Morphology, development and probable systematic position of Cercariaeum crassum Wesenberg-Lund, 1934 (Digenea), a parasite of Pisidium amnicum in eastern Finland

Cercariaeum crassum Wesenberg-Lund, 1934 is redescribed at the cercariaeum stage and the daughter-rediae and cercaria are also described on the basis of new material from Pisidium amnicum collected in the Liikasepuro River (eastern Finland). The species is allocated to the family Allocreadiidae, although its generic affiliation remains unknown. The probable life-cycle (based on the developmental stages observed in daughter-redia) appears to eliminate the cercarial stage and, instead, a cercariaeum (a type of cercaria without a tail) may develop directly from germ balls or, rarely, through the stages of an ophthalmoxiphidiocercaria that transforms into a young caudate cercariaeum. Their morphology and development are shown to be consistent with the family Allocreadiidae. The probable lack of a second intermediate host in the life-cycle is discussed.     

Monophyly and systematic position of Glypthelmins (Digenea), based on partial lsrDNA sequences and morphological evidence

Species composition and systematic placement within the order Plagiorchiida has been controversial. Species number in Glypthelmins Stafford, 1905, a genus of cosmopolitan parasites of anurans, has varied between 19 and 28 species, depending on the taxonomic treatment. The present study performs a phylogenetic analysis using partial lsrDNA sequences to test the monophyly of the genus, and compares new sequences obtained with those published for different plagiorchiids to clarify the systematic position of Glypthelmins within the order Plagiorchiida. Maximum parsimony (MP) and maximum likelihood (ML) analyses result in identical tree topology. The single MP tree (L=1587, CI=0.40, RI=0.76) includes several clades with high bootstrap and Bremer support values. Glypthelmins sensu lato as traditionally classified is paraphyletic. Based on molecular and/or morphological evidence, the taxonomic diagnosis for Glypthelmins is emended, only eight species are retained in the genus, and re-establishment of the genera Choledocystus Pereira & Cuocolo, 1941 and Rauschiella Babero, 1951 is proposed, resulting in the following new combinations: Choledocystus simulans (Teixeira de Freitas, 1941) comb. nov., C. vitellinophilum (Dobbin, 1958) comb. nov.; Rauschiella chaquensis (Mañé-Garzón & Holcman-Spector, 1967) comb. nov., R. lenti (Teixeira de Freitas, 1941) comb. nov., R. linguatula (Rudolphi, 1819) comb. nov., R. poncedeleoni (Razo-Mendivil & León-Règagnon, 2001) comb. nov., R. robusta (Brooks, 1976) comb. nov., R. rugocaudata (Yoshida, 1916) comb. nov., R. staffordi (Tubangui, 1928) comb. nov. In the phylogenetic reconstruction, Glypthelmins sensu stricto forms the sister group of Haematoloechus Looss, 1899.     

Phylogenetic analysis of the suborder plagiorchiata (Platyhelminthes, Digenea) based on partial lsrDNA sequences

The phylogenetic relationships and systematic position of the members of the suborder Plagiorchiata, one of the derived and most diverse groups of Digenea, have always been controversial. Here, we present a phylogeny of this group based on the analysis of partial sequences of the lsrDNA in 28 species representing 13 families of Plagiorchiata, as well as four outgroups. Our results show that the Plagiorchiata, as considered by most authors, is not monophyletic, and that the superfamilies Opecoeloidea, and most probably Dicrocoelioidea and Gorgoderoidea, may have to be removed from this suborder. According to our results, the Plagiorchiata includes only parasites of terrestrial vertebrates. We find the Plagiorchiata to be composed of two well-supported clades which can be ranked as superfamilies: (1) Plagiorchioidea, including the Plagiorchiidae, Haematoloechidae, Telorchiidae, Brachycoeliidae and Leptophallidae; and (2) Microphalloidea containing the Microphallidae, Prosthogonimidae, Lecithodendriidae and Pleurogenidae. The genetic analysis also allowed revision of the position of several taxa of Plagiorchiata, including: (1) a confirmation of the position of the Brachycoeliidae within the Plagiorchiata; (2) a close phylogenetic relationships of Macrodera with Paralepoderma, Leptophallus and Metaleptophallus; (3) the grouping of Opisthioglyphe and Telorchis within a distinct and strongly supported clade; and (4) the placement of Allassogonoporus amphoraeformis within the Pleurogenidae, and not close to Lecithodendriidae. Some systematic changes, corresponding to these results, are proposed.     

