On the evolution and morphology of the rotiferan trophi, with a cladistic analysis of Rotifera

The phylogeny of Rotifera was examined in different computer‐generated cladistic analyses, including Seisonidea, Bdelloidea, Flosculariacea, Collothecacea and all ploimids treated on family level. The analyses were based on a character matrix solely dealing with morphological characters, primarily based on the trophi morphology. Limnognathia maerski (Micrognathozoa), Rastrognathia macrostoma and Gnathostomula paradoxa (Gnathostomulida) were used as outgroups. The cladistic analysis performed by paup produced 288 most parsimonious trees. peewee analyses produced between 140 and 432 trees, depending on the concavity value. The monophyly of Eurotatoria, Monogononta and Ploima was confirmed in all obtained trees. All analyses suggested a division of Ploima into major clades. One clade corresponded to Transversiramida while the other contained all other ploimid taxa and recognized Antrorsiramida as a monophylum. Based on the obtained results a scenario for the trophi evolution is proposed. The analyses suggested that the presence of an incus is synapomorphic for Gnathifera while mallei are synapomorphic for Micrognathozoa and Rotifera. The ancestral rotifer trophi probably resembled those in Harringia (Asplanchnidae).     

Gnathostomulid phylogeny inferred from a combined approach of four molecular loci and morphology

The phylogeny of the obscure metazoan phylum Gnathostomulida has previously only been addressed with cladistic analyses of morphological data. In the present study DNA sequence data from four molecular loci, including 18S rRNA, 28S rRNA, histone H3 and cytochrome c oxidase subunit I, are added to a revised morphological data matrix. The data set represents 23 gnathostomulid species that are analyzed under direct optimization using parsimony as the optimality criterion. The results obtained from analyzing the four molecular loci and combined morphological and molecular data under different parameter sets are generally very congruent, and differ only on minor points. The results clearly support gnathostomulid monophyly, as well as the basal division of Gnathostomulida into Filospermoidea and Bursovaginoidea. Filospermoidea were represented by species of Haplognathia and Cosmognathia, and generic monophyly is supported for both groups. Within Bursovaginoidea, Conophoralia (= Austrognathiidae) and Scleroperalia appear as sister groups. Monophyly of Mesognathariidae was confirmed as well, whereas the relationships between species of Gnathostomulidae and Onychognathiidae were contradicted by the molecular data when compared to morphological observations. ©The Willi Hennig Society 2006.     

A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae)

We present the first cladistic analysis focused at the tribal and subfamily level of the orb-weaving spider family Araneidae. The data matrix of 82 characters scored for 57 arancid genera of 6 subfamilies and 19 tribes (and 13 genera from 8 outgroup families) resulted in 16 slightly different, most parsimonious trees. Successive weighting corroborated 62 of the 66 informative nodes on these cladograms; one is recommended as the 'working' araneid phylogcny. The sister group of Araneidae is all other Araneoidea. Araneidae comprises two major clades: the subfamily Araneinae, and the 'argiopoid' clade, which includes all other subfamilies and most tribes (((Gasteracanthinae, Caerostreae), (((Micratheninae, Xylcthreae), Eruyosaccus ), (Eurycorminae, Arciinae)), Cyrlarachninae), ((Argiopinae, Cyrtophorinae), Arachnureae)). Cyrtarachneae and Mastophoreae are united in a new subfamily, Cyr-tarachninae. The spiny orb-weavers alone (Gasteracanthinae and Micratheninae) are not monophyletic. The mimetid subfamily Arciinae and the 'tetragnathid' genus Zygiella are araneids, but .Nephila (and other tetragnathids) are not. On the preferred tree, web decorations (stabilimenta) evolved 9 times within 15 genera, and were lost once. The use of silk to subdue prey evolved once in cribellate and four times in ecribillate orb weavers. Sexual size dimorphism evolved once in nephilines, twice in araneids, and reverted to monomorphism five times. Evolution in other genitalic and somatic characters is also assessed; behavioral and spinneret features arc most consistent (male genitalia, leg and prosomal features least consistent) on the phylogeny.     

