A classification of the Ericaceae: subfamilies and tribes

The variation shown by 60 characters (morphological, anatomical, embryological, chemical and cytological) is tabulated for the whole of the Ericaceae. The reliability of the characters and the observations on them are discussed, and it is emphasized that many characters are difficult to use because they are rarely restricted to one taxon. Six subfamilies are recognized. Within the Rhododendroideae seven tribes are recognized, the monogeneric Daboecieae being new; Epigaea is transferred to the Rhododendroideae from the Andromedeae and Diplarche from the Diapensiaceae. Calluna is placed in a new monotypic tribe within the Ericoideae. The extended Vaccinioideae includes the Arbutoideae; five tribes are recognized, of which the Enkiantheae and Cassiopeae are new. The Pyroloideae and Monotropoideae are left unchanged: a new subfamily, the Wittsteinioideae, includes the deviant monotypic genus Wittsteinia. A key to subfamilies, and to genera of the Pyroloideae, Rhododendroideae and superior-ovaried Vaccinioideae is given.     

A chemotaxonomic survey of flavonoids and simple phenols in leaves of the Ericaceae

Leaves of 344 species of the Ericaceae and of 37 species in three related families have been surveyed for flavonoids and simple phenols. The yellow flavonol gossypetin was found in 158 ericaceous species and is of chemotaxonomic interest within the Rhododendroideae (where it occurs in 11 of 19 genera) and in the Vaccinioideae (in 3 of 22 genera). 5-0-methylflavonols appear in 81 species, again mainly in the Rhododendroideae (eight genera); whereas 3,5-di-O-methylquercetin, caryatin, is present only in Bejaria, Phyllodoce and Rhododendron. Dihydroflavonols. common in Rhododendron , show a sporadic distribution elsewhere in the family. Of the simpler phenols surveyed, gentisic acid is common but methyl salicylate is present mainly in Gaultheria. Hydroquinone appears in ten related genera, methylhydroquinone in three and orcinol in seven. The chemical evidence generally supports Stevens' recent classification of the family, particularly his inclusion of Epigaea in the Rhododendroideae and his transfer of Diplarche from the Diapensiaceae to the Ericaceae. There is also chemical evidence to support the separation of Calluna and Cassiope , the inclusion of the Aibutoideae within the Vaccinioideae and the retention of the Pyroloideae and Monotropoideae within the Ericaceae. In addition, chemistry indicates that Phyllodoce should be placed nearer to the Rhodoreae than in Stevens' system and that Cassiope and Harrimanella , placed together in the Cassiopeae by Stevens, are distinctly different.     

Phylogenetic analysis of the Monocotylidae (Monogenea) inferred from 28S rDNA sequences

The current classification of the Monocotylidae (Monogenea) is based on a phylogeny generated from morphological characters. The present study tests the morphological phylogenetic hypothesis using molecular methods. Sequences from domains C2 and D1 and the partial domains C1 and D2 from the 28S rDNA gene for 26 species of monocotylids from six of the seven subfamilies were used. Trees were generated using maximum parsimony, neighbour joining and maximum likelihood algorithms. The maximum parsimony tree, with branches showing less than 70% bootstrap support collapsed, had a topology identical to that obtained using the maximum likelihood analysis. The neighbour joining tree, with branches showing less than 70% support collapsed, differed only in its placement of Heterocotyle capricornensis as the sister group to the Decacotylinae clade. The molecular tree largely supports the subfamilies established using morphological characters. Differences are primarily how the subfamilies are related to each other. The monophyly of the Calicotylinae and Merizocotylinae and their sister group relationship is supported by high bootstrap values in all three methods, but relationships within the Merizocotylinae are unclear. Merizocotyle is paraphyletic and our data suggest that Mycteronastes and Thaumatocotyle, which were synonymized with Merizocotyle after the morphological cladistic analysis, should perhaps be resurrected as valid genera. The monophyly of the Monocotylinae and Decacotylinae is also supported by high bootstrap values. The Decacotylinae, which was considered previously to be the sister group to the Calicotylinae plus Merizocotylinae, is grouped in an unresolved polychotomy with the Monocotylinae and members of the Heterocotylinae. According to our molecular data, the Heterocotylinae is paraphyletic. Molecular data support a sister group relationship between Troglocephalus rhinobatidis and Neoheterocotyle rhinobatidis to the exclusion of the other species of Neoheterocotyle and recognition of Troglocephalus renders Neoheterocotyle paraphyletic. We propose Troglocephalus incertae sedis. An updated classification and full species list of the Monocotylidae is provided.     