The systematic position of Leptorhynchoides (Kostylew, 1924) and Pseudoleptorhynchoides (Salgado-Maldonado, 1976), inferred from nuclear and mitochondrial DNA gene sequences

The systematic relationships of acanthocephalans, including Leptorhynchoides and Pseudoleptorhynchoides that occur in freshwater and marine fishes in Neartic and Neotropical regions, are enigmatic. Leptorhynchoides (3 species) and Pseudoleptorhynchoides (1 species) are presently classified in the Rhadinorhynchidae. However, recent molecular and morphological phylogenies have challenged the monophyly of this family. Sequences of nuclear ribosomal DNA (large subunit, small subunit regions) and the mitochondrial cytochrome c oxidase subunit I gene of Leptorhynchoides thecatus and Pseudoleptorhynchoides lamothei were used in phylogenetic analyses with available sequences of 26 other acanthocephalans. Maximum parsimony and maximum likelihood analyses were identical in placing both genera in the Illiosentidae. Bootstrap analyses also indicate that placement of these genera with members of Illiosentidae is reliably supported.     

Clarification of the systematic position of Hippophae goniocarpa (Elaeagnaceae)

New information on (1) Hippophae goniocarpa Y.S. Lian et al . ex Swenson et Bartish and (2) H. goniocarpa ssp. litangensis (Lian et X.L. Chen ex Swenson et Bartish) Lian et K. Sun from China is provided and illustrated. Based on morphological characters and distribution, H. litangensis Y.S. Lian et X.L. Chen ex Swenson et Bartish is re-described as a subspecies. H. goniocarpa (including ssp. litangensis ) is a distinct species and should be placed in Section Gyantsensis of Hippophae , although it has been suggested that sspp. litangensis and goniocarpa are the result of hybridization between H. rhamnoides ssp. sinensis and H. neurocarpa ssp. neurocarpa , and H. rhamnoides ssp. yunnanensis and H. neurocarpa ssp. stellatopilosa , respectively. The type subspecies of H. goniocarpa is distributed in Songpan and Hongyuan Counties of Sichuan Province and Qilian County of Qinghai Province, at 3500–3750 m, whereas ssp. litangensis is found only in Jiawa, Litang County, Sichuan Province at c . 3700 m. A key including the four taxa of sect. Gyantsensis is provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 425−430.     

Differentiation of European freshwater bucephalids (Digenea: Bucephalidae) based on karyotypes and DNA sequences

Three species of bucephalid digeneans are known in European freshwater habitats. In this study parthenitae of Rhipidocotyle campanula (Dujardin, 1845) and R. fennica Gibson, Taskinen & Valtonen, 1992, infecting unionid bivalves, and adult Bucephalus polymorphus von Baer, 1827 from perch (Perca fluviatilis L.) were investigated using karyological analysis and DNA sequencing. Our previously published data on genetic characteristics of parthenitae of B. polymorphus from Dreissena polymorpha Pallas were used for comparative analysis. Ribosomal DNA sequences (ITS2 and 28S rDNA) were used to estimate the phylogenetic relationships of the three bucephalid species. Very close phylogenetic affinity between investigated species was revealed; the sequence difference between the two species of Rhipidocotyle Diesing, 1858 (3.78% based on 28S) was comparable with intergeneric differences observed in comparisons of B. polymorphus with R. campanula and R. fennica (3.43% and 4.49% based on 28S, respectively). A high degree of similarity was noted in karyotype structure of the two species of Rhipidocotyle. The diploid chromosome sets consist of 14 bi-armed chromosomes with the first pair of metacentric elements markedly larger than the remaining chromosomes. This chromosome set structure is also specific to B. polymorphus. One specimen of Anodonta anatina L. was infected with tetraploid R. fennica (4n = 28). On the basis of karyotype characters and molecular data, species of the genus Rhipidocotyle cannot be recognised as more closely related to each other than to B. polymorphus. Our findings of Lithuanian and Ukrainian populations of unionid mussels infected with R. fennica provide evidence that this species occurs not only in Finland but also in Central and Eastern Europe. Previous reports of B. polymorphus in unionids in these regions are equivocal because of possible confusion with R. fennica.     

