microscopy; (iii) DNA sequencing based on the nuclear encoded ribosomal 18S and D8‐D10 28S regions; (iv) toxicity via mouse bioassay; and (v) toxin profile as determined by Liquid Chromatography‐Mass Spectrometry. Both the morphological and phylogenetic data indicated that these strains represent a new species of Gambierdiscus, G. lapillus sp. nov. (plate formula Po, 3′, 0a, 7″, 6c, 7‐8s, 5?, 0p, 2″″ and distinctive by size and hatchet‐shaped 2′ plate). Culture extracts were found to be toxic using the mouse bioassay. Using chemical analysis, it was determined that they did not contain maitotoxin (MTX1) or known algal‐derived ciguatoxin analogs (CTX3B, 3C, CTX4A, 4B), but that they contained putative MTX3, and likely other unknown compounds.     

Pythiopina,an enigmatic subtribe of darkling beetles (Coleoptera: Tenebrionidae: Pedinini): taxonomic revision,microtomography, ecological niche models and phylogenetic position

Morphological, anatomical, and distributional data concerning the S outh A frican endemic beetle subtribe P ythiopina (T enebrionidae: P edinini) are revised. Five species, representing two genera, are recognized. Included in this total is one new species (Meglyphus mariae K amiński sp.n. ). The following species are placed in synonymy: Meglyphus ciliatipes [=Meglyphus calitzensis syn.n. ]; Meglyphus laenoides [=Meglyphus andreaei syn.n. ; =Meglyphus namaqua syn.n. ]. Microtomographic models for all valid P ythiopina species, including the holotype of the newly described species, are presented and analysed. Endoskeleton morphology (specifically characters of the tentorium and metendosternite) proved to be informative at the specific and generic levels. An identification key is provided to all known species of the subtribe. Environmental niche models are presented for the majority of species. A molecular phylogeny of P edinini based on six genetic loci (28S : D 1–D 3 region; 28S : D 4–D 5 region, COII , A rgK , CAD 2, wg) was also produced to explore the phylogenetic position of P ythiopina. This analysis is the first to include representatives of all seven subtribes of P edinini, and supports a sister relationship between P ythiopina and the P alaearctic subtribe D endarina. Results also suggest the existence of a second pair of sister taxa within P edinini (in addition to M elambiina) with an amphitropical A frican distribution. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:285AD87A‐46B1‐4FE9‐BC57‐949EA1F70D49 .     

A molecular phylogenetic framework for the subfamily Ocenebrinae (Gastropoda,Muricidae)

The Ocenebrinae is a subfamily of marine predatory gastropods known as oyster and mussel drills. Their current phylogenetic framework is traditionally based on shell and radular characters, but a consensus on relationships among genera is still lacking. We investigated the molecular phylogeny of Ocenebrinae using 50 species and DNA data from one nuclear (28S) and two mitochondrial (COI and 16S) genes, the largest data set so far assembled for this subfamily. We found support for the monophyly of the Ocenebrinae, and species were divided into four major lineages. Within groups, genera had similar geographic distributions, suggesting that except in a few cases, species diversification within clades occurred without range expansions. We discuss the phylogenetic distribution of a labral tooth and a sealed siphonal canal, two characteristic ocenebrine features. We also show that Ocinebrina species in the north‐eastern Pacific are not monophyletic with north‐eastern Atlantic and Mediterranean species, and that the Ocinebrina edwardsii species complex belongs to Ocenebra.     

Molecular phylogeny of the fungus gnat family Mycetophilidae (Diptera,Mycetophiliformia)

Abstract A molecular phylogeny of the fungus gnat family Mycetophilidae based on the nuclear 18S, 28S, and the mitochondrial 16S rRNA genes is presented. The total alignment included 58 taxa and 1704 bp. The family was recovered as monophyletic in parsimony and Bayesian analyses. In the Bayesian analysis, Mycetophilinae and its two tribes, Mycetophilini and Exechiini, were monophyletic with good statistical support. The subfamily Mycomyinae was found consistently in a sister‐group relationship to Mycetophilinae. Gnoristinae was rendered paraphyletic, subtending Mycomyinae and Mycetophilinae. Within Gnoristinae, the genera Coelosia Winnertz, Boletina Staeger, Gnoriste Meigen group with Docosia Winnertz, usually considered to be a member of Leiinae. No support was found for the monophyly of the subfamilies Sciophilinae and Leiinae.     

