First description of larvae of Radulinopsis derzhavini Soldatov et Lindberg, 1930 (Scorpaeniformes: Psychrolutidae) with remarks on melanin colouration and relationships with related species

The Derzhavin's sculpin (Radulinopsis derzhavini Soldatov et Lindberg, 1930) is a psychrolutids species that leads a cryptic life in coastal waters of the Japan Sea and the southern Okhotsk Sea. To date, larvae of this species have remained unknown; therefore, their biology is poorly understood. In the present study, the early developmental stages of R. derzhavini are described for the first time. In Peter the Great Bay, Japan Sea, this species is characterized by a spring spawning season (May) and a short pelagic period of larval development, usually from mid-May to the last 10 days of June. Species identification of the described larvae was confirmed by incubation, rearing of larvae in captivity, and genotyping of a fragment of the mitochondrial cytochrome c oxidase I (COI) gene. In addition, the development of pigmentation in larvae of this and related species was compared. Morphological analysis of both adults and larvae, together with complementary molecular genetics, confirms the previously obtained conclusions that the species of the genera Radulinopsis, Radulinus, Asemichthys, and Astrocottus form a natural group with monophyly by all types of data.     

PHYLOGENY OF THE GENUS ROSTANGA (NUDIBRANCHIA), WITH DESCRIPTIONS OF THREE NEW SPECIES FROM SOUTH AFRICA

Rostanga elandsia sp. nov., Rostanga aureamala sp. nov. andRostanga phepha sp. nov. are characterized by having the radulawith slender innermost lateral teeth, which lack denticles onthe inner side of the cusp and have a single denticle on theouter side. The outermost lateral teeth of these three speciesare elongate, but shorter than in other species of the genus.In addition, R. aureamala is the only species of the genus withrachidian teeth and R. phepha is unique within the genus Rostanga byvirtue of its white coloration with dark spots. A phylogenetic analysis shows that the three new species fromSouth Africa and Rostanga setidens (Odhner, 1939) are the sistergroup of the rest of the genus. The species from Japan and MarshallIslands (North Pacific Ocean) are basal in the sister cladecontaining the other species of Rostanga Bergh, 1879. The tropicalIndo-Pacific species of Rostanga are not monophyletic. The Atlanticand Eastern Pacific species form a monophyletic, derived clade,being the sister group of Rostanga australis Rudman & Avern,1989, which has a narrow range restricted to south eastern Australia.The widespread Indo-Pacific species Rostanga bifurcata Rudman& Avern, 1989, is the sister group of Rostanga dentacus Rudman& Avern, 1989, also widesprad in the tropical western Pacific. This phylogeny suggest s a viariant origin of the Sourth African, Atlantic-EasternPacific, and probably North Pacific species, whereas in thetropical Indo-Pacific most sister speceis are sympatric. (Received 16 May 1999; accepted 31 July 2000)     

Historical transoceanic dispersal of a freshwater shrimp: the colonization of the South Pacific by the genus Paratya (Atyidae)

Aim To infer the phylogenetic relationships within the freshwater shrimp genus Paratya Miers, 1882 (Atyidae) and to use these data to answer biogeographical questions about the location, timing and form of evolution of this genus in the South Pacific. Location Paratya are spread throughout various freshwater habitats in the western Pacific, with a disjunct northern range in the North Pacific (Japan, Korea, Ryukyu Islands, Siberia) and South Pacific (Australia, New Zealand, New Caledonia, Lord Howe, Norfolk Island). Methods Specimens were obtained from throughout its range. Mitochondrial sequences of cytochrome oxidase subunit I and 16S ribosomal DNA were analysed using phylogenetic techniques to identify whether landmasses are monophyletic and what the relationships are between landmasses. Molecular clock dating methods were used to date divergences between taxa. Results Each landmass was recovered as monophyletic. Japan/Ryukyu Islands is the most basal group, followed by New Zealand. Australian specimens form a sister group to a clade made up of two groups (New Caledonia and Lord Howe/Norfolk Island). The oldest divergence within the genus (between North and South Pacific) took place 12–19 Ma. Main conclusions The geographical origin of the genus (either Gondwana or Laurasia) is unclear. Dispersal occurred between the North and South Pacific long after the split up of Gondwana. Dispersal likely explains the presence of Paratya on each landmass in the South Pacific, from continent to isolated oceanic island. This dispersal is conjectured to have taken place through oceanic currents because of the amphidromous life cycle of some taxa of Paratya, given that amphyidromy is plesiomorphic in atyid shrimp.     

