The female postabdomen and genitalia of the basal moth family Heterobathmiidae (Insecta: Lepidoptera): Structure and phylogenetic significance

Female Heterobathmia have the segments behind VIII forming a compact ‘terminal unit’ with a large saddle-shaped dorsal plate and a membranous ventroposterior surface bearing the separate gonopore and anus. While females of most of the nine known species are overall similar, Heterobathmia valvifer is unique amongst lepidopterans in possessing paired ventral appendages (‘ovipositor valves’) arising from the intersegmental groove following segment VIII; evidence from musculature contradicts an interpretation of these appendages structures as ‘true’ ovipositor valves. The ventroposterior wall of the terminal unit in H. valvifer bears paired sclerites, possible homologues of the ‘ventral rods’ in basal Lepidoptera-Glossata. In Heterobathmia megadecella sclerites on paired longitudinal elevations in comparable positions probably are/include homologues of these sclerites. Their similarity with paired sclerotizations in the corresponding region of hydrobiosid caddisflies is noted. A prominent frame-like sclerotization in the genital chamber, located in front of the spermathecal duct origin, is present only in H. megadecella.Putative heterobathmiid autapomorphies include an enlarged ‘subgenital plate’ on venter VIII, absence of apophyses on segment VIII, shortened apophyses on the terminal unit, multilobed accessory glands (but their ‘type 1’ secretory epithelium is plesiomorphic at this level), a conspicuous papilla in the chamber cuticle bearing the opening of the ductus bursae on its apex, and inwards-pointing spines in the ductus bursae. A variably developed thickening of the anterior genital chamber intima is another putative family autapomorphy, while an extreme thickening of the posterior intima seen in Heterobathmia pseuderiocrania is not of general occurrence in heterobathmiids. A sistergroup relationship between Heterobathmiidae and Glossata is supported by their fully developed ‘2-compartment section’ of the spermathecal duct and losses of some likely lepidopteran groundplan muscles.     

First Potwarmus from the Miocene of Saudi Arabia and the early phylogeny of murines (Rodentia: Muroidea)

Potwarmus is recorded for the first time in Saudi Arabia. The material comes from the Middle Miocene of the Hofuf Formation at Al‐Jadidah. This species, Potwarmus flynni sp. nov. , is described and compared with all species currently placed in the genera Potwarmus, Dakkamys, Paradakkamys, Vallaris, and some species of ‘Myocricetodon’ (i.e. the so‐called ‘primitive dendromurids’) as well as with two of the most primitive definitive murines (namely Antemus chinjiensis and Progonomys debruijni). Potwarmus flynni sp. nov. differs from the above‐mentioned species in having the anterior part of the m1 strongly reduced. A cladistic analysis provides evidence that this new Arabian taxon is close to Potwarmus primitivus and Potwarmus sp. nov. from Jebel Zelten (Libya), and also that this genus is unlikely to be the sister taxon to Antemus. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 664–679.     

Miniatures,morphology and molecules: Paedocypris and its phylogenetic position (Teleostei,Cypriniformes)

We review the morphological and molecular evidence that Mayden & Chen recently used to infer that the developmentally truncated fish genus Paedocypris is not a member of the teleost order Cypriniformes or carp‐like fishes, but is ‘the basal sister group to all Cypriniformes’. This hypothesis contradicts several previous studies that used molecular sequence data or morphological characters. A review of the morphological characters that Mayden & Chen discussed and mapped onto their ‘simplified tree’ shows that these, analysed alone, rather support a close relationship of the cyprinids Sundadanio, Danionella, and Paedocypris. We also present four additional analyses of morphological data, which all contradict Mayden & Chen's result. Despite its highly reductive skeleton, posing a serious problem when analysing its phylogenetic position with skeletal characters, the presence in Paedocypris of the basioccipital masticatory plate is compelling evidence that it is a member of the Cyprinoidei (Cyprinidae plus Psilorhynchidae). Our reanalysis and exploration of their molecular sequence data shows that only a single gene, EGR3, of the six nuclear genes analysed by Mayden & Chen, is responsible for the position of Paedocypris as ‘the basal sister group to all Cypriniformes’. Three independent methods to visualize and analyse phylogenetic signal and conflict of data sets (phylogenetic networks, splits analysis methods or SAMS, and site‐wise likelihood analyses) reveal a high level of character conflict and noise in Mayden & Chen's data set. The ‘basal’ position of Paedocypris seems to be the outcome of the interplay of two long‐branch effects. We apply the same analytical methods to the data set from Rüber et al.'s molecular analysis of the phylogenetic position of Paedocypris and discuss our findings. We conclude that none of the molecular data sets compiled to date can establish the phylogenetic position of Paedocypris with confidence. Morphological data suggest that Paedocypris and Danionella are sister genera, and that their closest relative is Sundadanio, although the position of these three miniatures among cyprinoids is still unclear. © 2014 The Linnean Society of London     