New data on the phylogeny of Ariantinae (Pulmonata,Helicidae) and the systematic position of Cylindrus obtusus based on nuclear and mitochondrial DNA marker sequences

The phylogenetic relationships among genera of the subfamily Ariantinae (Pulmonata, Helicidae), especially the sister‐group relationship of Cylindrus obtusus, were investigated with three mitochondrial (12S rRNA, 16S rRNA, Cytochrome c oxidase subunit I) and two nuclear marker genes (Histone H4 and H3). Within Ariantinae, C. obtusus stands out because of its aberrant cylindrical shell shape. Here, we present phylogenetic trees based on these five marker sequences and discuss the position of C. obtusus and phylogeographical scenarios in comparison with previously published results. Our results provide strong support for the sister‐group relationship between Cylindrus and Arianta confirming previous studies and imply that the split between the two genera is quite old. The tree reveals a phylogeographical pattern of Ariantinae with a well‐supported clade comprising the Balkan taxa which is the sister group to a clade with individuals from Alpine localities. Additional lineages representing samples from southern Alpine localities as well as from Slovakia split from more basal nodes, but their relationships are not clearly resolved. To achieve more definitive conclusions concerning the geographical origin of Ariantinae, still more sequence data are needed to obtain a tree with better resolution of basal nodes. The genetic data also provided new insights concerning the genus Cepaea, which was used as one of the outgroup taxa. Cepaea vindobonensis is only distantly related to Cepaea nemoralis and Cepaea hortensis, the latter two being more closely related to Eobania vermiculata. Thus, in our tree, the genus Cepaea is paraphyletic.     

Phylogeny of the Bovidae (Artiodactyla, Mammalia), based on mitochondrial ribosomal DNA sequences.

Portions of the 12S and 16S mitochondrial ribosomal genes for 16 species representing nine tribes in the mammal family Bovidae were compared with six previously published orthologous sequences. Phylogenetic analysis of variable nucleotide positions under different constraints and weighting schemes revealed no robust groupings among tribes. Consensus trees support previous hypotheses of monophyly for four clades, including the traditional subfamily Bovinae. However, the basal diversification of bovid tribes, which was largely unresolved by morphological, immunodiffusion, allozyme, and protein sequence data, remains unresolved with the addition of DNA sequence data. The intractability of this systematic problem is consistent with a rapid radiation of the major bovid groups. Several analyses of our data show that monophyly of the Bovidae, which was weakly supported by previous morphological and molecular work, is questionable.     

Molecular phylogenetic position of Japanese Abies (Pinaceae) based on chloroplast DNA sequences

The phylogenetic positions of Japanese Abies species (A. firma, A. homolepis, A. veitchii, A. sachalinensis, and A. mariesii) were revealed based on nucleotide sequences of chloroplast DNA. First, rbcL sequences of 24 Abies species worldwide were analyzed to clarify the phylogenetic position of the Japanese Abies within this genus. While 4 of the Japanese species formed a monophyletic group with almost no sequence divergence, A. mariesii was placed into a clearly different group with North American species. Second, to clarify the relationships among the species in the monophyletic group, sequences of rbcL, matK, and six spacer regions (total ca. 5 kb) were analyzed for 12 species of the group, with A. mariesii as an outgroup. Although a unique sequence and some species-specific sequences were detected in A. mariesii and in A. firma and A. homolepis, respectively, A. veitchii and A. sachalinensis gave identical sequences for all the sequenced regions.     

Systematic position and phylogenetic relationships of the family Omphalometridae (Digenea, Plagiorchiida) inferred from partial lsrDNA sequences

The phylogenetic relationships and systematic position of the digenean genus Omphalometra Looss, 1899 and several other closely related genera, have always been controversial and opinions of different authors on the systematic rank and content of this group have varied greatly. Molecular analysis based on the partial sequences of the large subunit ribosomal DNA gene of representatives of the genera Omphalometra, Rubenstrema and Neoglyphe as well as previously published sequences of members of five families of Plagiorchioidea, has demonstrated: (1) close phylogenetic relationships between these three genera, and (2) a strong support of their position within the family Plagiorchiidae as a well-defined separate clade considered here as a subfamily Omphalometrinae. Molecular data do not support the close affinities of the members of Omphalometrinae and genus Opisthioglyphe as has been suggested by majority of previous authors. Among Omphalometrinae, Omphalometra flexuosa (a parasite of moles, Talpidae) occupies a basal position in relation to Rubenstrema exasperatum and Neoglyphe locellus (both parasitic in shrews, members of the more evolutionary advanced family Soricidae). An extremely low level of lsrDNA sequence divergence between Neoglyphe and Rubenstrema suggests very close phylogenetic relationships of these two genera. Results of the molecular analysis are briefly discussed in comparison with the previously published systems.     