螟黄足盘绒茧蜂复合群支序系统学研究（膜翅目：茧蜂科：小腹茧蜂亚科）

对螟黄足盘绒茧蜂复合群Cotesia flavipes complex (膜翅目：茧蜂科：小腹茧蜂亚科)分布全世界的5个种和外群侧沟茧蜂Microplitis 及荻茧蜂Diolcogaster 的25个性状,以及复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂Cotesia kariyai、粉蝶盘绒茧蜂Cotesia glomerata 的24个性状,分别进行比较研究,并运用支序分析的方法探讨该复合群内5个种种间的系统发育关系。支序分析表明螟黄足盘绒茧蜂复合群是一单系群,二化螟盘绒茧蜂C. chilonis 和大螟盘绒茧蜂C. sesamiae 近缘,芦螟盘绒茧蜂C. chiloluteelli 和汉寿盘绒茧蜂C. hanshouensis 近缘,螟黄足盘绒茧蜂C. flavipes相对独立。 以上研究表明无论是以近缘属作外群还是以同属其它种作外群,所得结果基本上都能反映螟黄足盘绒茧蜂复合群各种之间的分类地位。     

Phylogeny and biogeography of the Vitrinidae (Gastropoda: Stylommatophora)

The phylogeny of the Vitrinidae is reconstructed in a cladistic analysis based on characters of the genitalia, the copulation behaviour and the radula. The genera with an atrial stimulator turned out to be the earliest branches of the Vitrinidae, whereas the genera with a glandula amatoria form a monophyletic, taxonomically apomorphic group. The differences between the proposed phylogeny and previous hypotheses are discussed. The ancestral areas of the Vitrinidae and its sister group, the limacoid slugs Boettgerillidae–Limacidae–Agriolimacidae, are estimated using weighted ancestral area analysis. The Vitrinidae and the limacoid slugs might have originated by a vicariance event between Central Europe and the Near East. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 134 , 347–358.     

Cladistic analysis of the Lythraceae sensu lato based on morphological characters

GRAHAM, S. A., CRISCI, J. V. & HOCH, P. C, 1993. Cladistic analysis of the Lythraceae sensu lato based on morphological characters. The Lythraceae s. I. comprise 31 mostly highly distinctive genera distributed worldwide. Cladistic analyses of the family were undertaken using 26 characters from anatomy, floral morphology, pollen, and seed morphology. Of four outgroup hypotheses, the Onagraceae offered the best heuristic assessment, generating 3746 trees with a tree length of 83 and consistency index and (C.I.) of 0.41. A strict consensus tree from the 3746 trees produced a basal monophyletic group of five genera (Sonneratia, Duabanga, Punka, Lagerstroemia, Lawsonia) characterized by anthotelic (determinate) inflorescences and wet stigmas, and a second clade with blastotelic (indeterminate) inflorescences, dry stigmas, and reduced carpel number. Successive weighting with Onagraceae as outgroup generated five most parsimonious trees with a tree length of 164 and C.I. of 0.75. Internal branches are weakly supported by only seven non-homoplasious characters. Significant results of the analysis include: (1) recognition of the two major clades and a total of seven monophyletic groups within the family; (2) the paraphyly of subfamily Lythroideae (Lythraceae s. s .); and (3) indication that genera endemic to the New World have been derived from more than one ancient Old World evolutionary line. The current taxonomic classification of the family is not closely correlated with results of the cladistic analysis.     

Phylogeny of fossil and extant glypheid and litogastrid lobsters (Crustacea,Decapoda) as revealed by morphological characters

A phylogenetic analysis of a total of 31 species: 27 fossil species from seven families (Glypheidae, Litogastridae, Mecochiridae, Pemphicidae, Erymidae, Clytiopsidae, Chimaerastacidae), and four extant species from three families (Glypheidae, Nephropidae, Stenopodidae) is proposed. Most of the genera considered are coded exclusively based upon their type species and, as much as possible, based upon the type specimens. The cladistic analysis demonstrates that the glypheidean lobsters (infraorder Glypheidea) form a monophyletic group including two superfamilies: Glypheoidea and Pemphicoidea new status. Glypheoidea includes three families: Glypheidae, Mecochiridae and Litogastridae. Litogastridae is the sister group of the clade Glypheidae + Mecochiridae. Pemphicoidea includes a single family: Pemphicidae. A new classification of Glypheidea is proposed and currently known genera are rearranged based upon the phylogenetic analysis.     