Rerooting the rDNA gene tree reveals phoronids to be ‘brachiopods without shells’; dangers of wide taxon samples in metazoan phylogenetics (Phoronida; Brachiopoda)

Molecular phylogenetics has resulted in conflicting accounts of the relationship between phoronids and brachiopods. Taxonomically comprehensive analyses of brachiopod and phoronid ribosomal DNA sequences (rDNAs) rooted with short‐branched mollusc sequences uniformly find that phoronids nest within brachiopods as the sister of the three extant inarticulate lineages. Here, this is called the ‘alternate’ topology because it does not match traditional, morphology‐based ideas. Many other analyses of protein‐coding genes and/or rDNAs place phoronids elsewhere, often as the sister group of all brachiopods, better matching ‘traditional’ ideas. However, these analyses generally are based on data from small selections of brachiopods and phoronids, include data from a wide range of other metazoan taxa, and are rooted with distant outgroups. Here, I show that outgroup rooting of brachiopods and phoronid rDNAs is unreliable, and instead find the root position with procedures that are free from all distortions caused by distantly related taxa, i.e. by Bayesian and maximum likelihood relaxed‐clock analyses of a purely ingroup alignment. All such analyses confirm the ‘alternate’ topology: phoronids belong within the Brachiopoda as the sister group of the inarticulates. In addition, nine factors are identified that (singly or in combination) can cause misreporting of the phylogenetic signal in wide taxon‐range analyses of both rDNA and amino acid sequence data. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012.     

Embryology of Hortonioideae and Monimioideae (Monimiaceae,Laurales): characteristics of the ‘lower’ monimioids

We investigated the embryology of the ‘lower’ monimioids, i.e. Monimioideae (Monimia, Palmeria and Peumus) and Hortonioideae (Hortonia), which are poorly described embryologically. Our results show that, contrary to what has been reported in the literature, ‘lower’ monimioids show very little variation in their embryological characters. Comparisons with Mollinedioideae (a large derived subfamily in Monimiaceae) and other families in Laurales show that the ‘lower’ monimioids are relatively consistent in sharing predominantly isobilateral tetrads of microspores and megaspores, a non‐specialized chalaza, and a mesotestal–endotestal seed coat (with tracheoidal cells of the meso‐ and endotesta). It is likely that, while the shared successive cytokinesis during meiosis of microspore mother cells supports the Monimiaceae–Hernandiaceae–Lauraceae clade obtained by molecular evidence, no synapomorphies exist to support a sister‐group relationship of Monimiaceae with Hernandiaceae or Lauraceae. Instead, the lack of hypostase in ovules and/or young seeds, the lack of endosperm in mature seeds and the amoeboid tapetum in the anther are likely synapomorphies of Hernandiaceae and Lauraceae. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 228–241.     

Ultrastructural organization of the genital tracts in amphigonic females of Euceraphis betulae Koch (Aphididae: Calaphidinae)

The ultrastructure of the genital tracts in amphigonic females of Aphidoidea is described for the first time, using Euceraphis betulae Koch (Aphididae: Calaphidinae) as a representative. The female reproductive apparatus consists of two ovaries, each one with three/four meroistic telotrophic ovarioles; two sac‐like accessory glands lie laterally to a sac‐like seminal receptacle, opening into the dorso‐medial part of the common oviduct by means of a spermathecal duct. A marked secretory activity takes place in the epithelial cells of all the investigated tracts as shown by ultrastructural observation of many organelles involved in this process. No evident golgian area was observed in the cytoplasm of these cells. Extensive smooth endoplasmic reticulum, whose probable role is here discussed, was observed in epithelial cells of the wall of the accessory gland. Spermathecal duct and seminal receptacle had peculiar features that could be related to different secretory activities carried out by these two parts of the spermatheca.     