On the systematic position of the genus Petroschmidtia Taranetz et Andriashev, 1934 (Perciformes: Zoarcidae)

Following a comparative analysis of the morphological characters of the representatives of the genera Petroschmidtia and Lycodes, some previously proposed diagnostic characters of Petroschmidtia are confirmed and ascertained and several additional characters are suggested. The conclusion on the taxonomic validity of the genus Petroschmidtia is confirmed. Petroschmidtia differs from Lycodes in the following features: teeth and tooth plates on the vomer and palatinum are lacking; bone fontanels of the supraorbital canal on the frontale are lacking; the vomer is shortened; the ethmoidalia lateralia are narrowed, bearing a pocketlike cavity inside the lateral lobes; the fan-like expanded outgrowth of the supraoccipitale is short, blunt; the dentale is shortened; the radialia are dorsoventrally flattened, the lowest of which is the largest. The genus Petroschmidtia is assumed to be morphologically more closely related to Lycodes than any other of the Lycodinae.     

Filling the gaps in the classification of the Digenea Carus, 1863: systematic position of the Proterodiplostomidae Dubois, 1936 within the superfamily Diplostomoidea Poirier, 1886, inferred from nuclear and mitochondrial DNA sequences

The Diplostomida Olson, Cribb, Tkach, Bray & Littlewood, 2003 is the less diverse order of the two orders within the subclass Digenea Carus, 1863 and is currently classified into three superfamilies, i.e. Brachylaimoidea Joyeux & Foley, 1930, Diplostomoidea Poirier, 1886, and Schistosomatoidea Stiles & Hassall, 1898. Although the suprageneric-level relationships have been elucidated with the use of molecular markers, the lack of representation of some groups obscure the phylogenetic relationships among families, rendering the classification unstable. Here, we tested the phylogenetic position of the family Proterodiplostomidae Dubois, 1936 based on partial 28S rDNA and complete 18S rDNA sequences for Crocodilicola pseudostoma (Willemoes-Suhm, 1870), a crocodile parasite that has been found as a progenetic metacercaria parasitising the pale catfish Rhamdia guatemalensis (Günther) in Mexico and in other siluruforms in the Neotropics. We augmented the representation of the species, genera and families within the Diplostomida, including mostly representatives of the superfamily Diplostomoidea, and assembled a dataset that contains 49 species for the 28S rRNA gene, and 45 species for the 18S rRNA gene. Additionally, we explored the phylogenetic signal of the mitochondrial gene cox1 in reconstructing the phylogenetic relationships of selected members of the superfamily. Our analyses showed that the family Proterodiplostomidae is the sister taxon to the paraphyletic Diplostomidae Poirier, 1886 and Strigeidae Railliet, 1919, with Cyathocotylidae Mühling, 1898 + Brauninidae Wolf, 1903 as their sister group. Analysis of concatenated 18S + 28S sequences revealed the Liolopidae Odhner, 1912 as the basal group of the superfamily Diplostomoidea, although analyses of independent datasets showed that the position of this family remains uncertain. Analysis based on cox1 unequivocally resolved the Proterodiplostomidae as the sister taxon to the Diplostomidae and Strigeidae, although the Cyathocotylidae was nested in a different clade, along with brachylaimoids and schistosomatoids.     