Cladistic analysis of Enkianthus with notes on the early diversification of the Ericaceae

The genus Enkianthus (Ericaceae) has been subject to a cladistic analysis of morphological, anatomical, embryological, and cytological data, using two species of Clethra as outgroup. In order to evaluate the monophyly of Enkianthus , the genera Epigaea and Phyllodoce from the subfamily Rhododendroideae, and the genus Andromeda from the subfamily Vaccinioideae were also included in an initial analysis which yielded 20 equally parsimonious cladograms. The results indicated that Enkianthus is monophyletic, and a subsequent analysis with only the outgroup taxa and 14 species of Enkianthus yielded two equally parsimonious cladograms with a more resolved topology. In order to obtain strictly monophyletic taxa, the current subgeneric classification of Enkianthus is revised, recognizing four sections: Enkianthus (9 spp.), Andromedina (2 spp.), Meisteria (2 spp.), and Enkiantella (4 spp.). The paper provides a key, illustrations of representative taxa, a cladogram, and strict consensus trees.     

Molecular systematics of Iridaceae: evidence from four plastid DNA regions

Iridaceae are one of the largest families of Lilianae and probably also among the best studied of monocotyledons. To further evaluate generic, tribal, and subfamilial relationships we have produced four plastid DNA data sets for 57 genera of Iridaceae plus outgroups: rps4, rbcL (both protein-coding genes), the trnL intron, and the trnL-F intergenic spacer. All four matrices produce similar although not identical trees, and we thus analyzed them in a combined analysis, which produced a highly resolved and well-supported topology, in spite of the fact that the partition homogeneity test indicated strong incongruence. In each of the individual trees, some genera or groups of genera are misplaced relative to morphological cladistic studies, but the combined analysis produced a pattern much more similar to these previous ideas of relationships. In the combined tree, all subfamilies were resolved as monophyletic, except Nivenioideae that formed a grade in which Ixioideae were embedded. Achlorophyllous Geosiris (sometimes referred to Geosiridaceae or Burmanniaceae) fell within the nivenioid grade. Most of the tribes were monophyletic, and Isophysis (Tasmanian) was sister to the rest of the family; Diplarrhena (Australian) fell in a well-supported position as sister to Irideae/Sisyrinchieae/Tigridieae/Mariceae (i.e., Iridoideae); Bobartia of Sisyrinchieae is supported as a member of Irideae. The paraphyly of Nivenioideae is suspicious due to extremely high levels of sequence divergence, and when they were constrained to be monophyletic the resulting trees were only slightly less parsimonious (<1.0%). However, this subfamily also lacks clear morphological synapomorphies and is highly heterogeneous, so it is difficult to develop a strong case on nonmolecular grounds for their monophyly.     

Phylogenetic relationship among Libellula, Ladona and Plathemis (Odonata: Libellulidae) based on DNA sequence of mitochondrial 16S rRNA gene

Abstract. The type genus for the dragonfly family Libellulidae is Libellula. At present, Libellula s.l. includes twenty-nine species, whose distribution is largely Nearctic. Whether two other libellulid taxa, Ladona and Plathemis, should be considered synonyms of Libellula, subgenera of Libellula, or separate genera, has been a subject of intermittent debate for over a century. Earlier proposals concerning Ladona and Plathemis were based on a limited number of morphological characters and lacked rigorous phylogenetic analyses. Therefore, we used the DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony, maximum likelihood and neighbour-joining analyses to explore whether Ladona and Plathemis are monophyletic lineages distinct from Libellula. We obtained ≈ 415 bp of DNA sequence from twenty-three taxa including thirteen species of Libellula s.s., all three recognized species of Ladona, the two species of Plathemis and representatives of four other libellulid genera. Tetragoneuria williamsoni (Odonata: Corduliidae) was included as the outgroup. Parsimony analysis suggested that Ladona and Plathemis are monophyletic lineages distinct from Libellula s.s. with a sister group relationship between Libellula and Ladona. The monophyly of Ladona, Plathemis and Libellula was supported in > 90% of bootstrap replications and in trees five to ten steps longer than the most parsimonious trees. Relationships inferred from maximum likelihood and neighbour-joining analyses also supported the monophyly of Ladona and Plathemis. The four other libellulid genera included in the study formed a monophyletic clade distinct from Libellula, Ladona and Plathemis. Based on our analysis, we propose that Ladona and Plathemis be considered either genera or subgenera within Libellulidae.     