A newly discovered sperm transport system in the female of Lygaeidae bugs

In the female reproductive system of the relatively large hemipteran, the western conifer seed bug Leptoglossus occidentalis (Heidemann), a cuticle‐lined tube extends medially along the surface of the vagina from the proximal end of the spermathecal complex anteriorly to the base of the common oviduct. This medial tube houses the proximal end of the spermathecal duct, thereby enabling the transport of material from the spermatheca at the distal end of the spermathecal complex, past the vagina (or bursa copulatrix) and directly to the common oviduct. The proximal portion of the spermathecal complex also contains an insemination duct that is separate from the spermathecal duct. The insemination duct allows the male intromittent organ to extend from the vagina to the spermatheca without navigating through the spermathecal duct. The reproductive systems of two previously studied Hemiptera, the milkweed bug Oncopeltus fasciatus (Dallas) and the box elder bug Leptocoris trivittatus (Say), possess a similar cuticle‐lined medial tube housing the spermathecal duct. This new information provides a clearer understanding of sperm transport in the female reproductive system of Lygaeidae bugs, and helps to clarify the path of the male organ during copulation, as well as the movement of sperm during egg laying.     

The interrelationships of Devonian lungfishes (Sarcopterygii: Dipnoi) as inferred from neurocranial evidence and new data from the genus Soederberghia Lehman, 1959

New data are presented on the neurocranial complex (endocranium plus intimately associated bones of the palate) of the Late Devonian (Famennian) ‘rhynchodipterid’ lungfish Soederberghia groenlandica from the Aina Dal Formation (Celsius Bjerg Group) of East Greenland. Only the otic and occipital regions of the braincase are ossified in Soederberghia. The neurocranium of this genus shares a series of derived features with ‘Griphognathus’whitei, including a cranial centrum, fenestrate lateral cristae, and gutters on the ventral surface of the parasphenoid interpreted as accommodating the lateral dorsal aortae. The interrelationships of early lungfishes have been the subject of considerable disagreement. New data from Soederberghia are coupled with a data set focused on the character‐rich neurocranial complex in order to examine the systematic utility of this underexploited morphological system. Different methods of phylogenetic inference (maximum parsimony, Bayesian) return broadly consistent results. The Early to Middle Devonian forms Dipnorhynchus, Stomiahykus and Uranolophus are placed among the most basal of lungfishes. ‘Holodontids’ plus ‘rhynchodipterids’ (comprising Griphognathus and Soederberghia) occupy an apical position, and are separated from the earliest lungfishes by a paraphyletic assemblage of taxa generally identified as ‘chirodipterids’ and ‘dipterids.’ This finds broad agreement with the results of previous cladistic studies focused on non‐neurocranial data sets, but diverges from functional‐adaptive scenarios that posit three lineages of early lungfishes based on aspects of the dentition. As currently defined, both ‘chirodipterids’ and the genus Chirodipterus are heterogeneous assemblages; this analysis fails to find support for the monophyly of either. While this study indicates that Griphognathus is probably paraphyletic, it nevertheless supports a close relationship between the nominal species of this genus and Soederberghia. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151 , 115–171.     

Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)

With about 800 Recent species, ‘miters’ are a widely distributed group of tropical and subtropical gastropods that are most diverse in the Indo‐West Pacific. They include the two families Mitridae and Costellariidae, similar in shell morphology and traditionally treated as close relatives. Some genera of deep‐water Ptychatractidae and Volutomitridae are close to miters in shell morphology, and the term ‘mitriform gastropods’ has been introduced to refer to Mitridae, Costellariidae, and this assortment of convergent forms. The present study aimed at the reconstruction of phylogenetic relationships of mitriform gastropods based on representative taxon sampling. Four genetic markers [cytochrome c oxidase subunit I (COI), 16S and 12S rRNA mitochondrial genes, and H3 (Histone 3) nuclear gene] were sequenced for over 90 species in 20 genera, and the molecular data set was supplemented by studies of radula morphology. Our analysis recovered Mitridae as a monophyletic group, whereas the genus Mitra was found to be polyphyletic. Of 42 mitrid species included in the analysis, 37 formed a well‐supported ‘core Mitridae’ consisting of four major clades, three of them consistent with the subfamilies Cylindromitrinae, Imbricariinae, and Mitrinae, and Strigatella paupercula standing out by itself. Basal to the ‘core Mitridae’ are four minor lineages, with the genus Charitodoron recognized as sister group to all other Mitridae. The deep‐water family Pyramimitridae shows a sister relationship to the Mitridae, with high support for a Pyramimitridae + Mitridae clade. Our results recover the monophyly of the Costellariidae, which form a well‐supported clade that also includes Ptychatractidae, Columbariinae, and Volutomitridae, but not Mitridae. Most derived and diverse amongst Costellariidae are species of Vexillum, characterized by a bow‐shaped, multicuspidate rachidian tooth. Several previously unrecognized deep‐water costellariid lineages are revealed. Their members retain some plesiomorphies – in particular a tricuspidate rachidian tooth – that makes them morphologically intermediate between ptychatractids and Vexillum. The taxa of Ptychatractidae included in the analysis are not monophyletic, but form three well‐supported, unrelated groupings, corresponding respectively to Ceratoxancus + Latiromitra, Exilia, and Exiliodea. None of them shows an affinity to Pseudolividae. © 2015 The Linnean Society of London     

