scholarly journals Phylogeny of the subfamilies of Ichneumonidae (Hymenoptera)

2019 ◽  
Vol 71 ◽  
pp. 1-156 ◽  
Author(s):  
Andrew M.R. Bennett ◽  
Sophie Cardinal ◽  
Ian D. Gauld ◽  
David B. Wahl

A combined morphological and molecular phylogenetic analysis was performed to evaluate the subfamily relationships of the parasitoid wasp family Ichneumonidae (Hymenoptera). Data were obtained by coding 135 morphological and 6 biological characters for 131 exemplar species of ichneumonids and 3 species of Braconidae (the latter as outgroups). The species of ichneumonids represent all of the 42 currently recognized subfamilies. In addition, molecular sequence data (cytochrome oxidase I “DNA barcoding” region, the D2 region of 28S rDNA and part of the F2 copy of elongation factor 1-alpha) were obtained from specimens of the same species that were coded for morphology (1309 base pairs total). The data were analyzed using parsimony and Bayesian analyses. The parsimony analysis using all data recovered previously recognized informal subfamily groupings (Pimpliformes, Ophioniformes, Ichneumoniformes), although the relationships of these three groups to each other differed from previous studies and some of the subfamily relationships within these groupings had not previously been suggested. Specifically, Ophioniformes was the sister group to (Ichneumoniformes + Pimplformes), and Labeninae was placed near Ichneumoniformes, not as sister group to all Ichneumonidae except Xoridinae. The parsimony analysis using only morphological characters was poorly resolved and did not recover any of the three informal subfamily groupings and very few of the relationships were similar to the total-evidence parsimony analysis. The molecular-only parsimony analysis and both Bayesian analyses (total-evidence and molecular-only) recovered Pimpliformes, a restricted Ichneumoniformes grouping and many of the subfamily groupings recovered in the total-evidence parsimony analysis. A comparison and discussion of the results obtained by each phylogenetic method and different data sets is provided. It is concluded that the molecular characters produced results that were relatively consistent with traditional, non-phylogenetic concepts of relationships between the ichneumonid subfamilies, whereas the morphological characters did not (at least not by themselves). The inclusion of both molecular and morphological characters using parsimony produced a topology that was the closest to the traditional subfamily relationships. The method of analysis did not greatly affect the overall topology for the molecular-only analyses, but there were differences between Bayesian and parsimony results for the total-evidence analyses (especially near the root of the tree). The Bayesian results did not seem to be altered very much by the inclusion of morphological characters, unlike in the parsimony analysis. In summary, the following groups were supported in multiple analyses regardless of the characters used or method of tree-building: Pimpliformes, higher Ophioniformes, higher Pimpliformes, (Claseinae + Pedunculinae), (Banchinae + Stilbopinae), Campopleginae, Cremastinae, Diplazontinae, Ichneumoninae (including Alomya), Labeninae, Ophioninae, Poemeniinae, Rhyssinae, and Tersilochinae sensu stricto. Conversely, Ctenopelmatinae and Tryphoninae were never recovered without inclusion of other taxa. Based on the hypothesis of relationships obtained by the total-evidence parsimony analysis, the following formal taxonomic changes are proposed: Alomyinae Förster (= Alomya Panzer and Megalomya Uchida) is once again synonymized with Ichneumoninae and is now considered a tribe (Alomyinirev. stat.); and Notostilbops Townes is transferred from Stilbopinae to Banchinae, tribe Atrophini.

Author(s):  
T.S. Kemp

The vast majority of living and fossil mammals are placentals. Today there are about 4,400 species, which are traditionally organised into 18 Orders, with an extra one if the Pinnipedia are separated from the Carnivora, and a twentieth if the recently extinct Malagasy order Bibymalagasia is recognised as such. There have been many attempts to discover supraordinal groupings from amongst these Orders based on morphological characters, though few proposals have been universally accepted. It is only with the advent of increasingly large sets of molecular sequence data in the last few years that a reasonably robust resolution looks imminent, although these contemporary analyses are remarkably and controversially at odds with the traditional ones. Novacek et al. (1988) summarised the then current situation regarding supraordinal classification of placentals, a time at which morphology was still dominant but molecular data was at the threshold of significance. They accepted a basal group Edentata that combined the Xenarthra of the New World with the Pholidota of the Old, based on a few cranial characters, loss of the anterior teeth, and reduction of the enamel of the remaining ones. This left the rest of the living placentals as a monophyletic group Epitheria, sharing such apparently minor characters as the shape of the stapes bone in the ear. They found very little resolution within the Epitheria, and concluded that there was a polychotomy of no less than nine lineages arranged as a ‘star’ phylogeny. No remnant of the previously recognised taxon Ferungulata, created by Simpson (1945) for the Carnivora plus the ungulate orders Artiodactyla, Perissodactyla, Proboscidea, Hyracoidea, Sirenia, and Tubulidentata remained. On the other hand, three supra ordinal taxa of earlier authors did survive. One was Gregory’s (1910) Archonta, consisting of generally conservative forms and by now composed of the Primates, Dermoptera, Scandentia, and Chiroptera, but excluding the Lipotyphla. The second was Glires, originating with Linnaeus (1758) and widely accepted ever since, for the Rodentia and Lagomorpha; Novacek et al. (1988) tentatively placed the Macroscelidea as the sister-group of the Glires. The third supraordinal taxon recognised was, like Glires, well-established if not universally accepted.