Phylogenetic structure in the grass family (Poaceae) as inferred from chloroplast DNA restriction site variation

A cladistic analysis of chloroplast DNA restriction site variation among representatives of all subfamilies of the grass family (Poaceae), using Joinvillea (Joinvilleaceae) as the outgroup, placed most genera into two major clades. The first of these groups corresponds to a broadly circumscribed subfamily Pooideae that includes all sampled representatives of Ampelodesmeae, Aveneae, Brachypodieae, Bromeae, Diarrheneae, Meliceae, Poeae, Stipeae, and Triticeae. The second major clade includes all sampled representatives of four subfamilies (Panicoideae [tribes Andropogoneae and Paniceae], Arundinoideae [Arundineae], Chloridoideae [Eragrostideae], and Centothecoideae [Centotheceae]). Within this group (the “PACC” clade), the Panicoideae are resolved as monophyletic and as the sister group of the clade that comprises the other three subfamilies. Within the latter group, Danthonia (Arundinoideae) and Eragroslis (Chloridoideae) are resolved as a stable monophyletic group that excludes Phragmites (Arundinoideae); this structure is inconsistent with the Arundinoideae being monophyletic as currently circumscribed. The PACC clade is placed within a more inclusive though unstable clade that includes the woody Bambusoideae (Bambuseae) plus several disparate tribes of herbaceous grasses of uncertain affinity that are often recognized as herbaceous Bambusoideae (Brachyelytreae, Nardeae, Olyreae, Oryzeae, and Phareae). Among eight most-parsimonious trees resolved by the analysis, four include a monophyletic Bambusoideae sensu lato (comprising Bambuseae and all five of these herbaceous tribes) as the sister group of the PACC clade; in the other four trees these bambusoid elements are not resolved as monophyletic, and the PACC clade is nested among these tribes. These results are consistent with those of previous analyses that resolve a basal or near-basal branch within the family between Pooideae and all other grasses. However, resolution by the present analysis of the PACC clade, which includes Centothecoideae, Chloridoideae, and Panicoideae, but excludes Bambusoideae, is inconsistent with the results of previous analyses that place Bambusoideae and Panicoideae in a monophyletic group that excludes Centothecoideae and Chloridoideae.     

小腹茧蜂亚科的雄外生殖器及族级单元系统发育的研究

对分布在东洋区和古北区的小腹茧蜂亚科 (膜翅目: 茧蜂科)21个属的67个种及外群折脉茧蜂属 (膜翅目: 茧蜂科) 2个种的雄外生殖器的5个性状进行了比较研究。在形态学研究的基础上,通过选用头部、胸部和腹部(包括雌雄外生殖器的性状) 等34个性状,运用支序分析的方法探讨了分布在东洋区和古北区的小腹茧蜂亚科21个属以及它们所属的族间的系统发育关系,并对Mason (1981) 的分类系统进行了重新评价。雄外生殖器和支序分析基本上证实并恢复了由Mason (1981) 确定的2个主要分支,即绒茧蜂族Apantelini+小腹茧蜂族Microgastrini和拱茧蜂族Fornicini+盘绒茧蜂族Cotesiini+侧沟茧蜂族MicroplitinI。绒茧蜂族Apantelini、拱茧蜂族Fornicini和侧沟茧蜂族Microplitini为单系群也被支持,但小腹茧蜂族Microgastrini和盘绒茧蜂族Cotesiini是否为单系群尚难于在树形图中体现,而且族内各属间的分支关系有变动。因此,尽管Mason的族级分类单元有一些欠缺,但仍是可信、实用的,不同意Walker等 (1990) 认为不应再使用Mason分族系统的观点。     

Molecular systematics of the Catesbaeeae-Chiococceae complex (Rubiaceae): flower and fruit evolution and biogeographic implications

The classification of the Catesbaeeae and Chiococceae tribes, along with that of the entire Rubiaceae, has long been debated. The Catesbaeeae-Chiococceae complex (CCC) includes approximately 28 genera and 190 species primarily concentrated in the Greater Antilles (nearly 70% of the species), Central and South America, and in the western Pacific (three genera). Previous molecular studies, with broad sampling of the Rubiaceae, have shown the CCC to be a monophyletic group. The present study is a more detailed examination of the generic relationships within the CCC using two data sets, the nuclear ribosomal ITS regions and the trnL-F chloroplast intron and spacer. Maximum parsimony analyses lend further support to the previous hypotheses that the CCC is monophyletic and sister to Strumpfia maritima. However, within the complex several genera do not form monophyletic groups. Previous studies of the Rubiaceae suggest that the ancestral fruit type in the CCC is a multiseeded capsule. Indehiscent, fleshy fruits appear to have evolved three to four times within this lineage. Changes in floral morphologies within the complex tend to correspond to cladogenesis among and within genera. Finally, molecular analyses suggest one or possibly two long-distance dispersals from the Americas to the western Pacific.     

