Evolutionary relationships among eight species of South American phyllotine rodents (Rodentia: Muridae) based on allozymic data

Rodents of the tribe Phyllotini represents one of the main radiations of South American Sigmodontini. Phylogenetic relationships among species of this highly diversified group are poorly known. In this paper we analyse evolutionary relationships among eight phyllotine species belonging to the genera Calomys , Graomys , Phyllotis and Eligmodontia , on the basis of allozymic polymorphisms. Most of the differences among species were in allele frequencies and not of allele class. Neighbour-joining and maximum likelihood methods place P. xanthopygus in the same group as E. typus and G. griseoflavus , in agreement with results obtained by several authors on the basis of morphological characters. Parsimony analysis of 0–9 coded data suggest that the genus Calomys is paraphyletic, but with a low bootstrap support. In the tree based on genetic distance data, the genus also appears as paraphyletic. The maximum likelihood method yields a tree where Calomys is monophyletic, but this phylogeny is supported by only two out of 78 alleles analysed. Calomys hummelincki and C. venustus occupy a basal position among Calomys species. Calomys musculinus and C. lepidus are the most closely related species of the genus, with C. laucha as sister to them. These relationships are strongly supported by bootstrap percentages.     

Suprageneric relationships of galliform birds (Aves, Galliformes): a cladistic analysis of morphological characters

Of the basal clades of extant birds (Neornithes) the 'landfowl' or galliforms (Aves, Galliformes) are the most speciose. Cladistic analysis of more than 100 morphological characters coded at the generic level for most putative galliform genera confirms that the megapodes ('mound builders'; Megapodiidae) are the most basal clade within the order. They are followed successively by the curassows, guans and chachalacas (Cracidae), which comprise the sister-group to all other extant Galliformes (i.e. Phasianoidea). Within this large 'phasianoid' clade, analyses suggest that the guineafowl (Numididae) are the most basal taxon, although monophyly of this 'family' is not strictly supported on the basis of the morphological characters employed. An additional major clade within the phasianoid Galliformes is recovered by this analysis, comprising the traditional groupings of New World quails (Odontophoridae) and Old World quails ('Perdicini'), yet only monophyly of the former is supported unambiguously by morphological characters. Relationships within the remainder of the phasianoid taxa, including the grouse (Tetraonidae), turkeys (i.e. Meleagris / Agriocharus spp.) as well as other 'pavonine' galliforms (i.e. peafowl; Pavo , Afropavo , Rheinardia , Argusianus and Polyplectron spp.) remain largely unresolved on the basis of morphological characters, yet monophyly of the major subdivisions is supported here. Although there are a number of important differences, especially with regard to relationships within the nonquail phasianoids, the results of this morphological phylogenetic (cladistic) analysis are broadly congruent both with traditional classifications and existing molecular hypotheses of galliform phylogenetic relationships.     

Relationships among extant and fossil echimyids (Rodentia: Hystricognathi)

The echimyid rodents are the most diverse group of Neotropical hystricognaths, with approximately 40 extant and fossil genera. Craniodental characters are proposed in order to formulate hypotheses of phylogenetic relationships within the Echimyidae. A data matrix of 54 taxa and 50 characters is constructed and submitted to parsimony analyses using PAUP and WinClada programs. Analysis of the complete data set results in 47 448 most parsimonious trees 107 steps long. These trees are summarized in a strict consensus tree, which is taken as the main phylogenetic hypothesis resulting from this study. The monophyly of several currently recognized supraspecific taxa is not corroborated. These are: the subfamilies Eumysopinae, Echimyinae, Myocastorinae and Adelphomyinae; and the genera Proechimys , Echimys and Makalata . Conversely, the monophyly of Dactylomyinae and Trinomys is supported. New associations are proposed: (1) a clade comprising the extant Carterodon , Clyomys and Euryzygomatomys and the fossil Pampamys and Theridomysops placed at the base of the crown-group Echimyidae; (2) a clade uniting Proechimys , Hoplomys and Trinomys , which is the sister-taxon of (3) a clade including Mesomys , Lonchothrix , Myocastor and a clade with extant dactylomyines and echimyines and associated fossil taxa. Based on this phylogenetic hypothesis, patterns of tooth evolution in Echimyidae are discussed, and minimum ages for the divergence events within the family are estimated.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 445–477.     