2020 ◽  
Vol 190 (3) ◽  
pp. 1002-1019 ◽  
Author(s):  
Donald L J Quicke ◽  
Sergey A Belokobylskij ◽  
Yves Braet ◽  
Cornelis van Achterberg ◽  
Paul D N Hebert ◽  
...  

Abstract A new tribe of braconid wasps provisionally included in the Rhyssalinae, Laibaleini trib. nov., type genus Laibalea gen. nov. (type species Laibalea enigmatica sp. nov.), from Kenya and the Central African Republic, is described. A molecular dataset, with emphasis on basally derived taxa based on four gene fragments (28S D2–D3 expansion region, COI barcode, elongation factor 1-alpha and 16S ribosomal DNA), was analysed both alone and in combination with a morphological dataset. Molecular phylogenetic placement of the new species into an existing subfamily is complicated by the extreme sequence divergence of the three sequences obtained for Laibalea. In both the combined sequence analysis and the combined DNA plus morphological tree, Laibalea is recovered as a sister group to the Rhyssalinae plus all non-cyclostome lineage braconids excluding Mesostoinae, Maxfischeriinae and Aphidiinae. A consensus of morphological characters and molecular analyses suggests inclusion of Laibalea either in the otherwise principally Holarctic subfamily Rhyssalinae or perhap more basally, in the principally Gondwanan Mesostoinae s.l., although we cannot exclude the possibility that it might represent a separate basal lineage. We place Laibalea in its own tribe, provisionally included in Rhyssalinae. The DNA sequence data are presented for several genera for the first time. Avga, the type genus of Avgini, is shown not to belong to Mesostoinae s.l. or Hormiinae, but its exact relationships remain uncertain. The generic compositions of Rhyssalinae and Mesostoinae s.l. are revised. Anachyra, Apoavga, Neptihormius, Neoavga and Opiopterus are shown to belong to Mesostoinae s.s. A key to the tribes of Rhyssalinae is provided.


2006 ◽  
Vol 20 (5) ◽  
pp. 503 ◽  
Author(s):  
Gregory D. Edgecombe ◽  
Gonzalo Giribet

Scutigeromorpha (‘house centipedes’) play a pivotal role in myriapod systematics in being the sister group to all other chilopods, but their internal phylogeny has not been comprehensively appraised since K. W. Verhoeff’s morphological investigations a century ago. Relationships between the three families of Scutigeromorpha are inferred based on a combined analysis of approximately 5.5 Kb of sequence data from five molecular markers (complete 18S rRNA, a 2.2-Kb fragment of 28S rRNA, 16S rRNA, cytochrome c oxidase subunit I, histone H3) and 33 ingroup morphological characters. Molecular data are available for 19 ingroup terminals representing 14 morphospecies that include the genera Scutigerina, Madagassophora (family Scutigerinidae), Sphendononema (family Pselliodidae), Scutigera, Thereuopoda, Thereuopodina, Thereuonema, Allothereua and Parascutigera (family Scutigeridae). Morphology resolves the southern African–Malagasy Scutigerinidae as sister to all other Scutigeromorpha, whereas rival sister-group relationships between the Neotropical–Afrotropical Pselliodidae and Scutigerinidae + Scutigeridae or Pselliodidae + Scutigerinidae and Scutigeridae are resolved by the molecular and combined analyses. Monophyly of Scutigeridae and Thereuoneminae are stable across a broad range of analytical parameters. Thereuoneminae is composed of two stable clades: an Allothereua + Parascutigera group, and a grouping of Thereuopoda, Thereuonema and Thereuopodina. Molecular and combined analyses resolve the genus Scutigerina and the morphospecies Scutigerina weberi as paraphyletic, in both cases with a Malagasy clade excluding populations from southern Africa.