Phylogenetic analysis of the Monocotylidae (Monogenea) inferred from 28S rDNA sequences

The current classification of the Monocotylidae (Monogenea) is based on a phylogeny generated from morphological characters. The present study tests the morphological phylogenetic hypothesis using molecular methods. Sequences from domains C2 and D1 and the partial domains C1 and D2 from the 28S rDNA gene for 26 species of monocotylids from six of the seven subfamilies were used. Trees were generated using maximum parsimony, neighbour joining and maximum likelihood algorithms. The maximum parsimony tree, with branches showing less than 70% bootstrap support collapsed, had a topology identical to that obtained using the maximum likelihood analysis. The neighbour joining tree, with branches showing less than 70% support collapsed, differed only in its placement of Heterocotyle capricornensis as the sister group to the Decacotylinae clade. The molecular tree largely supports the subfamilies established using morphological characters. Differences are primarily how the subfamilies are related to each other. The monophyly of the Calicotylinae and Merizocotylinae and their sister group relationship is supported by high bootstrap values in all three methods, but relationships within the Merizocotylinae are unclear. Merizocotyle is paraphyletic and our data suggest that Mycteronastes and Thaumatocotyle, which were synonymized with Merizocotyle after the morphological cladistic analysis, should perhaps be resurrected as valid genera. The monophyly of the Monocotylinae and Decacotylinae is also supported by high bootstrap values. The Decacotylinae, which was considered previously to be the sister group to the Calicotylinae plus Merizocotylinae, is grouped in an unresolved polychotomy with the Monocotylinae and members of the Heterocotylinae. According to our molecular data, the Heterocotylinae is paraphyletic. Molecular data support a sister group relationship between Troglocephalus rhinobatidis and Neoheterocotyle rhinobatidis to the exclusion of the other species of Neoheterocotyle and recognition of Troglocephalus renders Neoheterocotyle paraphyletic. We propose Troglocephalus incertae sedis. An updated classification and full species list of the Monocotylidae is provided.     

STUDIES OF THE MONOTROPOIDEAE (ERICACEAE). FLORAL NECTARIES: ANATOMY AND FUNCTION IN POLLINATION ECOLOGY

The morphological and vascular characteristics of the nectaries of the species of the Monotropoideae (Ericaceae) reflect the morphology and declination of the flowers. These, in turn, are related to the pollination systems of the species. The nectaries of members of the Monotropoideae exhibit a range in form from slender, elongate projections, to short, stout ones, or to low ridges between the staminal bases.     

Mitochondrial and nuclear markers support the monophyly of Dolichopodidae and suggest a rapid origin of the subfamilies (Diptera: Empidoidea)