Phylogenetic relationships within the South American fish family Anostomidae (Teleostei,Ostariophysi, Characiformes)

Analysis of a morphological dataset containing 152 parsimony‐informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov. , described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synaptolaemus. Rhytiodus and Schizodon together formed a well‐supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis‐ and trans‐Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre‐dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 70–210.     

Phylogenetic relationships of the monozoic tapeworms (Eucestoda: Caryophyllidea) inferred from morphological characters

Phylogenetic relationships of all genera of the order Caryophyllidea, possibly the earliest branching group of true tapeworms (Platyhelminthes: Eucestoda) and the only one that is monozoic, have been assessed for the first time. Results of this cladistic analysis, inferred from 30 unweighted morphological characters, are only partly congruent with the existing classification, which consists of four families based on the position of the inner longitudinal muscles in relation to the internal genital organs. Whereas all but five genera of the Caryophyllaeidae form a monophyletic clade, members of the Capingentidae are split, occurring within six unrelated groups. The Lytocestidae is also paraphyletic, as some genera appear in four unrelated clades. Archigetes appears in a derived clade, indicating that its direct (monoxenous) life-cycle involving only tubificid oligochaetes is secondarily derived and not plesiomorphic among the Eucestoda, as postulated by some authors.     

Phylogenetic relationships of thorny catfishes (Siluriformes: Doradidae) inferred from molecular and morphological data

Doradidae is a putatively monophyletic group of South American freshwater catfishes containing 30 extant genera and 72 valid species. Only one study to date has attempted to estimate phylogenetic relationships among doradids. This morphological analysis partitioned species into two basal genera ( Wertheimeria and Francisodoras ) and a crown group of three subfamilies (Platydoradinae, Astrodoradinae and Doradinae) whose relationships were unresolved. No subsequent work has been done to resolve the subfamilial trichotomy or to assess whether postulated intergeneric relationships are accurate. We address this problem with complete sequences (2.5 kilobases, kb) of mitochondrial 12S and 16S rRNA genes and partial (1.3 kb) sequences of the nuclear elongation factor-1 alpha (EF1α) gene from representatives of 23 doradid genera (43 species) and 13 outgroups from additional siluriform families. Phylogenetic analysis of these data yields strong support for the monophyly of Doradidae and Astrodoradinae (as well as other relationships), but otherwise shows significant conflict with morphological results. A partial re-examination of published morphological data indicates that many characters may have been incorrectly polarized and many taxa have incorrect state assignments. Our results provide a framework for ongoing efforts to describe the species-level diversity of this poorly understood neotropical family.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 551–575.     

Phylogenetic relationships among atheriniform fishes (Teleostei: Atherinomorpha)

We present evidence from adult and larval morphology for the monophyly and relationships of Atheriniformes, using other atherinomorphs, mugilids and acanthomorph fishes as outgroups. Atheriniformes is diagnosed by ten characters (larval: short preanal length, single mid-dorsal row of melanophores; adult: vomerine ventral face concave, long Al muscle tendon to lacrimal, two anterior infraorbital bones, pelvic-rib ligament, pelvic medial plate not extended to anterior end, and second dorsal-fin spine flexible). We recognize six families within the order, the hierarchical relationships among which are: (Atherinopsidae (Notocheiridae (Melanotaeniidae (Atherionidae (Phallostethidae, Atherinidae))))). Other major conclusions include: (1) Atherinopsidae (Menidiinae, Atherinopsinae) is diagnosed by 20 characters (e.g. ethmomaxillary ligament attached to palatine dorsal process, ventral postcleithrum with two dorsal rami); (2) Melanotaeniidae (Bedotiinae (Melanotaeniinae (Telmatherinini, Pseudomugilini))) is diagnosed by six characters (e.g. absence of second dorsal-fin spine, sexual dimorphism in body colour and median-fin development, greater body depth); (3) Dentatherina is in Phallostethidae; (4) Atherinidae (Atherinomorinae (Craterocephalinae, Atherininae)) is diagnosed by three characters (lacrimal notch, ventral postcleithrum between first and second pleural ribs, pelvic ventral spine); (5) Atherinidae and Phallostethidae form the Atherinoidea clade diagnosed by seven characters (e.g. interopercle dorsal process absent, dorsal wings of urohyal absent, ventral postcleithrum laminar, pelvic medial plate extended to anterior end, presence of anal plate). Bedotia, Rhodes , and melanotaeniines are shown to be derived within atheriniforms rather than the plesiomorphic sister groups to a paraphyletic 'atherinoid' group. We also demonstrate that groups traditionally placed in Atherinidae (Menidiinae, Atherininae, Atherioninae, etc.) comprise a paraphyletic assemblage.     