Convoluted history and confusing morphology: Molecular phylogenetic analysis of dicrocoeliids reveals true systematic position of the Anenterotrematidae Yamaguti, 1958 (Platyhelminthes,Digenea)

The Dicrocoeliidae is a highly diverse family of digeneans parasitic in amniotic tetrapods. Detailed molecular phylogenetic analysis of dicrocoeliids is lacking and only a few dicrocoeliids from mammals have been included in previous studies. Sequence data were previously absent for the Anenterotrematidae that shares several morphological characteristics with dicrocoeliids. We examined phylogenetic affinities of several newly sequenced (nuclear 28S rDNA) taxa of dicrocoeliids and anenterotrematids collected from small mammals in Ecuador, Panama, Peru, USA and Vietnam. Our analyses demonstrated that the two anenterotrematid genera (Anenterotrema, Apharyngotrema) belong to the Dicrocoeliidae, placing the Anenterotrematidae into synonymy with the Dicrocoeliidae. Molecular data combined with morphological examination of type and new specimens provided evidence that Parametadelphis and Apharyngotrema are junior synonyms of Metadelphis, with all Metadelphis species lacking a digestive system. Phylogenetic analysis demonstrates that reduction of the alimentary tract in Lutztrema and its loss in Anenterotrema and Metadelphis represent at least two independent evolutionary events. Genera Brachylecithum, Brachydistomum, and Lyperosomum proved to be non-monophyletic, each likely representing more than a single genus. Furthermore, phylogenetic analysis did not support monophyly of the two largest subfamilies of the Dicrocoeliidae (Dicrocoeliinae and Leipertrematinae) with the other two subfamilies not included in this study. Therefore, we propose to abandon the current subfamily division of the Dicrocoeliidae. Analysis of host associations indicates multiple host-switching events throughout evolution of dicrocoeliids. Lastly, analysis of dicrocoeliid geographic distribution revealed that nearly all major clades included taxa from more than a single zoogeographic realm with the exception of the clade Anenterotrema?+?Metadelphis, found only in the Neotropics.     

A redescription of Pintneriella Yamaguti, 1934 (Cestoda: Trypanorhyncha and an examination of its systematic position

Specimens of Pintneriella musculicola Yamaguti, 1934 are redescribed based on paratype material and specimens subsequently allocated to the species by its describer. The cestode species is characterised by two bothridia, a typical heteroacanthous armature with a distinctive basal swelling and basal armature, a space between hook files 1 and 1', nine hooks per principal row, no intercalary hooks and bands of hooklets or chainettes but a distinctive space on the external surface of the tentacle between hook files 9 and 9'. The strobilia is unknown. The features described indicate that Pintneriella is a valid genus within the Heteracanthoidea. Its currently known morphological features do not allow its allocation to an existing family.     

Nested clade analysis of Dunnia sinensis (Rubiaceae), a monotypic genus from China based on organelle DNA sequences

Dunnia sinensis, a monotypic genus of theRubiaceae endemic to the southeast mainlandChina, is an endangered species due to habitatdestruction over the past decades. Informationon levels and apportionment of geneticvariation across populations and geographicalregions is fundamental to conservation. In thepresent study, we used organelle DNA variationand nested phylogeographic analyses to test theisolation-by-distance model in this specieswith wind-mediated seed-dispersal and todistinguish ongoing gene flow from historicalprocesses. As expected, low levels of geneticvariation were detected at the ribosomal ITSregion of mtDNA ( = 0.0019± 0.0002) and the atpB-rbcLintergenic spacer of cpDNA ( =0.0022 ± 0.0009) in the rare species. Sixand seven haplotypes of mt- and cpDNA wereidentified from 125 individuals, respectively,according to the reconstructed neighbor-joiningtrees. Both data sets suggested consistentphylogenies that recovered two differentiatedlineages corresponding to western (Yangchun)and eastern (four others populations) portionsof the range. Hierarchical analyses of themolecular variance (AMOVA) of mt- and cpDNAindicated that molecular variance wasattributable to the difference between regions(ct = 0.911 and 0.771 for mt- and cpDNA,respectively) in D. sinensis. Based ongeographic distributions of haplotypes in thehaplotype networks, significant geneticdifferentiation between the two geographicregions, which can be seen as evolutionarilyconservation units, was associated withhistorical fragmentation. In contrast, limitedgene flow with occasional long-range dispersalshaped the apportionment of organelle DNAalleles among populations of the easternregion, within which two incompletely isolatedphylogeographic groups can be recognized asconservation units for management.     

