Ultrastructural and immunocytochemical investigation of acoel sperms with 9 + 1 axoneme structure: new sperm characters for unraveling phylogeny in Acoela

Acoel sperm characters proved useful in deciphering acoel taxonomy. The phylogenetic value of sperm characters in closely related sub-groups or in a monophyletic taxon has not yet been assessed. We have investigated sperm ultrastructure in seven members of the monophyletic taxon Childia sensu (Tekle et al. J Zool Sys Evol Res 43(1):72–90, 2005) and in their closest relatives, the Mecynostomidae (four taxa). All members of Childia examined show little variation in their sperm ultrastructure. The common characters of Childia taxa are: 9 + 1 axoneme structure, the presence of six distal cytoplasmic microtubules in the absence of axial or cortical ones, long nucleus and extensive nucleus–flagella overlap. We have identified a new set of cytoplasmic microtubules lying in the centriolar end of the sperm cell, distal microtubules. The origin and phylogenetic significance of this character is discussed. The types and arrangement of cytoplasmic granules could be used as phylogenetic characters at a low taxonomic level. A loose membrane amorphous core type of granule was found to be a synapomorphy for the following clade within the taxon Childia: C. crassum + C. groenlandica + C. vivipara + C. brachyposthium + C. macroposthium. Sausage shaped granules are plesiomorphic among the taxa examined. The rest of the granule characters were found to be homoplasious. Sperm ultrastructural characters have again proven their concordance with molecular phylogeny. The only morphological synapomorphies known for the sister taxa Childia–Mecynostomidae, in the molecular phylogeny, are characters derived from sperm ultrastructure: distal microtubules arranged in two groups of three microtubules each and a 9 + 1 axoneme structure. The spermatozoa of Childia and Mecynostomidae show 9 + 1 axoneme configuration, seemingly similar to the 9 + ‘1’ axoneme pattern of the Platyhelminthes—Trepaxonemata. Using electron-microscope immunocytochemistry, we have demonstrated that, unlike the central cylinder of trepaxonematans, the central cylinder of the 9 + 1 axonemal pattern in acoels is immunoreactive to tubulin and contains a single central microtubule. Therefore, the 9 + 1 patterns in acoels and trepaxonematans are homoplasious.     

Evolutionary morphology of whip spiders: towards a phylogenetic system (Chelicerata: Arachnida: Amblypygi)*

The aim of this study is to present a cladogram and phylogenetic system and to use this to discuss the phylogeny and biogeography of the Amblypygi. A total of 29 morphological structures were studied, their plesiomorphic and apomorphic characters or character states were identified, and the resulting data matrix was analysed. As a result, the ‘old’Charontidae or Pulvillata emerge as a paraphyletic group; the genus Paracharon is the sister group of all other amblypygids, which are now termed Euamblypygi. The ‘new’Charontidae (sensu Quintero: the genera Stygophrynus and Charon) are the sister group of the Phrynida or Apulvillata; together they form the Neoamblypygi. The relationships of the genera of the Charinidae cannot be resolved with the available data. They may be a paraphyletic group. The genus Catageus is a possible candidate for being the sister group of the Neoamblypygi. The new system allows a discussion of the phylogeny and biogeography of whip spiders. It also points to unresolved taxa and thus indicates the questions future research should address.     

The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families

A phylogenetic analysis of Adephaga is presented. It is based on 148 morphological characters of adults and larvae and focussed on a placement of the recently described Meruidae, and the genus‐level phylogeny of the smaller aquatic families Gyrinidae, Haliplidae and Noteridae. We found a sister group relationship between Gyrinidae and the remaining adephagan families, as was found in previous studies using morphology. Haliplidae are either the sister group of Dytiscoidea or the sister group of a clade comprising Geadephaga and the dytiscoid families. Trachypachidae was placed as the sister group of the rhysodid‐carabid clade or of Dytiscoidea. The monophyly of Dytiscoidea including Meru is well supported. Autapomorphies are the extensive metathoracic intercoxal septum, the origin of the metafurca from this structure, the loss of Mm. furcacoxalis anterior and posterior, and possibly the presence of an elongated subcubital setal binding patch. Meruidae was placed as sister group of the Noteridae. Synapomorphies are the absence of the transverse ridge of the metaventrite, the fusion of abdominal segments III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7 and 9. The Meru‐noterid clade is the sister group of the remaining Dytiscoidea. The exact position of Aspidytes within this clade remains ambiguous: it is either the sister group of Amphizoidae or the sister group of a clade comprising this family and Hygrobiidae + Dytiscidae. The sister group relationship between Spanglerogyrinae and Gyrininae was strongly supported. The two included genera of Gyrinini form a clade, and Enhydrini are the sister group of a monophylum comprising the remaining Enhydrini and Orectochilini. A branching pattern (Peltodytes + (Brychius + Haliplus)) within Haliplidae was confirmed. Algophilus, Apteraliplus and the Haliplus‐subgenus Liaphlus form a clade. The generic status of the two former taxa is unjustified. The Phreatodytinae are the sister group of Noterinae, and Notomicrus (+ Speonoterus), Hydrocoptus, and Pronoterus branch off successively within this subfamily. The search for the larvae of Meru and a combined analysis of morphological and molecular data should have high priority. © The Willi Hennig Society 2006.     