Abstract. The Dolichopodidae is a species‐rich dipteran group with almost 7000 described species. The monophyly of the subfamilies and their relationships remain largely unknown because the polarities of key morphological characters are unclear and molecular data are available only for 9 of the 19 proposed subfamilies. Here we test whether molecular data from two nuclear (18S, 28S) and four mitochondrial (12S, 16S, Cytb, COI) genes can resolve the higher‐level relationships within the family. Our study is based on 76 Oriental species from 12 dolichopodid subfamilies and uses eight species of Empididae and Hybotidae as outgroups. Parsimony and likelihood analyses confirm the monophyly of the Dolichopodidae, as well as the monophyly of five of the ten subfamilies represented by more than two species [Sympycninae, Sciapodinae, Dolichopodinae, Hydrophorinae (excluding tribe Aphrosylini), Neurigoninae]. There is strong support for restoring the tribe Aphrosylini as a separate subfamily Aphrosylinae. The monophyly of Medeterinae, Peloropeodinae and Diaphorinae is dependent on which tree reconstruction technique is used, how indels are coded, and whether the fast‐evolving sites are excluded. Overall, we find that our sample of Oriental species is largely compatible with the subfamily concepts that were developed for the northern temperate fauna. However, our data provide little support for relationships between the subfamilies. Branch lengths, saturation, and distance plots suggest that this is probably the result of the rapid origin of dolichopodid subfamilies over a relatively short time. We find that genera that are difficult to place into subfamilies based on morphological characters are generally also difficult to place using molecular data. We predict that a dense, balanced taxon sample and protein‐encoding nuclear genes will be needed to resolve the higher‐level relationships in the Dolichopodidae.     

Phylogenetic relationships of the Suidae (Mammalia,Cetartiodactyla): new insights on the relationships within Suoidea

Orliac, M. J., Antoine, P. ‐O., Ducrocq, S. (2010). Phylogenetic relationships of the Suidae (Mammalia, Cetartiodactyla): new insights on the relationships within Suoidea. —Zoologica Scripta, 39, 315–330. In most analyses, both molecular and morphological phylogenies of the Cetartiodactyla support the monophyly of Suoidea. However, the evolutionary history of this superfamily remains poorly known primarily due to long‐lasting debates about the taxonomic content and relationships of the suoid families and subfamilies. Despite their crucial position in the reconstruction of the phylogeny of Cetartiodactyla, Suoidea themselves have received little attention in those phylogenies, and no extensive analysis of the group has been performed so far. We therefore examine the phylogeny of the Suidae through the first phylogenetic analysis of Suoidea, including recent and fossil representatives of all four putative families. The results support the monophyly of the traditional suid subfamilies and indicate the Sanitheriidae as sister taxon to the Suidae clade. The evolutionary history within Suidae reveals its complexity, with major convergences involving important morphological structures such as the auditory region or the upper male canine. Divergent signals gathered from either dental or cranio‐mandibular features are responsible for two long‐lasting unresolved issues within Suoidea: the question of the relationships between ‘Old World’ and ‘New World’ peccaries remaining unsolved, as well as the position and familial status of the mid‐Tertiary tayassuid Perchoerus.     

New insight into the systematics of Tomoceridae (Hexapoda,Collembola) by integrating molecular and morphological evidence

Tomoceridae is common but among the most problematic groups of Collembola. Its position within Collembola and the relationships within the family remain obscure. This also extends to the generic division of the subfamily Tomocerinae that remains controversial. This study examines these issues by integrating both molecular and morphological evidence. Our molecular phylogeny based on rDNA sequences supports the monophyly of Tomoceridae and the sister relationship between Tomocerinae and Lepidophorellinae. Reconstructions and tree topology tests constraining monophyly did not resolve the relationships between Tomoceridae and other collembolan groups. We also examined the morphology of the first instar (primary) larvae, which has significant phylogenetic value among higher Collembola. Mapping primary chaetotaxy onto our molecular phylogeny provided further evidence for the unique position of Tomoceridae within Entomobryomorpha and Collembola. The monophyly and subfamilial classification within Tomoceridae were validated here, whereas its position among Collembola will need further studies in a broader consideration across the major collembolan orders. Within Tomocerinae, the monophyly of Pogonognathellus was demonstrated, but the status of Tomocerus and Tomocerina is still to be resolved.     

Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem

No qualitative cladistic analysis has been performed previously for the subfamily classification of Pompilidae (Hymenoptera). In 1994 Shimizu proposed six subfamilies, but their validity and relationships remain inconclusive. The objective of this study was to perform a quantitative analysis of phylogenetic relationships of the Pompilidae, with emphasis on testing the validity of proposed subfamilies. Two cladistic analyses were performed based on morphological evidence. First, a maximum-parsimony analysis of Shimizu's original morphological data matrix (72 taxa by 54 characters) was conducted, with the data subjected to a heuristic search for the first time with phylogenetic software. The resulting strict-consensus cladogram yielded a monophyletic Ceropalinae that was sister group to a large polytomy containing members of the remaining five subfamilies. In a second analysis, several of Shimizu's characters were re-examined, and new characters and more taxa were added to the data set. Terminal taxa were coded as species rather than as generic abstractions, and 20 additional morphological characters were introduced. The analysis was based on 77 morphological characters derived from the adults of 84 taxa. This second analysis suggested that Notocyphinae sensu Shimizu (1994) was nested within Pompilinae and that Epipompilinae sensu Shimizu (1994) was nested within Ctenocerinae; neither should retain their status as a separate subfamily. Lastly, Chirodamus s .s., which historically has been a member of the Pepsinae, is placed within the Pompilinae with reservations rather than erecting a new subfamily. After these allowances were made, a strict consensus tree gave the following relationships: (Ceropalinae + (Pepsinae + (Ctenocerinae + Pompilinae))).     

Molecules,morphology, missing data and the phylogenetic position of a recently extinct madtom catfish (Actinopterygii: Ictaluridae)

Taxa missing large amounts of data pose challenges that may hinder the recovery of a well‐resolved, accurate phylogeny and leave questions surrounding their phylogenetic position. Systematists commonly have to contend with one or two species in a group for which there is little or no material available suitable for recovering molecular data. It is unclear whether these taxa can be better placed using analyses based on morphological data only, or should be included in broader analyses based on both morphological and molecular data. The extinct madtom catfish Noturus trautmani is known from few specimens for which molecular data are unavailable. We included this taxon in parsimony and Bayesian analyses of relationships of madtom catfishes based on a combination of morphological and molecular data. Results indicate that using a combination of morphological and molecular data does a better job at providing a phylogenetic placement for N. trautmani than morphology alone, even though it is missing all of its molecular characters. We provide a novel hypothesis of relationships among Noturus species and recommendations for classification within the group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 60–75.     

A phylogenetic analysis of the Orchidaceae: evidence from rbcL nucleotide

Cladistic parsimony analyses of rbcL nucleotide sequence data from 171 taxa representing nearly all tribes and subtribes of Orchidaceae are presented here. These analyses divide the family into five primary monophyletic clades: apostasioid, cypripedioid, vanilloid, orchidoid, and epidendroid orchids, arranged in that order. These clades, with the exception of the vanilloids, essentially correspond to currently recognized subfamilies. A distinct subfamily, based upon tribe Vanilleae, is supported for Vanilla and its allies. The general tree topology is, for the most part, congruent with previously published hypotheses of intrafamilial relationships; however, there is no evidence supporting the previously recognized subfamilies Spiranthoideae, Neottioideae, or Vandoideae. Subfamily Spiranthoideae is embedded within a single clade containing members of Orchidoideae and sister to tribe Diurideae. Genera representing tribe Tropideae are placed within the epidendroid clade. Most traditional subtribal units are supported within each clade, but few tribes, as currently circumscribed, are monophyletic. Although powerful in assessing monophyly of clades within the family, in this case rbcL fails to provide strong support for the interrelationships of the subfamilies (i.e., along the spine of the tree). The cladograms presented here should serve as a standard to which future morphological and molecular studies can be compared.     

Further progress on the phylogeny of Noctuoidea (Insecta: Lepidoptera) using an expanded gene sample