Patterns of morphological evolution in the mandible of the house mouse Mus musculus (Rodentia: Muridae)

The worldwide distributed house mouse, Mus musculus, is subdivided into at least three lineages, Mus musculus musculus, Mus musculus domesticus, and Mus musculus castaneus. The subspecies occur parapatrically in a region considered to be the cradle of the species in Southern Asia (‘central region’), as well as in the rest of the world (‘peripheral region’). The morphological evolution of this species in a phylogeographical context is studied using a landmark‐based approach on mandible morphology of different populations of the three lineages. The morphological variation increases from central to peripheral regions at the population and subspecific levels, confirming a centrifugal sub‐speciation within this species. Furthermore, the outgroup comparison with sister species suggests that M. musculus musculus and populations of all subspecies inhabiting the Iranian plateau have retained a more ancestral mandible morphology, suggesting that this region may represent one of the relevant places of the origin of the species. Mus musculus castaneus, both from central and peripheral regions, is morphologically the most variable and divergent subspecies. Finally, the results obtained in the present study suggest that the independent evolution to commensalism in the three lineages is not accompanied by a convergence detectable on jaw morphology. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 635–647.     

Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages

The phylogenetic relationships and generic assignments of ‘Ochlerotatus’ and related taxa of uncertain taxonomic position in the classification of Aedini previously proposed by the authors in 2004 and 2006 are explored using 297 characters from eggs, fourth‐instar larvae, pupae, adults and immature habitat coded for 158 exemplar species. The ingroup comprises 54 species and the outgroup includes four non‐aedine species and 100 aedine species, 21 of which were previously classified as incertae sedis. Data are analysed in a total‐evidence approach using implied weighting. The analysis produced 158 most parsimonious cladograms. The strict consensus tree (SCT) corroborates the monophyly of the 30 generic‐level taxa recognized previously that are included in the analysis. Overall, the results show remarkable congruence with those obtained previously despite differences in the taxa and morphological characters analysed in this and the two previous studies. All species of Ochlerotatus s.s., subgenus ‘Ochlerotatus’sensu auctorum, Geoskusea, Levua, Pseudoskusea and Rhinoskusea included in the analysis fall within a single clade that is treated as genus Ochlerotatus; thus, the last four taxa are restored to their previous subgeneric rank within this genus. Nine additional subgenera, of which four are new, are proposed for monophyletic clades of Ochlerotatus species based on the strength of character support and application of the principle of equivalent rank. Acartomyia stat. nov. , Culicelsa stat. nov. , Gilesia stat. nov. , Protoculex stat. nov. and Chrysoconops stat. nov. are resurrected from synonymy with Ochlerotatus; and Empihals subgen. nov. (type species: Culex vigilax Skuse), Pholeomyia subgen. nov. (type species: Aedes calcariae Marks), Buvirilia subgen. nov. (type species: Aedes edgari Stone & Rosen) and Sallumia subgen. nov. (type species: Aedes hortator Dyar & Knab) are described as new. The sister group of Ochlerotatus includes a number of species that were previously regarded as incertae sedis in ‘Oc. (Finlaya)’ and ‘Oc. (Protomacleaya)’. Based on previous observations, refined relationships and new character support, three additional genera are recognized for species previously included in ‘Finlaya’, i.e. Danielsia stat. nov . (type species: Danielsia albotaeniata Leicester), Luius gen. nov. (type species: Aedes fengi Edwards) and Hopkinsius gen. nov. (type species: Aedes ingrami Edwards). Additionally, Alloeomyia subgen. nov. (type species: Culex pseudotaeniatus Giles) and Yamada subgen. nov. (type species: Aedes seoulensis Yamada) are introduced as subgenera of Collessius and Hopkinsius, respectively. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 29–114.     