Molecular evidence for repetitive parallel evolution of shell structure in Clausiliidae (Gastropoda, pulmonata)

The division of clausiliid genera, using the type of clausilial apparatus (CA) as the decisive criterion, is ambiguous. Two types of CA can be distinguished: the normal (N) type and the Graciliaria (G) type. Morphological resemblance between species with different types of CA led to the hypothesis that the CA type is homoplasious. Therefore sequence variation, phylogenetic relationships, and the evolution of the CA were studied in the genera Albinaria, Isabellaria, and Sericata. Phylogenetic relations were inferred from parsimony and neighbor-joining analyses of the nucleotide sequences of both internal transcribed spacers (ITS1 and ITS2) of the rDNA of 36 species. The variation among the sequences was great: 21.8% of the sequences were ambiguously aligned and excluded from the analysis. A high GC content in the unambiguously aligned portions and a substitutional bias toward a higher GC content are indicators of substitutional constraints in the spacers. We analyzed the data in several ways: using both spacers together and separately, weighting all mutations equally, correcting for transition/transversion bias by weighting, and using transversions only. In all resulting trees, Isabellaria is not a monophyletic group. Its division into two clades is supported by over 40 mutations and one large indel. Clade 1 consists of Isabellaria and Sericata and clade 2 consists of Isabellaria and Albinaria species. The present distribution of the CA type was plotted on the tree and its most parsimonious evolution was reconstructed. The CA type was shown to be highly homoplasious. In clade 1 and clade 2 both types of CA were found; depending on the ancestral state, either the G or the N type evolved several times in parallel. These results contribute decisively to the current debate on the morphological diagnoses of Albinaria, Sericata, and Isabellaria as monophyletic taxa.     

水蛇亚科形态学特征的支序分析

水蛇亚科属于游蛇科,包含10个属。其中7个属为单型属。选取水蛇亚科14个形态学特征进行支序分析,并利用计算机软件Hennig86对水蛇亚科中8个属之间的系统发育关系进行初步探讨,结果显示水蛇亚科分为两支Gerarda和Fordonia两个属构成姊妹群,Cerberus、Erpeton和Homalopsis三个属也构成单系群,与Vorisetal(2002)的分子系统树相同,但Cantoria属的地位则与Vorisetal(2002)的明显不同。     

Pattern of Phylogenetic Relationships among Members of the Tribe Melitaeini (Lepidoptera: Nymphalidae) Inferred from Mitochondrial DNA Sequences

We report a cladistic analysis of 77 butterfly species of the tribe Melitaeini (Lepidoptera: Nymphalidae) based on mitochondrial DNA gene sequences. We sequenced ca. 536 bp from the 16S ribosomal DNA (rDNA) and a 1422-bp sequence from the cytochrome oxidase I gene. Alignments are critical to statements of homology, especially when aligning rDNA sequences. We aligned the 16S sequences using conventional methods and direct optimization. We found that direct optimization of the sequences produced the best alignments and our preferred phylogenetic hypothesis. Our results suggest that many of the previously proposed genera are paraphyletic and we conclude that there are four monophyletic groups of species in our cladogram: the Euphydryas group, the Phyciodes group, the Chlosyne group, and the Melitaea group. The following genera are found to be paraphyletic: Castilia, Chlosyne, Didymaeformia, Eresia, Melitaea , and Thessalia . In addition, recognition of the monophyletic genera Cinclidia, Mellicta , and Telenassa would render other genera paraphyletic. Our phylogenetic hypothesis indicates that the melitaeines originated in the Nearctic and have colonized the Neotropics three times and the Palaearctic twice.     