Major progress has been made recently toward resolving the phylogeny of Noctuoidea, the largest superfamily of Lepidoptera. However, numerous questions and weakly supported nodes remain. In this paper we independently check and extend the main findings of multiple recent authors by performing maximum‐likelihood analyses of 5–19 genes (6.7–18.6 kb) in 74 noctuoids representing all the families and a majority of the subfamilies. Our results strongly support the six family system of Zahiri et al., with the former Lymantriidae and Arctiidae subsumed within the huge family Erebidae, and Noctuidae restricted largely to the subfamilies with so‐called trifine hindwing venation. Our data also strongly corroborate monophyly of the set of four families with quadrifid forewing venation, to the exclusion of Notodontidae, and removal from the latter of Oenosandridae. Other among‐family relationships, however, remain unsettled. Our evidence is equivocal on the position of Oenosandridae, which are sister group to either Notodontidae alone or to all other noctuoids. Like other recent nuclear gene studies, our results also provide no strong support for relationships among the four quadrifid forewing families. In contrast, within families our analyses significantly expand the list of robustly resolved relationships, while introducing no strong conflicts with previous molecular studies. Within Notodontidae, for which we present the largest molecular taxon sample to date, we find strong evidence for polyphyly for some, or all, recent definitions of the subfamilies Thaumetopoeinae, Pygaerinae, Notodontinae and Heterocampinae. Deeper divergences are incompletely resolved but there is strong support for multiple ‘backbone’ nodes subtending most of the subfamilies studied. Within Erebidae, we find much agreement and no strong conflict with a recent previous study regarding relationships among subfamilies, and somewhat stronger support. Although many questions remain, the two studies together firmly resolve positions for over half the subfamilies. Within Noctuidae, we find no strong conflict with previous molecular studies regarding relationships among subfamilies, but much stronger resolution along the ‘backbone’ of the phylogeny. Combining information from multiple studies yields strongly resolved positions for most of the subfamilies. Finally, our results strongly suggest that the tribes Pseudeustrotiini and Prodeniini, currently assigned to the largest subfamily, Noctuinae, do not belong there. In sum, our results provide additional corroboration for the main outlines of family‐level phylogeny in Noctuoidea, and contribute toward resolving relationships within families.     

Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure

The Monotropoideae (Ericaceae) are nonphotosynthetic plants that obtain fixed carbon from their fungal mycorrhizal associates. To infer the evolutionary history of this symbiosis we identified both the plant and fungal lineages involved using a molecular phylogenetic approach to screen 331 plants, representing 10 of the 12 described species. For five species no prior molecular data were available; for three species we confirmed prior studies which used limited samples; for five species all previous reports are in conflict with our results, which are supported by sequence analysis of multiple samples and are consistent with the phylogenetic patterns of host plants. The phylogenetic patterns observed indicate that: (i) each of the 13 plant phylogenetic lineages identified is specialized to a different genus or species group within five families of ectomycorrhizal Basidiomycetes; (ii) mycorrhizal specificity is correlated with phylogeny; (iii) in sympatry, there is no overlap in mature plant fungal symbionts even if the fungi and the plants are closely related; and (iv) there are geographical patterns to specificity.     

Molecular phylogeny of Potamotrygonocotyle (Monogenea,Monocotylidae) challenges the validity of some of its species

Fehlauer‐Ale, K. H. & Littlewood, D. T. J. (2011). Molecular phylogeny of Potamotrygonocotyle (Monogenea, Monocotylidae) challenges the validity of some of its species. —Zoologica Scripta, 40, 638–658. The marine‐derived stingrays Potamotrygonidae are the only chondrichthyans landlocked to freshwaters of Central and South America. The family includes approximately 22 described species organized in four genera widely distributed across the main Atlantic and Caribbean continental drainages. Investigations into the parasite fauna of potamotrygonids have mainly focused on cestodes, with a few studies addressing the biodiversity of monogeneans. Potamotrygonocotyle (Monogenea, Monocotylidae) is composed of 12 species, exclusively found in the gills of species of Potamotrygonidae. This study presents molecular phylogenetic analyses of this group of monogeneans distributed throughout La Plata and Amazonas basins, with the purpose of readdressing the phylogeny of Monocotylidae based on 28S rDNA sequences and of unravelling the phylogeny of its species using data from mitochondrial gene cytochrome c oxidase subunit I and nuclear gene internal transcribed spacer 1. The phylogenetic status of the five tested monocotylid subfamilies and most of their internal relationships are concordant with the results of a previous study, and the monophyletic status of Potamotrygonocotyle based on molecular data is corroborated for the first time. However, the placement of the genus within Monocotylidae is not resolved, as its sister‐group relationship with Neoheterocotyle and Troglocephalus is uncertain. Investigations into the relationships within Potamotrygonocotyle support the monophyletic status of nine nominal species and suggest the existence of cryptic lineages for the remaining three. Molecular analyses reveal distinct sister‐groups relationships in comparison with a previously published phylogeny for the genus based on morphological data. Finally, the surveys of this study expand the known distribution range of some members of Potamotrygonocotyle.     