Phylogenetic position of the house dust mite subfamily Guatemalichinae (Acariformes: Pyroglyphidae) based on integrated molecular and morphological analyses and different measures of support

Based on multilocus phylogenetic analyses (18S, 28S, EF1‐α, SRP54, HSP70, CO1, 10 860 nt aligned), we show that the house dust mite subfamily Guatemalichinae is nested within non‐onychalgine pyroglyphid mites and forms the sister group to the genus Sturnophagoides (bootstrap support 100, posterior probability 1.0). Because high bootstrap support values may be misleading in the presence of incongruence, we evaluate robustness of the Guatemalichinae+Sturnophagoides clade using: (1) internode certainty indices to estimate the frequency of conflicting bipartitions in maximum‐likelihood bootstrap trees, (ii) consensus networks to investigate conflict among different loci; and (iii) statistical hypothesis testing based on information theory, both multi‐scale and regular bootstrap. Results suggest that this grouping is very well supported given the data. The molecular analyses were integrated with detailed morphological study using scanning electron and light microscopy. We suggest that the subfamilial status of Guatemalichinae should be reconsidered, and this lineage should be placed within the subfamily Dermatophagoidinae. The latter subfamily is currently accepted in the literature as a monophyletic group but was here inferred as paraphyletic and was not supported by any morphological synapomorphy. The paraphyly involved the most species‐rich and medically important genus, Dermatophagoides. Our findings suggest the need for a comprehensive revision of the higher‐level relationships of pyroglyphid house dust mites using both DNA sequences and morphology coupled with a broad taxonomic sampling.     

Molecular analysis and comparative morphology to resolve a complex of cryptic Xiphinema species

Gutiérrez‐Gutiérrez, C., Palomares‐Rius, J.E., Cantalapiedra‐Navarrete, C., Landa, B.B., Esmenjaud, D. & Castillo, P. (2010). Molecular analysis and comparative morphology to resolve a complex of cryptic Xiphinema species. —Zoologica Scripta, 39, 483–498. During nematode surveys in cultivated and natural environments in southern Spain nine populations of parthenogenic Xiphinema species tentatively identified as Xiphinema cf. pyrenaicum and one population morphologically close to Xiphinema turcicum were detected. Surveys in southern France also identified one population resembling X. pyrenaicum. We developed a comparative study among these related Xiphinema species, including topotypes of two species of this group previously synonymized, viz. Xiphinema hispanum and Xiphinema sphaerocephalum, by considering morphological and morphometrical features together with molecular data from nuclear ribosomal RNA genes (D2‐D3 expansion segments of 28S, ITS1, and partial 18S). Morphological and morphometrical results identified eight of the Spanish populations as Xiphinema nuragicum (previously synonymized with X. pyrenaicum) whereas the ninth population was identified as Xiphinema adenohystherum (also synonymized with X. pyrenaicum). The species X. adenohystherum, X. nuragicum, X. pyrenaicum, and X. sphaerocephalum were shown to be morphologically almost indistinguishable but clearly separated by phylogenetic analyses, thus constituting a complex of cryptic species. Consequently, X. adenohystherum, X. nuragicum, and X. sphaerocephalum were re‐established as valid species. Similarly, X. hispanum (morphologically similar to X. aceri) was also shown as a valid species. Xiphinema turcicum, morphologically related to X. pyrenaicum complex by its rounded tail, uterus with a pseudo‐Z‐differentiation and small spines, was phylogenetically distant to these species based on D2‐D3 expansion segments of 28S and ITS1, which suggests a morphological convergence in their evolution.     

A scolopocryptopid centipede (Chilopoda: Scolopendromorpha) from Mexican amber: synchrotron microtomography and phylogenetic placement using a combined morphological and molecular data set

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryptopinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojovelensis sp. nov. , displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non‐invasive 3D synchrotron microtomography using X‐ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein‐coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein‐coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopocryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 768–786.