Another perspective on cytoevolution in Lobelioideae (Campanulaceae)

The Lobelioideae is a cosmopolitan group whose cytoevolution is discussed on a model of primitively high diploid chromosome numbers, in which x = 14 is relatively plesiomorphic and x = 21 may be even more plesiomorphic. This model is suggested from the high frequency of lobelioid genera with x = 14, the probably plesiomorphic condition of x = 17 in the sister group Campanuloideae (Campanulaceae), and the primitive x = 15 in Stylidiaceae (Campanulales). It contrasts with that for a primitive x = 7 and paleopolyploidy to higher chromosome numbers. In our analysis, the genus Lobelia shows three broad cytoevolutionary groups, which probably have phylogenetic and infrageneric taxonomic significance: (1) woody diploids with x = 21 in Chile and woody diploids with x = 14 in Africa, Asia, and Hawaii; (2) herbaceous diploids with several series of dysploid chromosome numbers n = 19, 13, 12, 11, 10, 9, 8, 7, 6, mainly in Africa and Australia; (3) widespread and speciose herbaceous taxa based on a very derived n = 7, with recent frequent euploid rises (neopolyploidy) at or below the species level in subgenus Lobelia and allied or segregate genera. Other woody and herbaceous lobeliad genera have comparable cytoevolutionary patterns. New chromosome counts for Australian Lobelia, Pratia, and Isotoma illustrate the last two cytoevolutionary groups.     

On the thoracic anatomy of the Madagascan Heterogyrus milloti and the phylogeny of Gyrinidae (Coleoptera)

Gyrinidae is a group of beetles with a unique specialization of swimming on the water surface. Heterogyrus milloti Legros (Heterogyrinae) from Madagascar is a species with various preserved plesiomorphic features. The information on the morphology and biology was very limited until recently, and the thoracic anatomy remained largely unknown. Consequently, the aim of the present study is to describe external and internal thoracic features of Heterogyrus Legros in detail and to interprete them with respect to their phylogenetic and functional significance, with a special focus on the unusual flight apparatus of Gyrinidae. Characters documented with innovative techniques are compared to conditions found in other gyrinid genera and other groups of Adephaga, including characters of other body parts and larvae. A data matrix with 144 characters of adults, larvae and eggs was compiled and analysed cladistically. Gyrinidae excluding Spanglerogyrus Folkers (Heterogyrinae + Gyrininae) is supported by many apomorphies, mainly by a unique locomotor apparatus with paddle‐like middle and hind legs. The results confirm Heterogyrus as the earliest diverging branch in Gyrinidae except Spanglerogyrus, implying a sister‐group relationship between this genus and Gyrininae, a clade comprising Gyrinini, Dineutini and Orectochilini. The presence of an opening between the mesanepisternum and elytra, reduction of the lateral metafurcal arms, loss of the metathoracic M. furcacoxalis lateralis, and modifications of the head, including the dorsal shift of the upper subcomponent of the compound eyes, are synapomorphies of the three tribes. The monophyly of Gyrinini is moderately well‐supported, whereas Orectochilini is strongly supported by different characters including a highly simplified but functioning flight apparatus. A clade comprising Orectochilini and the dineutine genera is suggested by synapomorphies of adults and larvae. The monophyly of Dineutini was supported in a recent study, but not by the characters analysed here. Features of adults, larvae and eggs indicate that Gyrinidae are the sister group to the remaining adephagan families, as suggested in some earlier morphology‐based studies and recent analyses of large molecular datasets.     