Phylogenetic relationships of Microdontinae (Diptera: Syrphidae) based on molecular and morphological characters

The intrasubfamilial classification of Microdontinae Rondani (Diptera: Syrphidae) has been a challenge: until recently more than 300 out of more than 400 valid species names were classified in Microdon Meigen. We present phylogenetic analyses of molecular and morphological characters (both separate and combined) of Microdontinae. The morphological dataset contains 174 characters, scored for 189 taxa (9 outgroup), representing all 43 presently recognized genera and several subgenera and species groups. The molecular dataset, representing 90 ingroup species of 28 genera, comprises sequences of five partitions in total from the mitochondrial gene COI and the nuclear ribosomal genes 18S and 28S. We test the sister‐group relationship of Spheginobaccha with the other Microdontinae, attempt to elucidate phylogenetic relationships within the Microdontinae and discuss uncertainties in the classification of Microdontinae. Trees based on molecular characters alone are poorly resolved, but combined data are better resolved. Support for many deeper nodes is low, and placement of such nodes differs between parsimony and Bayesian analyses. However, Spheginobaccha is recovered as highly supported sister group in both. Both analyses agree on the early branching of Mixogaster, Schizoceratomyia, Afromicrodon and Paramicrodon. The taxonomical rank in relation to the other Syrphidae is discussed briefly. An additional analysis based on morphological characters only, including all 189 taxa, used implied weighting. A range of weighting strengths (k‐values) is applied, chosen such that values of character fit of the resulting trees are divided into regular intervals. Results of this analysis are used for discussing the phylogenetic relationships of genera unrepresented in the molecular dataset.     

Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence

This study is undertaken in order to evaluate specific hypotheses of relationship among extant and extinct sloths (Mammalia, Xenarthra, Tardigrada). Questions of particular interest include the relationship among the three traditional family groupings of extinct ground sloths and the monophyletic or diphyletic origin of the two genera of extant tree sloths. A computer‐based cladistic investigation of the phylogenetic relationships among 33 sloth genera is performed based upon 286 osteological characteristics of the skull, lower jaw, dentition and hyoid arch. Characters are polarized via comparisons with the following successive outgroups, all members of the supraordinal grouping Edentata: the Vermilingua, or anteaters; the Cingulata, or armadillos and glyptodonts; the Palaeanodonta; and the Pholidota, or pangolins. The results of the analysis strongly corroborate the diphyly of living tree sloths, with the three‐toed sloth Bradypus positioned as the sister‐taxon to all other sloths, and the two‐toed sloth Choloepus allied with extinct members of the family Megalonychidae. These results imply that the split between the two extant sloth genera is ancient, dating back perhaps as much as 40 Myr, and that the similarities between the two taxa, including their suspensory locomotor habits, present one of the most dramatic examples of convergent evolution known among mammals. The monophyly of the three traditional ground sloth families Megatheriidae, Megalonychidae and Mylodontidae is confirmed in the present study, and the late Miocene–Pleistocene nothrotheres are shown to form a clade. It is suggested that this latter clade merits recognition as a distinct family‐level grouping, the family Nothrotheriidae. The monophyly of the Megatherioidea, a clade including members of the families Megatheriidae, Megalonychidae and Nothrotheriidae, is also supported. Within Megatherioidea, the families Nothrotheriidae and Megatheriidae form a monophyletic group called the Megatheria. The relationships within the families Megatheriidae and Mylodontidae are fully and consistently resolved, although the hypothesized scheme of relationships among the late Miocene to Pleistocene members of the mylodontid subfamily Mylodontinae differ strongly from any proposed by previous authors. Within the family Megalonychidae, Choloepus is allied to a monophyletic grouping of West Indian sloths, although the relationships within this clade are not fully resolved. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 255–305.     