Kauri seeds and larval somersaults: The larval trunk of the seed mining basal moth Agathiphaga vitensis (lepidoptera: Agathiphagidae)

The trunk morphology of the larvae of the kauri pine (Agathis) seed infesting moth Agathiphaga is described using conventional, polarization, and scanning electron microscopy. The pine seed chamber formed by the larva is also described and commented on. The simple larval chaetotaxy includes more of the minute posture sensing setae, proprioceptors, than expected from the lepidopteran larval ground plan. The excess of proprioceptors is suggested to be necessary for sensory input concerning the larval posture within the seed chamber. The trunk musculature includes an autapomorphic radial ventral musculature made up of unique multisegmental muscles. The combined presence of additional proprioceptors and the unique ventral musculature is proposed to be related to the larval movement within the confined space of the seed chamber, especially to a proposed somersault movement that allows the larva to orientate itself within the chamber. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

The fine structure of the female reproductive system of Zorotypus caudelli Karny (Zoraptera)

The general structure of the female genital system of Zorotypus caudelli is described. The ovarioles are of the panoistic type. Due to the reduction of the envelope (tunica externa) the ovarioles are in direct contact with the hemolymph like in some other insect groups, Plecoptera included. The calices are much larger in Z. caudelli then in Zorotypus hubbardi and their epithelial cells produce large amounts of secretions, probably protecting the surface of the eggs deposited on the substrate. Eggs taken from the calyx bear a series of long fringes, which are missing in the eggs found in the ovariole, and in other zorapteran species. The long sperm of Z. caudelli and the long spermathecal duct are likely related to a sexual isolating mechanism (cryptic female choice), impeding female re-mating. The apical receptacle and the spermathecal duct - both of ectodermal origin - consist of three cell types. In addition to the cells beneath the cuticle lining the lumen, two other cell types are visible: secretory and canal cells. The cytoplasm of the former is rich in rough endoplasmic reticulum cisterns and Golgi complexes, which produce numerous discrete dense secretory bodies. These products are released into the receiving canal crossing the extracellular cavity of secretory cells, extending over a series of long microvilli. The secretion is transported towards the lumen of the apical receptacle of the spermatheca or to that of the spermathecal duct by a connecting canal formed by the canal cells. It is enriched by material produced by the slender canal cells. Before mating, the sperm cells are enveloped by a thick glycocalyx produced at the level of the male accessory glands, but it is absent when they have reached the apical receptacle, and also in the spermathecal duct lumen. It is likely removed by secretions of the spermatheca. The eggs are fertilized at the level of the common oviduct where the spermathecal duct opens. Two micropyles at the dorsal side of the equator level possibly facilitate fertilization. The presence of these two micropyles is a presumably derived feature shared with Phasmatodea. The fine structure of the female reproductive system of Z. caudelli does not allow to assess the phylogenetic position at the present stage of knowledge. The enlarged calyx and the temporary presence of long fringes on the eggs are potential autapomorphies of Z. caudelli or may indicate relationships with other Zorotypus species.     

Evolutionary trends on extant cat skull morphology (Carnivora: Felidae): a three‐dimensional geometrical approach

The skulls of 33 extant cat species were characterized through three‐dimensional geometric morphometrics using 20 landmarks. A principal component analysis (PCA) was performed with Procrustes fitted coordinates, and the PC‐scores were phylogenetically corrected by independent contrasts method. Three PCs allowed for the definition of five cat skull patterns. PC1: ‘snouted/massive‐headed cats’ (genus Panthera) opposing the ‘round‐headed small cats’ (genus Oncifelis, Prionailurus rubiginosus, Prionailurus bengalensis, among other small cats); PC2: ‘tapering‐headed cats’ (Neofelis nebulosa, Herpailurus yagouaroundi, Prionailurus planiceps) opposing the ‘stout‐headed cats’ (Acinonyx jubatus, Uncia uncia, Otocolobus manul, Felis margarita, and Felis nigripes); and PC3: ‘low profiled‐headed cats’ (mostly, Pr. planiceps). A sixth pattern, the ‘generalized skull’, observed in the Caracal lineage, genus Lynx, Leopardus pardalis, and Catopuma temminckii, indicates a morphological convergence among midsized‐cats. The morphological trends ‘snouted/massive’ and ‘round’ clearly denote a co‐evolution between size and shape. The other skull patterns evolved unrelatedly to the size (i.e. their allometric variations are not a size function). Nevertheless, each species comprises an amalgam of these patterns, so the influence of the size permeates, in some extent, the skull morphology along all cat lineages. The felid ecomorphological fit to hypercarnivory is obvious; however, different skull shapes in same‐sized species with similar habits, indicate that the variation in the skull morphology may result from phenotypic fluctuations, whose adaptive value (if indeed there is any) is still obscure. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 176–190.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