2019 ◽  
Vol 187 (1) ◽  
pp. 100-116 ◽  
Author(s):  
Andrew R Brinkworth ◽  
Robert Sansom ◽  
Matthew A Wills

Abstract Notwithstanding the rapidly increasing sampling density of molecular sequence data, morphological characters still make an important contribution to our understanding of the evolutionary relationships of arthropod groups. In many clades, characters relating to the number and morphological specialization of appendages are ascribed particular phylogenetic significance and may be preferentially sampled. However, previous studies have shown that partitions of morphological character matrices often imply significantly different phylogenies. Here, we ask whether a similar incongruence is observed in the appendage and non-appendage characters of arthropods. We apply tree length (incongruence length difference, ILD) and tree distance (incongruence relationship difference, IRD) tests to these partitions in an empirical sample of 53 published neontological datasets for arthropods. We find significant incongruence about one time in five: more often than expected, but markedly less often than in previous partition studies. We also find similar levels of homoplasy in limb and non-limb characters, both in terms of internal consistency and consistency relative to molecular trees. Taken together, these findings imply that sampled limb and non-limb characters are of similar phylogenetic utility and quality, and that a total evidence approach to their analysis is preferable.


2019 ◽  
Vol 69 (4) ◽  
pp. 638-659 ◽  
Author(s):  
Yimeng Li ◽  
Marcello Ruta ◽  
Matthew A Wills

Abstract Despite the increasing importance of molecular sequence data, morphology still makes an important contribution to resolving the phylogeny of many groups, and is the only source of data for most fossils. Most systematists sample morphological characters as broadly as possible on the principle of total evidence. However, it is not uncommon for sampling to be focused on particular aspects of anatomy, either because characters therein are believed to be more informative, or because preservation biases restrict what is available. Empirically, the optimal trees from partitions of morphological data sets often represent significantly different hypotheses of relationships. Previous work on hard-part versus soft-part characters across animal phyla revealed significant differences in about a half of sampled studies. Similarly, studies of the craniodental versus postcranial characters of vertebrates revealed significantly different trees in about one-third of cases, with the highest rates observed in non-avian dinosaurs. We test whether this is a generality here with a much larger sample of 81 published data matrices across all major dinosaur groups. Using the incongruence length difference test and two variants of the incongruence relationship difference test, we found significant incongruence in about 50% of cases. Incongruence is not uniformly distributed across major dinosaur clades, being highest (63%) in Theropoda and lowest (25%) in Thyreophora. As in previous studies, our partition tests show some sensitivity to matrix dimensions and the amount and distribution of missing entries. Levels of homoplasy and retained synapomorphy are similar between partitions, such that incongruence must partly reflect differences in patterns of homoplasy between partitions, which may itself be a function of modularity and mosaic evolution. Finally, we implement new tests to determine which partition yields trees most similar to those from the entire matrix. Despite no bias across dinosaurs overall, there are striking differences between major groups. The craniodental characters of Ornithischia and the postcranial characters of Saurischia yield trees most similar to the “total evidence” trees derived from the entire matrix. Trees from these same character partitions also tend to be most stratigraphically congruent: a mutual consilience suggesting that those partitions yield more accurate trees. [Dinosauria; homoplasy; partition homogeneity.]


Zootaxa ◽  
2011 ◽  
Vol 2984 (1) ◽  
pp. 67 ◽  
Author(s):  
LEANDRO C. S. ASSIS ◽  
MARCELO R. DE CARVALHO ◽  
QUENTIN D. WHEELER

David Wake and colleagues provided a thought-provoking review of the concept of homoplasy through the integration, within a phylogenetic framework, of genetic and developmental data (Wake et al. 2011). According to them (p. 1032) “Molecular sequence data have greatly increased our ability to identify homoplastic traits.” This is made clear, for example, in their flow chart for homoplasy detection (Figure 2, p. 1034), wherein homoplasy is discovered through the mapping of “traits of interest” onto a phylogram, a practice common in the molecular phylogenetic paradigm. The “mapping” is usually of morphological characters that are employed to support the chosen (molecular) topology, but which, as a consequence, do not themselves contribute to the formation of those topologies (Assis & Carvalho 2010).