Contrasting evolutionary history of hedgehogs and gymnures (Mammalia: Erinaceomorpha) as inferred from a multigene study

The subfamilies Erinaceinae and Galericinae of the extant family Erinaceidae are the only living representatives of the once diverse taxon Erinaceomorpha. In the present study, we performed the first multilocus analysis of phylogenetic relationships among genera of Erinaceidae and estimated the split times between and within the two subfamilies. The analyses of five nuclear and two mitochondrial genes produced a well‐resolved molecular phylogeny. Generally, the molecular tree is compatible with the morphology‐based taxonomy proposed by Frost, Wozencraft & Hoffmann with the exception of the position of Mesechinus, which is placed as the closest sister taxon of Hemiechinus. Another point of contradiction between molecular and morphological phylogenies is the position of Hylomys megalotis, which was consistently placed as the most basal branch among all gymnures in molecular analyses. Genetic relationships between Erinaceus and Atelerix remain unclear, suggesting a hard trichotomy among these two lineages and Hemiechinus + Paraechinus. Molecular dating suggests an ancient origin of the extant gymnure lineages, which date back to the late Eocene to early Oligocene. The age of the basal split within spiny hedgehogs is relatively recent and corresponds to the Miocene–Pliocene boundary. Possible changes to the erinaceid taxonomy are considered. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 499–519.     

Toward a Better Understanding of the Phylogeny of the Asplanchnidae (Rotifera)

We investigated the phylogenetic relationships of Family Asplanchnidae using both morphological and molecular data. The morphological database, comprising 23 characters from 19 taxa (15 Asplanchnidae and 4 outgroups), was compiled from a survey of the literature and our own observations; the molecular data (ITS and V4 region nuclear regions and mitochondrial cox1) was sequenced from specimens that we collected. Our analysis of the morphological data set (maximum parsimony) yielded 12 most-parsimonious trees with a tree length of 27 steps. From this analysis we conclude (1) Asplanchnidae is a monophyletic group as are the three genera comprising it, (2) there is no compelling support for the argument that Asplanchna should be separated into two discrete genera, and (3) there is some support for the proposal that Asplanchnidae and Synchaetidae are sister groups. Our analysis of the molecular data set supports the first two of these conclusions while the sister group of the family varied depending on the gene region analyzed and families and genera included. Current understanding of the phylogeny of Asplanchnidae is hampered by the need for additional informative morphological characters and a lack of molecular data for the genus Harringia and several other members of the Asplanchnidae.     

Critical Reexamination of Palynological Characters Used to Delimit Asclepiadaceae in Comparison to the Molecular Phylogeny Obtained from PlastidmatK Sequences

The family Asclepiadaceae (Dicotyledones) was created by Brown in 1810 by splitting in two the family Apocynaceae of Jussieu established in 1789. The morphological characters used to make this distinction were mainly palynological, such as presence of tetrads or pollinia and number and orientation of pollinia. Those characters, still used in higher taxonomic delimitation (families, subfamilies, and tribes), are here critically reexamined and compared to a molecular phylogeny obtained with one of the more variable plastid genes (matK) of 46 species in the order Gentianales. In this molecular phylogeny, Asclepiadaceae form a monophyletic group derived from within Apocynaceae. Each of the subfamilies of Asclepiadaceae is monophyletic and based on reliable palynological characters, but palynological characters are not useful to delimit tribes of the subfamily Asclepiadoideae. Based on the molecular data, these tribes have undergone parallelisms in several reproductive traits.