A new genus and species of long-clawed mouse (Rodentia: Muridae) from temperate rainforests of Chile

Morphological comparisons of long-clawed mice in southern South America reveal the existence of four genera, only three of which are named. The fourth genus is described and compared with Geoxus, Chelemys and Notiomys. External, cranial and dental characters indicate that the new mouse is most closely related to Geoxus , the raton topo or mole mouse. Geoxus and the new form may be distinguished using both qualitative and quantitative characters. The incipient specializations of the new mouse for fossorial life suggest that it represents an early offshoot along the line leading to the more highly specialized Geoxus. Discovery of a new mammalian genus in Chile, where naturalists have been active for more than two centuries, indicates the importance of continued biological surveys, even in ostensibly well-studied regions. The new mouse reinforces the strong differentiation of faunas inhabiting southern beech ( Nothofagus ) forests in South America, where generic endemism of rodents and marsupials reaches 67%.     

A morphological study of the Petunia integrifolia complex (Solanaceae)

BACKGROUND AND AIMS: Petunia inflata has been treated taxonomically in various ways: it has been described as an independent species, treated as a synonym of P. integrifolia, and also regarded as a subspecies of P. integrifolia. The present study was designed to resolve the ambiguity involving the P. integrifolia complex (P. integrifolia plus P. inflata). METHODS: Tentative identification (either integrifolia group or inflata group) was carried out in the field based on the observation of live specimens at the restricted type localities. The accuracy of the tentative identification was later tested with principal component and cluster analyses of data obtained by measuring 21 morphological characters on cultivated live specimens sourced from 113 natural populations of the P. integrifolia complex in Argentina, Brazil, Paraguay and Uruguay. KEY RESULTS: There was a clear, statistically significant gap between the morphological measurements of the two groups, ensuring the accuracy of identification carried out in the field except for a probable hybrid swarm. Previously, the condition of the pedicel in the fruiting state was considered an important character distinguishing between these two groups; however, the condition of the pedicel was rather variable in the integrifolia group. The two groups were found to have geographically distinct distributions: the integrifolia group occurred in southern regions, whereas the inflata group occurred in northern regions. CONCLUSIONS: Based on the available evidence, it is suggested that the two groups are allopatric species, P. integrifolia and P. inflata, in agreement with the opinion of Fries (1911).     

Phylogenetic relationships between isolated populations of the limestone‐dwelling microsnail Gyliotrachela hungerfordiana (Gastropoda: Vertiginidae)

The vertiginid species Gyliotrachela hungerfordiana, an obligate limestone‐dwelling microsnail, is relatively widespread and is found on a large number of isolated limestone hills in Peninsular Malaysia. To elucidate the pattern of colonization of these hills, we conducted a molecular phylogenetic analysis on G. hungerfordiana subpopulations from 15 separate limestone outcrops. As outgroups, we also included five related Peninsular Malaysian Vertiginidae (Gyliotrachela frequens, Gyliotrachela transitans, Gyliotrachela salpinx, Gyliotrachela depressispira and Paraboysidia tarutao), one population each. A combined analysis of nuclear (internal transcribed spacer 1) and mitochondrial (cytochrome c oxidase 1) sequences showed that (1) G. hungerfordiana is monophyletic; (2) there is a clear geographical pattern in the phylogenetic relationships between G. hungerfordiana populations, with genetic distances increasing with geographic distance; (3) this pattern is disturbed by a few long‐distance (north‐west to south‐east and north to south) colonizations.     

Phylogenetic relationships of basal hexapods among the mandibulate arthropods: a cladistic analysis based on comparative morphological characters

In this paper we propose a reappraisal of the relationships between the basal hexapod lineages (the former 'apterygote' insects) and the other major groups of mandibulate arthropods. It results from a cladistic analysis including 72 characters based on external morphology, internal anatomy and development. Detailed comments are provided on the various characters used and the scoring of their states. The 35 terminal taxa include 12 hexapods (9 of which are basal 'apterygote' representatives), 7 myriapods, 13 crustaceans, and 3 chelicerates taken as outgroups. The results of our analyses are discussed in detail for each of the taxonomic groupings, and compared with those recently obtained by other authors using different approaches based on morphological, palaeontological, developmental or molecular sequence data. Our results support the monophyly of the Mandibulata, Crustacea, Atelocerata (Tracheata) and Hexapoda, but the assemblage of Myriapoda appears poorly supported. A close relationship between Crustacea and Hexapoda, as hypothesized by several authors, is not found in any of our analyses. Within Hexapoda, the Protura and the Collembola appear as independent clades, whereas the two unresolved dipluran taxa are grouped with the monophyletic Ectognatha (Archaeognatha, Zygentoma and Pterygota).     