When rare species become endangered: cryptic speciation in myrmecophilous hoverflies

The myrmecophilous hoverfly, Microdon mutabilis, is listed as a ‘Rare’ or ‘Nationally Notable Species’ in UK Red Data Books. As an obligate social parasite, feeding only from ant colonies, its life‐style satisfies theoretical conditions under which cryptic speciation is predicted to evolve; namely, strong selection for nonmorphological adaptations that enhance its exploitation of a local subspecies or populations of its host. Samples of larvae and pupae in Ireland, Scotland and England showed that M. mutabilis exploits a single and different host ant species on different sites across its range. In nine southern English colonies, 95.6% of infested nests were of Myrmica scabrinodis whereas in six Irish and two Scottish colonies 100% and 94.2%, respectively, of the infested nests were of Formica lemani, despite M. scabrinodis being common at all sites. Although the adults from ‘scabrinodis’ (and lemani) populations are cryptic, morphometric measurements of pupae showed consistent diagnostic characters that were sufficiently distinct for these ecotypes to be classed as separate species. We conclude that M. mutabilis is the ‘lemani‐type’ and designate the ‘scabrinodis‐type’ as a new species, Microdon myrmicae spec. nov. Thus, one of the listed threatened species of the British Isles becomes two species, each possessing about half the number of populations and occupying half the range of the original ‘species’. Each also inhabits a different serai stage within grassland or heathland, and will require a different management regime if its declining populations are to be conserved. ‘M. mutabilis’ is reported with other host ant species on the European continent. In the light of our results, these may prove to be additional cryptic species. We suggest that cryptic speciation is apt to evolve in species, such as myrmecophiles, endoparasites and koinobiont parasitoids, whose life‐styles result in strong selection on their physiological or behavioural characters. The implications for Red Data Book classifications and for practical conservation are discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75 , 291–300.     

Coding characters from different life stages for phylogenetic reconstruction: a case study on dragonfly adults and larvae,including a description of the larval head anatomy of Epiophlebia superstes (Odonata: Epiophlebiidae)

The exclusive use of characters coding for specific life stages may bias tree reconstruction. If characters from several life stages are coded, the type of coding becomes important. Here, we simulate the influence on tree reconstruction of morphological characters of Odonata larvae incorporated into a data matrix based on the adult body under different coding schemes. For testing purposes, our analysis is focused on a well‐supported hypothesis: the relationships of the suborders Zygoptera, ‘Anisozygoptera’, and Anisoptera. We studied the cephalic morphology of Epiophlebia, a key taxon among Odonata, and compared it with representatives of Zygoptera and Anisoptera in order to complement the data matrix. Odonate larvae are characterized by a peculiar morphology, such as the specific head form, mouthpart configuration, ridge configuration, cephalic musculature, and leg and gill morphology. Four coding strategies were used to incorporate the larval data: artificial coding (AC), treating larvae as independent terminal taxa; non‐multistate coding (NMC), preferring the adult life stage; multistate coding (MC); and coding larval and adult characters separately (SC) within the same taxon. As expected, larvae are ‘monophyletic’ in the AC strategy, but with anisopteran and zygopteran larvae as sister groups. Excluding larvae in the NMC approach leads to strong support for both monophyletic Odonata and Epiprocta, whereas MC erodes phylogenetic signal completely. This is an obvious result of the larval morphology leading to many multistate characters. SC results in the strongest support for Odonata, and Epiprocta receives the same support as with NMC. Our results show the deleterious effects of larval morphology on tree reconstruction when multistate coding is applied. Coding larval characters separately is still the best approach in a phylogenetic framework. © 2015 The Linnean Society of London     