The gastropod–crustacean connection: towards the phylogeny and evolution of the parasitic copepod family Splanchnotrophidae

Amongst the most significant metazoan taxa associated with gastropod molluscs is the endoparasitic copepod family Splanchnotrophidae. Currently it contains five genera with highly modified morphology and exclusively infesting nudibranch and sacoglossan sea slug hosts. The present study is a first approach towards reconstructing their phylogeny and evolution. Cladistic analysis of 109 morphological characters including 24 known splanchnotrophid species resulted in a fully resolved strict consensus tree that is discussed in morphological, functional, and geographical frameworks. Alternative topologies are also explored. Originating from paraphyletic Philoblennidae, the Splanchnotrophidae emerge as sister group to the genus Briarella. Unique synapomorphies, such as the bizarre body shapes and successive reduction of mouthparts, are discussed as adaptive traits to endoparasitism that evolved only once within copepods infesting shell‐less heterobranch gastropods. The ancestrally Indo‐Pacific Splanchnotrophidae split up into a clade of the still Indo‐Pacific genera Ceratosomicola and Arthurius, sister to a clade composed of the monophyletic amphi‐American genus Ismaila and European Splanchnotrophus emerging from paraphyletic Lomanoticola. Although initial radiation of Briarella and Splanchnotrophidae is likely to have involved chromodoridid nudibranch hosts, later phylogenies of parasites and their hosts are incongruent; intriguingly, host shifts from nudibranch to only distantly related sacoglossan species occurred at least two times independently. Such remarkable ecological plasticity is assumed to have driven splanchnotrophid diversification. Topological hypotheses and historical biogeographical and evolutionary scenarios inferred herein can be tested by future molecular research. © 2013 The Linnean Society of London     

Phylogenetic evidence for loss of sound production and a shift in sexual recognition signals in Hamadryas butterflies (Nymphalidae: Biblidinae)

The neotropical butterfly genus Hamadryas Hübner comprises 20 species that exhibit an intriguing variation in their natural history traits. Although revised in 1983, no phylogenetic hypothesis was presented: the first phylogenetic hypothesis is estimated here based on 93 characters and including species from the three other genera in the tribe Ageroniini. The phylogeny is used to test the monophyly of the genus, establish the sister group of Hamadryas and identify its apomorphies. The tree allows the inference of patterns of character change in sound production and sexual dimorphism. Implied weights show that Hamadryas is monophyletic and corroborate Ectima Doubleday as a sister genus. Previously suggested subgenera for Hamadryas were non‐monophyletic, with the exception of the laodamia clade, supported by the presence of a complete sterigma. Sound production is inferred to be a derived condition in Hamadryas that has been lost in the laodamia clade. This, plus the presence of androconial organs and sexual dimorphism in the laodamia clade, suggests a shift in sexual recognition signalling. Furthermore, the phylogeny indicates that the colour pattern of males in the laodamia clade is novel, supporting a Darwinian origin of sexual dimorphism.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.     

Molecular systematics of social skinks: phylogeny and taxonomy of the Egernia group (Reptilia: Scincidae)

The lizards of the Egernia group of Australia and Melanesia include some of the most distinctive members of the family Scincidae in morphology (including giant size, spinose scalation), ecology and behaviour. Social behaviour, including long‐term recognition of individuals and kin, mate fidelity and home site fidelity, is amongst the most complex known in squamate reptiles and is the subject of an expanding number of studies. Lack of a sound phylogeny for the Egernia group has limited our ability to understand the evolution and patterns of variation in social behaviour within this group, and evidence for the monophyly of the largest genus, Egernia (64% of the species), has been lacking. We present data derived from nucleotide sequences that establish a phylogenetic framework for the Egernia group. We used two mitochondrial sequences, the protein‐encoding ND4 gene and a ribosomal gene, 12s rRNA, and two nuclear sequences, the protein‐encoding c‐mos, and non‐encoding intron 7 of β‐fibrinogen. Our phylogenetic analyses show that Corucia of the Solomon Islands is the sister group of the rest of the Egernia group. The genus Egernia is paraphyletic, including four well‐supported monophyletic units, one of which is the sister lineage of the Tiliqua lineage (Tiliqua plus Cyclodomorphus). We suggest a revised taxonomic scheme that recognizes the major monophyletic lineages in Egernia (s.l.) as distinct genera. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 781–794.     