Evolution of Spore Release Mechanisms in the Saprolegniaceae (Oomycetes): Evidence from a Phylogenetic Analysis of Internal Transcribed Spacer Sequences

Classical studies on spore release within the Saprolegniaceae (Oomycetes) led to the proposition that different mechanisms of sporangial emptying represent steps in an evolutionary transition series. We have reevaluated this idea in a phylogenetic framework using internal transcribed spacer sequences of four genera. These data were compared with the response to osmotic stress exhibited by each taxon.Saprolegniaemerges as the most basal genus, sister toAchlya, Thraustotheca,andDictyuchus. AchlyaandThraustothecaare most closely related, whileDictyuchusappears to have evolved along a separate evolutionary lineage. The resulting phylogenetic framework is consistent with the idea that the mechanism of sporangial emptying exhibited bySaprolegniarepresents the plesiomorphic condition from which the other mechanisms were derived independently. These alternative mechanisms of spore release may have resulted from a small number of mutations that inhibited axonemal development and altered the temporal and spatial expression of lytic enzymes that degrade the sporangial wall.     

Comparative osteology and phylogenetic systematics of fossil and living bony-tongue fishes (Actinopterygii, Teleostei, Osteoglossomorpha)

Several recent studies using analyses of morphological characters have addressed the interrelationships of Osteoglossomorpha, a group that sometimes is considered the living sister group of all other living teleosts. Many characters used in these studies were found to be poorly defined, to be coded incorrectly or illogically, or to display more variation than was described. The goal of this study is to address these concerns and contribute generally to knowledge of the morphology and systematic relationships of osteoglossomorphs. Analysis of 72 characters (65 informative) scored for 20 genera resulted in two most parsimonious cladograms (171 steps, CI = 0.6433, CI = 0.6139 excluding uninformative characters, HI = 0.3977, HI = 0.3861 excluding uninformative characters; RI = 0.7782; RC = 0.5006). Osteoglossomorpha is supported by both synapomorphies and homoplasies, although its monophyly was not truly tested in this analysis (only a single outgroup, Elops , was included in the analysis). The only difference in the topologies of these cladograms is in the position of ?Lycoptera (recovered as either the sister group of all other osteoglossomorphs sampled or of ?Eohiodon +Hiodon ). ?Ostariostoma is recovered as the sister group of all non‐hiodontiform osteoglossomorphs. Mormyrids are the sister group of notopterids + osteoglossids. This clade has not been found in other recent analyses. Mormyrids and notopterids usually are considered more closely related to each other than to any other group; characters not included here support this relationship and future consideration of these characters must be made. Although almost completely dichotomous, many nodes of the resulting trees lack rigorous support. For example, ?Palaeonotopterus is interpreted as the sister group of all mormyrids sampled, although for this taxon only 22% of characters could be scored. © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society , 2003, 137 , 1?100.     

Phylogenetic analysis of Trechitae (Coleoptera: Carabidae) based on larval morphology, with a description of first-instar Phrypeus and a key to genera

Abstract. Sixty-nine characters of larval structure of twenty-eight genera of the supertribe Trechitae (Coleoptera: Carabidae) were analysed phylogenetically. The monophyly of Trechitae is strongly supported with five unique synapomorphies. The monophyly of Zolini + Bembidiini + Pogonini is supported with two synapomorphies. We propose that the tribe Trechini is a sister group to them and its monophyly is supported with two unique synapomorphies. The inferred branching pattern of Trechini genera is (Perileptus + Thalassophilus) + (Amblystogenium + (Trechimorphus + (Trechus + Epaphius + Aepopsis + Trechisibus))); Perileptus is a member of Trechodina rather than Trechina. The monophyly of Zolini is not supported. The monophyly of Pogonini is supported with two unique synapomorphies; its sister group relationships remain obscure; the branching pattern of pogonine genera is (((Pogonus + Pogonistes) + Cardiaderus) + Thalassotrechus). No evidence for monophyly of the tribe Bembidiini (s. lato; including subtribes Bembidiina, Tachyina, Xystosomina, and Anillina) was found. The relationships of Phrypeus are obscure; no evidence could be found linking it with Bembidiina. Without Phrypeus, Bembidiina might be a monophylum with a single synapomorphy. Sinechostictus branches basal of (Bembidion + Asaphidion) and therefore should be treated as a separate genus. Tachyina and Xystosomina form a monophylum based on two unique synapomorphies; a close relationship with a monophyletic Anillina is suggested. Reduction of the number of claws from two to one in Trechitae has taken place twice: within Trechina (Trechus, Epaphius, Aepopsis and Trechisibus) and in (Zolini + Bembidiini + Pogonini). The previously unknown larvae of the isolated genus Phrypeus are described and illustrated. A key to all twenty-eight analysed Trechitae genera based on characters of larvae and a list of larval autapomorphies for each genus are provided.     