Phylogenetic relationships within Maldanidae (Capitellida,Annelida), based on morphological characters

The Maldanidae annelid worms are reviewed and the phylogenetic relationships of their subgroups provided, based on Hennigian principles and maximum parsimony. Characters were coded as binary and multistate (transformation series). We used 33 terminal taxa (species), and 50 characters. Characters were treated as unordered and of equal weight, and analysis was run in TNT. Three equally most-parsimonious trees were obtained with heuristic searches, with lengths of 64 steps; CI = 0.95, and RI = 0.98. The monophyly of Maldanidae was supported with 100% of bootstrap and jackknife values. As a result of our analysis, Arenicolidae remains the sister-group of Maldanidae, and both should be referred to Maldanomorpha. Maldanidae was supported by the following synapomorphies: dorsal prostomium; prostomium keel-shaped and fused to peristomium; torus globose behind median chaetigers; median chaetigers greatly elongated; number of pre-anal segments reduced. The subfamily Bogueinae was not monophyletic; Boguea and Boguella were included within Rhodininae. Clymenura, previously included in the Clymenurinae, was included within Euclymeninae. The taxa Notoproctinae, Maldaninae, Nicomachinae and Euclymeninae were grouped in the Maldanoplaca, a new taxon. Eight further new clades have been found, but were not named.     

Phylogenetic convergence and multiple shell shape optima for gliding scallops (Bivalvia: Pectinidae)

An important question in evolutionary biology is how often, and to what extent, do similar ecologies elicit distantly related taxa to evolve towards the same phenotype? In some scenarios, the repeated evolution of particular phenotypes may be expected, for instance when species are exposed to common selective forces that result from strong functional demands. In bivalved scallops (Pectinidae), some species exhibit a distinct swimming behaviour (gliding), which requires specific biomechanical attributes to generate lift and reduce drag during locomotive events. Further, a phylogenetic analysis revealed that gliding behaviour has independently evolved at least four times, which raises the question as to whether these independent lineages have also converged on a similar phenotype. Here, we test the hypothesis that gliding scallops display shell shape convergence using a combination of geometric morphometrics and phylogenetic comparative methods that evaluate patterns of multivariate trait evolution. Our findings reveal that the gliding species display less morphological disparity and significant evolutionary convergence in morphospace, relative to expectations under a neutral model of Brownian motion for evolutionary phenotypic change. Intriguingly, the phylomorphospace patterns indicate that gliding lineages follow similar evolutionary trajectories to not one, but two regions of morphological space, and subsequent analyses identified significant differences in their biomechanical parameters, suggesting that these two groups of scallops accomplish gliding in different ways. Thus, whereas there is a clear gliding morphotype that has evolved convergently across the phylogeny, functionally distinct morphological subforms are apparent, suggesting that there may be two optima for the gliding phenotype in the Pectinidae.     

Phylogenetic relationships among superfamilies of Cicadomorpha (Hemiptera: Auchenorrhyncha) inferred from the wing base structure

The infraorder Cicadomorpha is a monophyletic group of the order Hemiptera, suborder Auchenorrhyncha, and is composed of three superfamilies: Cercopoidea (spittle bugs), Cicadoidea (cicadas) and Membracoidea (leafhoppers and treehoppers). Phylogenetic relationships among the superfamilies have been highly controversial morphologically and molecularly, but recent molecular phylogenetic analyses provided support for Cercopoidea + Cicadoidea. In this study, we examined morphology of the wing base structure in Cicadomorpha and tested the previous phylogenetic hypotheses using the characters selected from the wing base. As a result, a sister‐group relationship between Cicadoidea and Cercopoidea was supported by three synapomorphies (presence of a projection posterior to the anterior notal wing process, presence of a novel notal process anterior to the posterior notal wing process, presence of a novel sclerite between the distal median plate and the base of anal vein). The present study provides the first unambiguous and prominent morphological support for Cicadoidea + Cercopoidea.     