2004 ◽  
Vol 73 (1-2) ◽  
pp. 3-163 ◽  
Author(s):  
Ronald A. Jenner

This paper critically assesses all morphological cladistic analyses of the Metazoa that were published during the last one and a half decades. Molecular and total evidence analyses are also critically reviewed. This study focuses on evaluating alternative phylogenetic positions of the ‘acoelomate’ worms: Platyhelminthes, Nemertea, and Gnathostomulida. This paper consists of two parts. In Part I, all recently proposed sister group hypotheses and the supporting synapomorphies for these phyla are evaluated. Discrepancies in the treatment of corresponding characters in different cladistic analyses are identified, and where possible, resolved. In Part II, the overall phylogenetic significance across the Metazoa of all characters relevant for placing the ‘acoelomate’ worms is examined. The coding and scoring of these characters for other phyla are evaluated, and uncertainties in our understanding are pointed out in order to guide future research. The characters discussed in this paper are broadly categorized as follows: epidermis and cuticle, reproduction and sexual condition, development, larval forms, coeloms and mesoderm source, nervous system and sensory organs, nephridia, musculature, digestive system, and miscellaneous characters. Competing phylogenetic hypotheses are compared in terms of several criteria: 1) taxon sampling and the fulfillment of domain of definition for each character; 2) character sampling; 3) character coding; 4) character scoring and quality of primary homology; 5) quality of the proposed diagnostic synapomorphies as secondary homologies. On the basis of this study I conclude that a sister group for the Platyhelminthes has not yet been unambiguously established. A clade minimally composed of Neotrochozoa (Mollusca, Sipuncula, Echiura, Annelida) emerges as the most likely sister group of the Nemertea on the basis of morphological and total evidence analyses. Finally, morphological data currrently favor a sister group relationship of Gnathostomulida and Syndermata (probably plus Micrognathozoa). In contrast, molecular or total evidence analyses have not identified a reliable sister group of Gnathostomulida.Further progress in our understanding of metazoan phylogeny crucially depends on the improvement of the quality of currently adopted cladistic data matrices. A thorough reassessment of many of the more than 70 morphological characters discussed here is necessary. Despite the recent compilation of comprehensive data matrices, the power to test competing hypotheses of higher-level metazoan relationships is critically compromised due to uncritical data selection and poor character study in even the most recently published cladistic analyses.


Paleobiology ◽  
1994 ◽  
Vol 20 (3) ◽  
pp. 259-273 ◽  
Author(s):  
Andrew B. Smith ◽  
D. T. J. Littlewood

Molecular data are becoming an indispensable tool for the reconstruction of phylogenies. Fossil molecular data remain scarce, but have the potential to resolve patterns of deep branching and provide empirical tests of tree reconstruction techniques. A total evidence approach, combining and comparing complementary morphological, molecular and stratigraphical data from both recent and fossil taxa, is advocated as the most promising way forward because there are several well-established problems that can afflict the analysis of molecular sequence data sometimes resulting in spurious tree topologies. The integration of evidence allows us to: (1) choose suitable taxa for molecular phylogenetic analysis for the question at hand; (2) discriminate between conflicting hypotheses of taxonomic relationship and phylogeny; (3) evaluate procedures and assumptions underlying methods of building trees; and (4) estimate rates of molecular evolution in the geological past. Paleontology offers a set of independent data for comparison and corroboration of analyses and provides the only direct means of calibrating molecular trees, thus giving insight into rates of molecular evolution in the geological past.


Zootaxa ◽  
2002 ◽  
Vol 87 (1) ◽  
pp. 1 ◽  
Author(s):  
FRANK LOUIS CARLE ◽  
KARL M. KJER

Phylogenetic analysis was performed on a set of 242 morphological characters. The taxon sample included 31 Libellula, and representative species from selected libeluline tribes, from all libellulid subfamilies, from all libelluloid families, from all anisopteran superfamilies, and Epiophlebia. Corduliinae was shown to be paraphyletic even among genera characterized by a well developed anal loop bisector. Sympetrini was found to be polyphyletic with Crocothemis the sister group to Libellulini. The traditional placement of Trameini, far from Libellulini is in doubt, because it is here placed as the sister group to Crocothemis + Libellulini. Kennedy’s phylogeny of Libellula was largely corroborated, with the following exceptions: the subgenera Libellula, Eolibellula, and Syntetrum form a monophyletic group which is the sister group to a clade including Belonia, Holotania, Neotetrum, and Eotainia subgenus nov. [type species Mesothemis composita Hagen]; and Eurothemis is determined to be the sister group of Ladona instead of Neotetrum. In addition we confirm Belonia to be monophyletic, and find Platetrum + Plathemis to form a monophyletic group, sister to Ladona + Eurothemis; these four subgenera together form the sister group to Libellula sensu stricto (s.s.).


Zootaxa ◽  
2008 ◽  
Vol 1792 (1) ◽  
pp. 1 ◽  
Author(s):  
GUSTAVO HORMIGA ◽  
LIHONG TU

The spider genus Putaoa new genus (Araneae, Pimoidae) is described to place two species of pimoids from China, Putaoa huaping new species (the type species) and P. megacantha (Xu & Li, 2007) new combination. Parsimony analysis of morphological characters provides support for the monophyly of Putaoa and for its sister group relationship to the genus Weintrauboa Hormiga, 2003 and corroborates the monophyly of Pimoidae.


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