Megachiropteran Evolution Studied with 12S rDNA and c-mos DNA Sequences

Recent studies of mitochondrial DNA sequences have indicated the requirement for substantial revisions of the morphological understanding of the phylogeny of Megachiroptera (Pteropodidae). There is disagreement between studies as to what these revisions might be. This investigation was undertaken to expand the number of studied species and to add the first data from a nuclear gene sequence. For 12S ribosomal DNA (aligned length of 405 positions), 75 Megachiroptera (50 species in 20 genera) and two outgroup species were sequenced. For the oncogene c-mos (aligned length of 488 bases), 56 Megachiroptera (42 species in 19 genera) were sequenced and three eutherians from GenBank used as outgroups.The root of the megachiropteran phylogeny cannot be determined with the present data. Nyctimene, the only studied insectivorous genus (Paranyctimene not being included), plus Notopteris, the only long-tailed megachiropteran, form the sister clade to the other genera in combined analyses. Several alternative rootings are not rejected by the data, suggesting a rapid early radiation. Generic distributions indicate that this may have occurred in Melanesia. The results confirm that the subfamily Macroglossinae is not monophyletic with the long tongued phenoptype arising at least twice and support the existence of a major clade including a monophyletic endemic African component and biogeographically neighboring genera such as Rousettus and Eonycteris. The phylogenetic position of one African genus, Eidolon, remains uncertain.A cynopterine section (excluding Nyctimene and Myonycteris) is supported, albeit weakly, as a monophyletic group. Pteropus and the related, possibly polyphyletic genus Pteralopex, are unexpectedly basal compared to previous molecular studies.     

Biogeography and diversification of the Pacific ant genus Lordomyrma Emery

Aim This study addresses the origins of terrestrial biodiversity of the Fijian islands using the ant genus Lordomyrma (Hymenoptera: Formicidae: Myrmicinae) as a model system. We derive the evolution of the genus and determine its closest extra-Fijian relatives from geological data, molecular phylogenetic reconstruction and divergence estimates. Location Ant taxa were sampled in the Southwest Pacific, Melanesia, Southeast Asia, Australia and mainland China. Methods Phylogeny and divergence estimates of the ant genus Lordomyrma based on four nuclear genes (28S, ArgK, LW Rh, CAD) plus data on Indo-Pacific geological history are used to address current hypotheses regarding the origins of the Fijian biota. Results The genus Lordomyrma probably originated in mainland Asia, with subsequent colonization of Australia and the Pacific. The Fijian Lordomyrma clade is monophyletic, and originated c. 8.8 Ma, when it diverged from a sister group in Papua New Guinea. Main conclusions The colonization of Fiji by Lordomyrma is probably a result of long-distance dispersal from New Guinea, possibly aided by island hopping across the Vitiaz Arc. The timeline of diversification in Lordomyrma is broadly congruent with the Miocene fragmentation of the Vitiaz Arc and the Pliocene emergence of Vanua Levu. The biotic shuttle hypothesis, which posits ‘Eua Island as the source of Fijian endemics, is rejected based on the sister relationship of Fiji and New Guinea lineages, as well as on the Miocene submergence of the terrane below sea level. The diversity of Fijian Lordomyrma results from the radiation of a single lineage, which diverged from a New Guinea sister group. The genus appears to have originated in Asia rather than in Australia.     

Physical gills in Elmidae (Coleoptera: Byrrhoidea): Structure and evolutionary pattern of plastron in Stenelmis and related genera

Many species within Elmidae (Coleoptera: Byrrhoidea) have plastrons composed of flattened setae. However, some genera display fine plastrons on the epicuticle, called plastron hairs. In Japanese elmids, members of the genera Stenelmis, Ordobrevia, Nomuraelmis and Leptelmis bear ventral plastron hairs. Based on a maximum likelihood tree including most Japanese genera within Elmidae, we found that these genera are monophyletic and that plastron hairs are a derived character in Elmidae. We also found that the genus Graphelmis bears jigsaw puzzle‐like plastron scales with plastron hair‐like projections, and is sister to the group with plastron hairs.     