Phylogenetic relationships of the family Axinellidae (Porifera: Demospongiae) using morphological and molecular data

Alvarez, B., Crisp, M.D., Driver, F., Hooper, J.N.A. & Van Soest, R.W.M. (2000). Phylogenetic relationships of the family Axinellidae (Porifera: Demospongiae) using morphological and molecular data. —Zoologica Scripta, 29, 169–198. Twenty‐seven species of marine sponges belonging to Axinellidae and related groups (Halichondriidae, Dictyonellidae, Agelasida) were selected to test the monophyly of Axinellidae and investigate their phylogenetic relationships using parsimony and maximum likelihood methods. Partial 28S rDNA sequences, including the D3 domain, and traditional morphological characters (mainly skeletal ones) were used independently to construct phylogenetic trees. Sequences were aligned using the appropriate model of secondary structure of the RNA and compared to that produced by the multiple sequence alignment program, ClustalW. The alignment using secondary structure constraints produced a better estimate of the phylogeny and was demonstrated to be an effective and objective method. Results of the cladistic analyses of the molecular and morphological data sets were not fully congruent; the morphological data suggest that Axinellidae is monophyletic, however, the molecular data suggest that it is nonmonophyletic. The single most‐parsimonious tree derived from the molecular data showed that species of Axinella (except A. polypoides) are united in a clade that is more closely related to members of Agelasida than to other species of Axinellidae; the remaining members of Axinellidae form a monophyletic group that is closely related to the families Dictyonellidae and Halichondriidae. The consensus tree of 20 most‐parsimonious trees from the morphological analysis, on the other hand, showed that all the sampled species of Axinellidae belong to a monophyletic group which is closely related to the species of Dictyonellidae and Halichondriidae. Only two branches were identical in both cladograms, the one uniting the species of Ptilocaulis and Reniochalina and the one with the species of Dictyonellidae. The robustness of the molecular and morphological trees (or parts of the trees), was tested using bootstrap, jack‐knife, PTP and T‐PTP tests. The results of the PTP test were significant indicating significant cladistic structure in both data sets. The bootstrap and jack‐knife values indicate that the molecular tree is in general better supported than the morphological one. The lack of morphological characters and the homoplastic nature of some may explain the weak support of the morphological tree. A T‐PTP test of nonmonophyly showed that the nonmonophyly of Axinellidae, as indicated by the results of the molecular analysis, is not significant; however, a T‐PTP test of monophyly of Axinellidae, as indicated by the morphological tree, produced significant results. This indicates that the monophyly of Axinellidae based on morphological data cannot be rejected; the family however, cannot be defined in terms of a unique diagnostic character common to all members of the ingroup. Tests of heterogeneity (reciprocal T‐PTP and partition homogeneity test) indicated that the data partitions are heterogeneous, which could be due to sampling errors (in either data set) or differences in the underlying phylogenies; therefore data were not combined in a single analysis. Further, both data sets are unequally sized (95 informative molecular characters vs. 16 informative morphological characters), which means that the molecular signal could swamp the morphological signal if the data is combined. Nonmonophyly of Axinellidae is supported by chemical and genetic evidence available in the literature and DNA sequences data of axinellid species from New Zealand. However, this needs to be confirmed using independent evidence from different genes (or gene regions), biochemistry, histology or cell ultrastructure. Therefore, no changes to the taxonomic position of the family in the higher classification are proposed at this stage.     

Phylogenetic relationships within the genus Pimephales as inferred from ND4 and ND4L nucleotide sequences

In phylogenetic analyses, conducted on the ND4L gene and part of the ND4 gene from species of the genus Pimephales , maximum parsimony yielded four trees, with the strict consensus providing no resolution of relationships among species. Maximum likelihood and minimum evolution methods yielded identical tree topologies, which differed from previous hypotheses of relationships for these species. If this topology is correct, it implies independent evolution of morphological characters, possibly associated with convergent trophic specialization.