Phylogeny of Hanabusaya (Campanulaceae), a Korean endemic,based on ITS sequences of nuclear ribosomal DNA

The phylogenetic position of the genus Hanabusaya was verified using the internal transcribed spacer (ITS) sequences data of 181 individuals of 176 taxa in Campanulaceae, including allied genera such as genus Campanula, Adenophora, Symphyandra, and Zeugandra. The phylogenetic analysis showed thatHanabusaya is more closely allied to Adenophora than to Symphyandra, Zeugandra, and Campanula. Accordingly, the connate anthers of Hanabusaya are considered to have evolved independently of Symphyandra or Zeugandra. However, Hanabusaya and sect. Remotiflorae of Adenophora were unresolved within a clade. Therefore, the relationship between Hanabusaya and its allies should be more accurately determined through a broader study on diverse genetic regions.     

Molecular phylogeny of the large carpenter bees, genus Xylocopa (Hymenoptera: apidae), based on mitochondrial DNA sequences

Carpenter bees, genus Xylocopa Latreille, a group of bees found on all continents, are of particular interest to behavioral ecologists because of their utility for studies of the evolution of mating strategies and sociality. This paper presents phylogenetic analyses based on sequences of two mitochondrial genes cytochrome oxidase 1 and cytochrome b for 22 subgenera of Xylocopa. Maximum-parsimony and maximum-likelihood methods were used to infer phylogenetic relationships. The analyses resulted in three resolved clades of subgenera: a South American group (including the subgenera Stenoxylocopa, Megaxylocopa, and Neoxylocopa), a group including the subgenera Xylocopa s.s. and Ctenoxylocopa, and an Ethiopean group (including the subgenera Afroxylocopa, Mesotrichia, Alloxylocopa, Platynopoda, Hoploxylocopa, and Koptortosoma). The relationships between the 11 other subgenera and the resolved clades are unclear. Within the Ethiopian group we found a clear separation of the African and the Oriental taxa and apparent polyphyly of the subgenus Koptortosoma. Using an evolutionary rate for ants, we investigated whether Gondwana vicariance or more recent dispersal events could best explain the present-day distribution of subgenera. Although some taxa show divergences that approach Gondwanan breakup times, most divergences between geographic groups are too recent to support a vicariance hypothesis.     

Molecular systematic analysis of the order Proteocephalidea (Eucestoda) based on mitochondrial and nuclear rDNA sequences

Two ribosomal DNA sequences were used to infer phylogenetic relationships among the Eucestoda order Proteocephalidea. A 437 bp portion of the 16S mitochondrial and a 1149 bp 5' portion of the nuclear large sub-unit rRNA molecule were sequenced for 53 proteocephalidean cestodes (representing nine subfamilies and 22 genera) and for one outgroup species. Parsimony and distance-based analyses of the two databases, alone and combined, failed to support the monophyly of the two traditionally accepted families, of numerous subfamilies (with the exception of the Rudolphielliinae and Othinoscolescinae which were validated in our analysis) and of various genera, including the genus Nomimoscolex (Woodland), Ophiotaenia (La Rue) as well as the type genus Proteocephalus (Weinland). Palaearctic Proteocephalus species nevertheless constituted a well-defined clade. The two genes globally yielded compatible results; however, the nuclear ribosomal gene provided a better resolution of relations among Proteocephalidea.     

A phylogenetic analysis of Pseudonaja (Hydrophiinae, Elapidae, Serpentes) based on mitochondrial DNA sequences

A phylogenetic analysis of mitochondrial ND4 and adjacent tRNA sequences for a geographically extensive series of specimens reveals nine major clades within Pseudonaja, of which six are largely coincident with nominal taxa (P. affinis, P. guttata, P. inframacula, P. ingrami, P. modesta, and P. textilis). The remaining three clades are composed of specimens presently referred to P. nuchalis. Two of these clades correspond with the "Darwin" and "Southern" morphs of previous authors, while the third clade incorporates the "Orange with black head" and "Pale head, grey nape" morphs. We are unable to confirm the presence of consistent karyotypic differences between "Orange with black head" and "Pale head, grey nape" specimens, however, P. inframacula, P. textilis, and P. nuchalis "Darwin" are found to exhibit distinctive karyotypes, as previously reported. These results, in conjunction with additional observations of karyotpic and morphological variation, are consistent with nine historically-independent lineages (i.e., species) within Pseudonaja. There is strong support for a clade composed of P. affinis, P. inframacula, P. ingrami, P. textilis, and the three P. nuchalis lineages, and for the relationships (P. inframacula, P. nuchalis "Southern") and (P. nuchalis "Darwin", P. nuchalis "Orange with black head"--"Pale head, grey nape" ).