Phylogeny of the World Anthophorini (Hymenoptera: Apoidea: Apidae)

Abstract.  The phylogenetic relationships of members of the long-tongued bee tribe Anthophorini were investigated. Twenty-six ingroup taxa, representing all known genera and the most common subgenera of the Anthophorini, were included in a cladistic analysis of the tribe based on 51 morphological characters. The analysis yielded the generic relationships: (( Habrophorula , Elaphropoda ) ( Habropoda ( Deltoptila ( Pachymelus ( Amegilla , Anthophora ))))). The monophyly of the Anthophorini and each of its genera was confirmed. A second tribe, Habropodini, as postulated by some authors, is paraphyletic. On the basis of the results of the cladistic analysis and the biogeographical data of Anthophorini, an evolutionary scenario is postulated in which all genera of Anthophorini probably evolved in the Late Cretaceous, except Amegilla and Anthophora , which originated in the Oligocene. The northern part of south-east Asia (India to south-east China) is regarded as the most probable place of origin and radiation of the Anthophorini. The New World was probably colonized three times independently by Habropoda (upper Cretaceous to Tertiary), by an ancestral lineage of Deltoptila (upper Cretaceous to Tertiary) and by Anthophora (Tertiary to Quaternary).     

Phylogeny and classification of Oleaceae based on rps16 and trnL-F sequence data

Phylogenetic relationships among 76 species of Oleaceae, representing all 25 recognized genera of the family, were assessed by a cladistic analysis of DNA sequences from two noncoding chloroplast loci, the rps16 intron and the trnL-F region. Consensus trees from separate and combined analyses are congruent and agree well with nonmolecular data (chromosome numbers, fruit and wood anatomy, leaf glycosides, and iridoids). The two debated genera Dimetra and Nyctanthes, previously suggested to belong to Verbenaceae (sensu lato) or Nyctanthaceae, are shown to belong to Oleaceae, sister to the hitherto genus incertae sedis Myxopyrum. This clade is also supported by anatomical and chemical data. The subfamily Jasminoideae is paraphyletic, and a new classification is presented. The subfamily level is abandoned, and the former Jasminoideae is split into four tribes: Myxopyreae (Myxopyrum, Nyctanthes, and Dimetra), Fontanesieae (Fontanesia), Forsythieae (Abeliophyllum and Forsythia), and Jasmineae (Jasminum and Menodora). The tribe Oleeae (previous subfamily Oleoideae) is clearly monophyletic, comprising the subtribes Ligustrinae (Syringa and Ligustrum), Schreberinae status novus (Schrebera and Comoranthus), Fraxininae status novus (Fraxinus), and Oleinae (12 drupaceous genera). An rps16 sequence obtained from Hesperelaea, known only from the type specimen collected in 1875, confirmed the placement of this extinct taxon in the subtribe Oleinae.     

Morphology and evolution of the nervous system in Gnathostomulida (Gnathifera,Spiralia)

Within Spiralia, Gnathifera may represent the deepest branching lineage comprising the jaw worms Gnathostomulida and their sister group Micrognathozoa + Syndermata. Yet, very few nervous system studies have been conducted on this lineage of microscopic, jaw-bearing worms, limiting our understanding of the evolution of this organ system in Spiralia. The nervous system of representatives from all major groups of Gnathostomulida was here mapped using confocal laser scanning microscopy and immunohistochemistry. Their intra-epidermal, unsegmented nervous systems comprise an anterior brain and three to five ventral and two to four dorsal longitudinal nerves, connected by few transverse commissures. Neurites of the stomatogastric nervous system were found lining the pharynx and connecting to a prominent buccal ganglion. Supposedly, sensory ciliated cells in the pharynx and the gut were documented for the first time. Based on these morphological results, primary homologies of neural structures in Gnathostomulida and other Gnathifera were hypothesized and thereafter tested using parsimony. This first neurophylogeny of Gnathostomulida resulted in a topology congruent with molecular data, supporting the monophyly of Bursovaginoidea, Conophoralia, and Scleroperalia. From this topology, the evolution of the gnathostomulid nervous system was reconstructed. It suggests a specialization and diversification of cords and serotonin-like immunoreactive cell patterns from a plesiomorphic neuroarchitecture of three unsegmented nerve cords and a compact anterior brain and buccal ganglion. These plesiomorphic states resemble the nervous system of Micrognathozoa, and possibly the ancestral states of Spiralia.