Phylogeny of the flyingfish family Exocoetidae (Teleostei, Beloniformes)

The phylogeny of the flyingfish family Exocoetidae is studied cladistically, using 41 morphological characters encompassing early life history, and external and internal features. The monophyly of the family is supported by 10 synapomorphies. Within the family,Oxyporhamphus is the sister group to all other genera, the monophyly of the latter being defined by 10 synapomorphies.Fodiator is the sister group of genera characterized by the presence of chin barbels in juveniles.Parexocoetus is the sister group ofExocoetus, Cypselurus, Prognichthys andHirundichthys, the latter being defined by four synapomorphies. In the latter group,Exocoetus is the sister group of the other three genera. The phylogeny of the Exocoetidae is characterized by the stepwise upgrading of gliding capability, with sequential modifications of the caudal, pectoral and pelvic fins. The subfamily Oxyporhamphinae is resurrected.     

Limb myology and phylogenetic relationships in the superfamily Dipodoidea (birch mice, jumping mice, and jerboas)

Limb muscles were dissected in seven genera, representing all six superfamilies, of dipodoid rodents and myoloic characters were used to construct a phylogenetic hpothesis of relationships within this cfade. Mologic differences among genera suported tie monophyly of the superfamily Dipodoidea reLtive to the outrou taxon and reveaEd thatSicista is the sister group to all other zapodid and dipodid enera. Tkis picement of Sicista differs markedly from its position in previous classifications where it has been regarded merely as a primitive zapodid genus. The phlograrn based on rnyologic characters also indicated that Cardiocranius is not a rimitive dipodid genus; it is the sister group to the subfamily Dipodinae. Although myologic differences among taxa were not sufficient to warrant the continued separation of zaodids and dipodids into two families, a new classification that places Sicista in its own family, ficistidae, and places the remaining zaodids and dipodids in the family Dipodidae, is proposed. Differences in karyology, genitaP morholoy, and postcranial osteological characters among dipodoid rodents are discussed in light or this pjylogeny.     

Comments on hexanchiform phylogeny (Pisces, Neoselachii)

The Hexanchiformes (Cow Sharks) are regarded as a monophyletic taxon. Cladistic analysis shows that among the various neoselachian taxa proposed so far as the sister group of the Hexanchiformes a sister group relationship between the Hexanchiformes and a (still unnamed) taxon comprising the Squaliformes and Pristiophoriformes appears as the most probable hypothesis. In addition, MAISEY and WOLFRAM'S (1984) concept of hexanchiform interrelationships is critically reviewed. An alternative cladogram of hexanchiform interrelationships is developed which includes Recent as well as fossil hexanchiform taxa. In this cladogram the living genera Hexancbus and Notorynchus are sister groups and both taxa together form the sister group of the Recent Heptranchias. The fossil taxa +Notidanoides, +“Hexanchus” gracilis, +Notidanodon and +Weltonia are arranged in the stem lineage of recent Hexanchiformes.     

Phylogenetics of the slipper orchids (Cypripedioideae, Orchidaceae): Nuclear rDNA ITS sequences