Evolution of sperm morphology in potamid freshwater crabs (Crustacea: Brachyura: Potamoidea)

We investigated sperm cells and spermatophores of four species of Old World freshwater crabs belonging to three different genera of the subfamily Potaminae (family Potamidae). Characters previously believed to be apomorphic for the potamid subfamily Potamiscinae were also found to occur in the Potaminae. To infer the morphological ancestral character state combination of the Potamidae, ancestral character state analysis of four different sperm traits was performed, based on a 16S rDNA phylogeny of the investigated species. Comparing molecular phylogeny and character state distribution, several cases of convergent evolution could be identified. The densely packed, coenospermic spermatophores and the occurrence of a ‘tongue‐and‐groove’ connection between operculum and acrosomal zones are probably apomorphies for the whole Potamidae. The spermatozoa of Socotrapotamon socotrense show several unique characters. We also analysed the evolution of acrosome size. The sperm cells of the Potamidae and their sister‐group Gecarcinucidae only slightly overlap in acrosome size. Within the investigated species, the ‘East Asia’ subclade (subfamily Potamiscinae) developed significantly larger acrosomes than the subfamily Potaminae. Our results suggest that the use of brachyuran acrosome morphology for phylogenetic inference at the family level is strongly affected by small sample size, and by convergent character evolution. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Phylogenetics and taxonomy of the New World leafy spurges,Euphorbia section Tithymalus (Euphorbiaceae)

The 480 species of leafy spurges, Euphorbia subgenus Esula, represent the main temperate radiation in the large genus Euphorbia. This group is distributed primarily in temperate Eurasia, but with smaller, disjunct centres of diversity in the mountains of the Old World tropics, in temperate southern Africa and in the New World. The majority of New World diversity (32 species) occurs in a single section, section Tithymalus. We analysed sequences of the nrITS and plastid ndhF, trnH‐psbA, trnS‐trnG and trnD‐trnT regions to reconstruct the phylogeny of section Tithymalus and to examine the origins and diversification of the species native to the New World. Our results indicate that the New World species of section Tithymalus form a clade that is sister to the widespread, weedy E. peplus. The New World species fall into two primary groups: a ‘northern annual clade’ from eastern North America and a diverse clade of both annual and perennial species that is divided into three subgroups. Within the second group, there is a small ‘southern annual clade’ from Texas and northern Mexico, a perennial ‘Brachycera clade’ from the western United States and northern Mexico, and a perennial ‘Esuliformis clade’ from montane areas of Mexico, Guatemala, Honduras and the Caribbean island of Hispaniola. Ancestral state reconstructions indicate that the annual habit probably evolved in the ancestor of E. peplus and the New World clade, with a subsequent reversal to the perennial habit. In conjunction with this phylogenetic framework, the New World species of section Tithymalus are comprehensively reviewed. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 191–228.     

Sperm storage by spermatodoses in the spermatheca of Trioza alacris (Flor, 1861) hemiptera,psylloidea, triozidae: A structural and ultrastructural study

Female insects generally store sperm received during mating in specific organs of their reproductive tract, i.e., the spermathecae, which keep the sperm alive for a long time until fertilization occurs. We investigated spermatheca morphology and ultrastructure in the psylloidean insect Trioza alacris (Flor, 1861 ) in which spheroidal sperm packets that we refer to as ‘spermatodoses’ are found after mating. The ectoderm‐derived epithelium of the sac‐shaped spermatheca that has a proximal neck, consists of large secretory and flat cuticle‐forming cells. Secretory cells are characterized by a wide extracellular cavity, bordered by microvilli, in which electron‐dense secretion accumulates before discharge into the spermathecal lumen. The cuticle‐forming cells produce the cuticular intima of the organ and a peculiar specialized apical structure, through which secretion flows into the lumen. At mating, the male transfers bundles of sperm cells embedded in seminal fluid into the spermathecal neck. Sperm cells proceed towards the spermathecal sac lumen, where they are progressively compacted and surrounded with an envelope that also encloses secretions of both male and female origin. We describe the formation of these sperm containing structures and document the contribution of the female secretion to spermatodose or female‐determined spermatophore construction. We also discuss the choice of the term ‘spermatodose’ for T. alacris and suggest it be used to refer to sperm masses constructed in the female reproductive organs, at least when they involve the contribution of female secretion. © 2011 Wiley Periodicals, Inc.