Molecular systematics of threadfin breams and relatives (Teleostei,Nemipteridae)

Threadfin breams and relatives of the family Nemipteridae comprise 69 currently recognized species in five genera. They are found in the tropical and subtropical Indo‐West Pacific and most are commercially important. Using recently developed molecule‐based approaches exploiting DNA sequence variation among species/specimens, this study reconstructed a comprehensive phylogeny of the Nemipteridae, examined the validity of species and explored the cryptic diversity of the family, and tested previous phylogenetic hypotheses. A combined data set (105 taxa from 41 morphospecies) with newly determined sequences from two nuclear genes (RAG1 and RH) and one mitochondrial gene (COI), and a data set with only COI gene sequences (329 newly obtained plus 328 from public databases from a total of 53 morphospecies) were used in the phylogenetic analysis. The latter was further used for species delimitation analyses with two different tools to explore species diversity. Our phylogenetic results showed that all the currently recognized genera were monophyletic. The monotypic genus Scaevius is the sister group of Pentapodus and they together are sister to Nemipterus. These three genera combined to form the sister group of the clade comprising Parascolopsis and Scolopsis. The validity of most of the examined species was confirmed except in some cases. The combined evidence from the results of different analyses revealed a gap in our existing knowledge of species diversity in the Nemipteridae. We found several currently recognized species contain multiple separately evolving metapopulation lineages within species; some lineages should be considered as new species for further assignment. Finally, some problematic sequences deposited in public databases (probably due to misidentification) were also revised in this study to improve the accuracy for prospective DNA barcoding work on nemipterid fishes.     

Molecular phylogenetics of tribe Synandreae, a North American lineage of lamioid mints (Lamiaceae)

The five mint genera Brazoria, Macbridea, Physostegia, Synandra and Warnockia (Lamioideae: Lamiaceae) are all North American endemics. Together with the monotypic European genus Melittis and the Asian genus Chelonopsis, these taxa have been classified as subtribe Melittidinae. Previous morphological studies have failed to uncover synapomorphic characters for this group. We sequenced the plastid trnL‐trnF region and trnS‐trnG spacer and the nuclear ribosomal 5S non‐transcribed spacer (5S‐NTS) to assess phylogenetic relationships within Melittidinae. Standard parsimony and direct optimization (POY) analyses show Melittis, the type genus of the subtribe, as sister to Stachys. Thus, the monophyly of subtribe Melittidinae is not supported either by molecular or morphological data. However, the North American endemics form a monophyletic group that can be recognized as the recircumscribed tribe Synandreae. The molecular relationships among these genera are corroborated by both morphological and cytological data. The expected close relationship between the south‐central endemics Warnockia and Brazoria and their sister relationship to the widespread genus Physostegia is confirmed. Nevertheless, most of the North American endemics are restricted to the south‐east of the continent. Dispersal westwards and northwards is correlated with an increase in chromosome numbers. No specific Eurasian origin (i.e., transatlantic or transpacific) can be determined, but Synandreae are clearly distinct from the large Stachys clade, and therefore represent a separate migration into North America. © The Willi Hennig Society 2007.     

The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

Phylogenetic relationships of the genus Hoplia Illiger (Scarabaeidae: Hopliinae)

Results of phylogenetic analysis based on 34 morphological characters of 24 species of 11 genera of Hopliinae from Europe, Japan, South Africa, Madagascar, North and Central America, indicates that the genus Hoplia is a monophyletic group with species distributed in Europe, Japan and America. Based in this analysis the Asiatic genus Ectinohoplia is the closest relative of the genus Hoplia, and the South American genus Barybas (Melolonthinae: Macrodactylini) is the sister group of Hopliinae.     

Phylogeny of Castanea (Fagaceae) based on chloroplast trnT-L-F sequence data

Species in the genus Castanea are widely distributed in the deciduous forests of the Northern Hemisphere from Asia to Europe and North America. They show floristic similarity but differences in chestnut blight resistance especially among eastern Asian and eastern North American species. Phylogenetic analyses were conducted in this study using sequences of three chloroplast noncoding trnT-L-F regions. The trnT-L region was found to be the most variable and informative region. The highest proportion of parsimony informative sites, more and larger indels, and higher pairwise distances between taxa were obtained at trnT-L than at the other two regions. The high A+T values (74.5%) in the Castanea trnT-L region may explain the high proportion of transversions found in this region where as comparatively lower A+T values were found in the trnL intron (68.35%) and trnL-F spacer (70.07%) with relatively balanced numbers of transitions and transversions. The genus Castanea is supported as a monophyletic clade, while the section Eucastanon is paraphyletic. C. crenata is the most basal clade and sister to the remainder of the genus. The three Chinese species of Castanea are supported as a single monophyletic clade, whose sister group contains the North American and European species. There is consistent but weak support for a sister–group relationship between the North American species and European species.     