Cypripedioideae (Orchidaceae) have been the subject of numerous taxonomic treatments with conflicting interpretations of relationships among the five genera and the 150–170 species. We have produced nuclear ribosomal ITS nucleotide sequences for nearly 100 slipper orchid species and used parsimony analysis to investigate their relationships. Our results demonstrate that each genus, as currently circumscribed, is monophyletic (Mexipedium andSelenipedium being represented by a single taxon). LikerbcL data, ITS sequences placeMexipedium sister toPhragmipedium. Relationships at the sectional level inPaphiopedilum are largely as described byCribb. However, the division ofPaphiopedilum into subgg.Brachypetalum andPaphiopedilum is not supported; subg.Brachypetalum is paraphyletic to subg.Paphiopedilum. Phragmipedium species are divided into the same three major clades as in the taxonomic scheme ofMcCook. The plicate-leaved genera,Cypripedium andSelenipedium, are successive sister groups to the rest of the subfamily, confirming generally held opinions that they display plesiomorphic characters compared to the conduplicate-leaved genera. A survey of karyotypes in the context of the ITS tree reveals a general trend toward increased chromosome number, probably brought about by centric fission. These data also accord with a previously suggested biogeographic hypothesis of a widespread Northern Hemisphere distribution, followed by range fragmentation due to Miocene cooling.     

Development of a phylogenetic model for the tribe Micrarctiini (Lepidoptera, Arctiidae) by the SYNAP method

A phylogenetic model for all the 18 genera of Micrarctiini (Lepidoptera, Arctiidae) of the World fauna is discussed. The SYNAP method was used for development of the model. For these purposes, 47 characters of the general appearance, male and female genitalia, and wing pattern were used. Based on these characteristics, an evolutionary trend from the plesiomorphic to apomorphic states was revealed. The genera Apantesis and Amurrhyparia, as well as Notarctia, differ from the other genera to the greatest extent. The other genera have formed two the main clades: Grammia + (Diacrisia + (Rhyparia + Rhyparioides)). Neoarctia + (Palearctia + Holoarctia + Chelis) and all the other genera have occupied separate positions. The latter group was divided into two clades: Ebertarctia + Tancrea + Divarctia with brachypterous females and Centrarctia + (Sibirarctia + (Micrarctia + Hyperborea)).     

Phylogeny of the Acanthocephala based on morphological characters

Only four previous studies of relationships among acanthocephalans have included cladistic analyses, and knowledge of the phylogeny of the group has not kept pace with that of other taxa. The purpose of this study is to provide a more comprehensive analysis of the phylogenetic relationships among members of the phylum Acanthocephala using morphological characters. The most appropriate outgroups are those that share a common early cell-cleavage pattern (polar placement of centrioles), such as the Rotifera, rather than the Priapulida (meridional placement of centrioles) to provide character polarity based on common ancestry rather than a general similarity likely due to convergence of body shapes. The phylogeny of 22 species of the Acanthocephala was evaluated based on 138 binary and multistate characters derived from comparative morphological and ontogenetic studies. Three assumptions of cement gland structure were tested: (i) the plesiomorphic type of cement glands in the Rotifera, as the sister group, is undetermined; (ii) non-syncytial cement glands are plesiomorphic; and (iii) syncytial cement glands are plesiomorphic. The results were used to test an early move of Tegorhynchus pectinarius to Koronacantha and to evaluate the relationship between Tegorhynchus and Illiosentis. Analysis of the data-set for each of these assumptions of cement gland structure produced the same single most parsimonious tree topology. Using Assumptions i and ii for the cement glands, the trees were the same length (length = 404 steps, CI = 0.545, CIX = 0.517, HI = 0.455, HIX = 0.483, RI = 0.670, RC = 0.365). Using Assumption iii, the tree was three steps longer (length = 408 steps, CI = 0.539, CIX = 0.512, HI = 0.461, HIX = 0.488, RI = 0.665, RC = 0.359). The tree indicates that the Palaeacanthocephala and Eoacanthocephala both are monophyletic and are sister taxa. The members of the Archiacanthocephala are basal to the other two clades, but do not themselves form a clade. The results provide strong support for the Palaeacanthocephala and the Eoacanthocephala and the hypothesis that the Eoacanthocephala is the most primitive group is not supported. Little support for the Archiacanthocephala as a monophyletic group was provided by the analysis. Support is provided for the recognition of Tegorhynchus and Illiosentis as distinct taxa, as well as the transfer of T. pectinarius to Koronacantha.     

亚洲蝮亚科蛇属间系统发生支序分析 (蛇亚目：蝰科)

在大量比较形态研究的基础上,选择了５个特征方面２８个性状,以支序分析方法探讨了分布于亚洲蝮亚科蛇５属１４种的系统发生关系,结果表明：该亚科蛇类可以划分为３个不同的类群,第一个类群包括尖吻蝮属、瘤鼻蝮属、红口腹属,它们具有较多的祖征,代表了该科中原始一类,基中红口蝮可能是最原始的一属;亚洲蝮属单独形成一个类群;第三个类群为原广义的烙铁头蛇属,包括竹叶青蛇属、原矛蝮属、烙铁头蛇属、黑绿烙铁头蛇属、莽山     

The skull of the Upper Cretaceous snake Dinilysia patagonica Smith‐Woodward, 1901, and its phylogenetic position revisited

The cranial anatomy of Dinilysia patagonica, a terrestrial snake from the Upper Cretaceous of Argentina, is redescribed and illustrated, based on high‐resolution X‐ray computed tomography and better preparations made on previously known specimens, including the holotype. Previously unreported characters reinforce the intriguing mosaic nature of the skull of Dinilysia, with a suite of plesiomorphic and apomorphic characters with respect to extant snakes. Newly recognized plesiomorphies are the absence of the medial vertical flange of the nasal, lateral position of the prefrontal, lizard‐like contact between vomer and palatine, floor of the recessus scalae tympani formed by the basioccipital, posterolateral corners of the basisphenoid strongly ventrolaterally projected, and absence of a medial parietal pillar separating the telencephalon and mesencephalon, amongst others. We also reinterpreted the structures forming the otic region of Dinilysia, confirming the presence of a crista circumfenestralis, which represents an important derived ophidian synapomorphy. Both plesiomorphic and apomorphic traits of Dinilysia are treated in detail and illustrated accordingly. Results of a phylogenetic analysis support a basal position of Dinilysia, as the sister‐taxon to all extant snakes. The fossil taxa Yurlunggur, Haasiophis, Eupodophis, Pachyrhachis, and Wonambi appear as derived snakes nested within the extant clade Alethinophidia, as stem‐taxa to the crown‐clade Macrostomata. The hypothesis of a sister‐group relationship between Dinilysia and Najash rionegrina, as suggested by some authors, is rejected by the results of our analysis. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 194–238.     

The Phylogeny of the Myrmecophagidae (Mammalia, Xenarthra, Vermilingua) and the Relationship of Eurotamandua to the Vermilingua

A cladistic investigation of the phylogenetic relationships among the three extant anteater genera and the three undoubted extinct myrmecophagid genera is performed based upon osteological characteristics of the skull and postcranial skeleton. One hundred seven discrete morphological characters are analyzed using the computer program PAUP. Characters are polarized via comparison to the successive xenarthran outgroups Tardigrada (represented by the living sloth Bradypus) and Cingulata (represented by the recent armadillos Dasypus and Euphractus). The analysis results in a single most-parsimonious tree (TL = 190, CI = 0.699, RI = 0.713). The tree corroborates the monophyly of the subfamilies Cyclopinae and Myrmecophaginae, the former including the extant Cyclopes and the Pliocene genus Palaeomyrmidon. Within the Myrmecophaginae the Miocene genus Protamandua is the sister taxon to a clade including the remaining three genera. The recent Tamandua is in turn the sister taxon to the extant Myrmecophaga plus the Pliocene genus Neotamandua. Contrary to the suggestions of recent authors, weak support is provided for the taxonomic distinctiveness of the latter genus from the recent Myrmecophaga. The monophyly of the Myrmecophagidae is supported by 15 unequivocal synapomorphies. The monophyly of the Cyclopinae and Myrmecophaginae is supported by 3 and 13 unambiguous synapomorphies, respectively. The enigmatic Eocene genus Eurotamandua, from the Messel fauna of Germany, is coded for the 107 morphological characters above and included in two subsequent PAUP analyses. The palaeanodont Metacheiromys is also added to these two analyses as a nonxenarthran outgroup to test for the possibility that Eurotamandua lies outside the Xenarthra. In the first analysis, Eurotamandua is constrained a priori to membership in the Vermilingua. The single most-parsimonious tree (TL = 224, CI = 0.618) that results places Eurotamandua as the sister group to the remaining anteater genera, contra Storch and Habersetzer's (1991) assignment of Eurotamandua to the vermilinguan subfamily Myrmecophaginae. Eurotamandua shares six unequivocal synapomorphies with other anteaters, including the absence of teeth and the presence of a lateral tuberosity on the fifth metatarsal. The remaining vermilinguans are united by 11 unequivocal synapomorphies, plus an additional 10 ambiguous synapomorphies. In the second analysis, the position of Eurotamandua is unconstrained. The resulting single most-parsimonious tree (TL = 219, CI = 0.632) places Eurotamandua outside Vermilingua as the sister group to the Pilosa (Vermilingua plus Bradypus). The monophyly of this node is supported by four unambiguous synapomorphies in the unconstrained analysis. Further manipulation of this second analysis shows that placement of Eurotamandua as the sister group to the Xenarthra or to the Palaeanodonta adds three steps to the shortest tree but is more parsimonious than its placement as a sister group to the Vermilingua is the previous analysis. The addition of pangolins to the analysis does little to alter the major phylogenetic conclusions of the study. The allocation of Eurotamandua to the Xenarthra, but as a sister group to the Pilosa, is a novel arrangement which leaves open the biogeographic question of how a xenarthran reached Western Europe during the Eocene.     