Phylogenetic relationships within the South American fish family Anostomidae (Teleostei,Ostariophysi, Characiformes)

Analysis of a morphological dataset containing 152 parsimony‐informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov. , described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synaptolaemus. Rhytiodus and Schizodon together formed a well‐supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis‐ and trans‐Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre‐dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 70–210.     

Phylogeny of western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae)

Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty‐nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty‐eight terminal taxa, twenty‐four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para‐ or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.     

MULTIGENE ANALYSES OF PHOTOSYNTHETIC EUGLENOIDS AND NEW FAMILY,PHACACEAE (EUGLENALES)1

Bayesian and maximum‐likelihood (ML) analyses of the combined multigene data (nuclear SSU rDNA, and plastid SSU and LSU rDNA) were conducted to evaluate the phylogeny of photosynthetic euglenoids. The combined data set consisted of 108 strains of photosynthetic euglenoids including a colorless sister taxon. Bayesian and ML analyses recovered trees of almost identical topology. The results indicated that photosynthetic euglenoids were divided into two major clades, the Euglenaceae clade (Euglena, Euglenaria, Trachelomonas, Strombomonas, Monomorphina, Cryptoglena, Colacium) and the Phacaceae clade (Phacus, Lepocinclis, Discoplastis). The Euglenaceae clade was monophyletic with high support and subdivided into four main clades: the Colacium, the Strombomonas and Trachelomonas, the Cryptoglena and Monomorphina, and the Euglena and Euglenaria clades. The genus Colacium was positioned at the base of the Euglenaceae and was well supported as a monophyletic lineage. The loricate genera (Strombomonas and Trachelomonas) were located at the middle of the Euglenaceae clade and formed a robust monophyletic lineage. The genera Cryptoglena and Monomorphina also formed a well‐supported monophyletic clade. Euglena and the recently erected genus Euglenaria emerged as sister groups. However, Euglena proxima branched off at the base of the Euglenaceae. The Phacaceae clade was also a monophyletic group with high support values and subdivided into three clades, the Discoplastis, Phacus, and Lepocinclis clades. The genus Discoplastis branched first, and then Phacus and Lepocinclis emerged as sister groups. These genera shared a common characteristic, numerous small discoid chloroplasts without pyrenoids. These results clearly separated the Phacaceae clade from the Euglenaceae clade. Therefore, we propose to limit the family Euglenaceae to the members of the Euglena clade and erect a new family, the Phacaceae, to house the genera Phacus, Lepocinclis, and Discoplastis.     

Polyphyly of Caddoidea,reinstatement of the family Acropsopilionidae in Dyspnoi,and a revised classification system of Palpatores (Arachnida,Opiliones)

Among the least studied harvestmen are the members of the family Caddidae sensu Shear, 1975 , a group of Opiliones with massive eyes and the putative sister group of the remaining Eupnoi. Caddids were originally described as two families, Caddidae and Acropsopilionidae, but these are currently treated as subfamilies of Caddidae. These minute arachnids are rarely collected and present some interesting biogeographical patterns, including a disjunct distribution between East Asia and eastern North America, and some of the few cases of trans‐Pacific genera in southern hemisphere Opiliones. We therefore obtained samples from most of the landmasses inhabited by Caddidae and undertook a phylogenetic study using nuclear and mitochondrial genes for as many samples as possible. Our results, based on a broad taxonomic sampling, surprisingly showed polyphyly of Caddidae, with the genus Caddo forming the sister group of the remaining Eupnoi, whereas the southern hemisphere genera, many of which were originally placed in Acropsopilionidae, within Dyspnoi, formed the sister clade of the remaining Dyspnoi. In addition, the more recently described genus Hesperopilio, from Western Australia and Chile, was unrelated to either Caddidae or Acropsopilionidae, despite having the supposedly diagnostic large ocularium, and instead appeared deeply nested within the Eupnoi superfamily Phalangioidea. Our results are robust to analytical treatment and to homology scheme (dynamic vs. static notions of homology), resulting in a new phylogenetic proposal for Eupnoi and Dyspnoi. Ancestral state reconstruction suggests that the ancestral Palpatores was probably a tiny harvestman with an enlarged ocularium and glandular palpal setae in its enlarged and armed palps. We take the following taxonomic actions: Acropsopilionidae is removed from synonymy under Caddidae and its family status and membership in Dyspnoi are restored. Hesperopilio Shear, 1996 is removed from Caddoidea/Caddidae and transferred to Phalangioidea, but it is not assigned to any family.