Evolution of the skeleton and musculature of the male genitalia in the family lycaenidae (Lepidoptera): II. Infratribe Polyommatina Swainson, 1827

Skeleton and musculature of valvae were examined in 19 species from the section Polyommatus (Lepidoptera, Lycaenidae) revised by Bálint and Johnson (1997). Functional organization of male genitalia in the section Polyommatus was shown to be relatively constant. Significant interspecific variation of the skeleton and muscles was found only in the structure of valvae. During their evolution, the valvae became elongated, the costal torsion and longitudinal medial fold appeared on their medial wall, the subcostal groove was formed, and the intravalval muscle differentiated into two muscles forming a chiasma. This muscular chiasma was formed at least twice in the evolution of the group, in Polyommatus Latreille, 1804 + Aricia [Reichenbach], 1819 and in Plebeius Kluk, 1802. The genus Chilades Moore, [1881] is the least advanced as concerns the functional organization of the valvae. The genera Polyommatus and Aricia are the most advanced and constitute a sister pair. The intermediate position is occupied by Plebeius Kluk, 1802 and probably Madeleinea Bálint, 1993. The genus Albulina Tutt, 1809 is a compound taxon, its species occupying an intermediate position between Polyommatus and Aricia on the one hand, and Plebeius on the other hand. With a sole exception, they are characterized by a plesiomorphic state of the valval musculature that does not allow their mutual phylogenetic relations to be evaluated consistently. The only examined species of the genus Agriades Hübner, [1819], A. glandon (De Prunner, 1798), also possesses undifferentiated musculature, but according to the characters of the vaginal area, its females are close to the branch Polyommatus + Aricia.     

Phylogeny,taxonomy, and evolution of Neotropical cichlids (Teleostei: Cichlidae: Cichlinae)

Despite recent progress on the higher‐level relationships of Cichlidae and its Indian, Malagasy, and Greater Antillean components, conflict and uncertainty remain within the species‐rich African, South American, and Middle American assemblages. Herein, we combine morphological and nucleotide characters from the mitochondrial large ribosomal subunit, cytochrome c oxidase subunit I, NADH dehydrogenase four, and cytochrome b genes and from the nuclear histone H3, recombination activating gene two, Tmo‐4C4, Tmo‐M27, and ribosomal S7 loci to analyse relationships within the Neotropical cichlid subfamily Cichlinae. The simultaneous analysis of 6309 characters for 90 terminals, including representatives of all major cichlid lineages and all Neotropical genera, resulted in the first well‐supported and resolved generic‐level phylogeny for Neotropical cichlids. The Neotropical subfamily Cichlinae was recovered as monophyletic and partitioned into seven tribes: Astronotini, Chaetobranchini, Cichlasomatini, Cichlini, Geophagini, Heroini, and Retroculini. Chaetobranchini + Geophagini (including the “crenicichlines”) was resolved as the sister group of Heroini + Cichlasomatini (including Acaronia). The monogeneric Astronotini was recovered as the sister group of these four tribes. Finally, a clade composed of Cichlini + Retroculini was resolved as the sister group to all other cichlines. The analysis included the recently described ?Proterocara argentina, the oldest known cichlid fossil (Eocene), which was placed in an apical position within Geophagini, further supporting a Gondwanan origin for Cichlidae. These phylogenetic results were used as the basis for generating a monophyletic cichline taxonomy. © The Willi Hennig Society 2008.