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Australia has a diversity of vectors and vector-borne human diseases. Mosquito-borne arboviruses are of greatest concern, but there are issues with other vector and pathogen systems. Mosquitoes were responsible for more than 35,000 cases of Ross River virus during 1991-1997. Barmah Forest virus is increasing nationwide, and unidentified bunyaviruses suspected of causing illness have been isolated. Cases of Murray Valley encephalitis have occurred in 14 of the past 20 years in northern Australia. Dengue is a continuing problem for northern Queensland, with various serotypes being active. Japanese encephalitis has appeared in the Torres Strait Islands and threatens mainland Australia. Although malaria is eradicated, almost 1,000 cases are imported annually and occasional cases of local transmission occur. With ticks, paralysis in children occurs annually in eastern Australia. Tick typhus (Queensland Tick Typhus--Rickettsia australis) occurs down the east coast, and (Flinders Island Spotted Fever--Rickettsia honei) in Bass Strait and probably Tasmania. Lyme disease is reported but its presence is controversial. Fleas were responsible for a recent outbreak of murine typhus (Rickettsia typhi) in Western Australia. Mites cause scrub typhus (Orientia tsutsugamushi), and there was a recent fatality in the Northern Territory. Overall, resources for investigation and control of vector-borne disease have generally been meager. However, various avenues of basic and applied research have been pursued, and have included investigations into mosquito ecology, vector competence, disease epidemiology, and vector control. Disease surveillance programs vary between states, and mosquito control programs are organized and effective in only a few regions. There are concerns for import of vectors such as Aedes albopictus and export of pathogens such as Ross River virus; the former has occurred but the species has not become established, and the latter has occurred and has resulted in a major outbreak in the South Pacific. The predicted scenarios of increased temperature and rainfall with global warming are also causing concern for increases in vector-borne diseases, particularly the endemic arboviruses. Interest by health authorities is gravitating more towards epidemiological reporting and less towards public health action. In many respects, humans have much to do to get "on top" of vectors and their pathogens "down under" in Australia.
Recently, the first part of the morphological revision of the Southeast Asian water monitor lizards of the Varanus salvator (Laurenti, 1768) species group provided a taxonomic overview over the members of this successful and widespread species complex (Koch et al. 2007). There, the Philippine taxa marmoratus, nuchalis and cumingi were reelevated to species status due to diagnostic morphological characteristics, e.g. significantly enlarged scales on the neck region. In this second part of the ongoing revision, these three species are re-investigated using additional voucher specimens and advanced statistical techniques including canonical variates analysis and principal component analysis. Our new investigations indicate that V. marmoratus represents a composite species, comprising at least three distinct taxa. Hence, the populations of the Sulu Archipelago (Tawi-Tawi Island) and those of the Palawan region are described as new species, viz. Varanus rasmusseni sp. nov. and V. palawanensis sp. nov., respectively. The allopatric island populations of V. cumingi inhabiting Samar, Leyte, and Bohol (the East Visayan subregion) show characteristic and geographically correlated colour patterns distinct from the type locality Mindanao (the second subregion of Greater Mindanao), warranting subspecific partition of this species. The new subspecies is named V. cumingi samarensis ssp. nov. In contrast, the taxonomic status of V. nuchalis remained unchanged, although this species shows some considerable variation in colour pattern. The systematic chapters are supplemented with notes about biology and conservation status. The hitherto underestimated diversity and zoogeography of Philippine water monitors is discussed in the light of Pleistocene sea level fluctuations. Finally, we introduce a scenario for the evolution and spread of Southeast Asian water monitor lizards and provide an identification key for the Philippine members of the V. salvatoI' complex.
Ubiquitylation is a three-step process, which results in the attachment of the small protein ubiquitin (Ub) to lysine residues on a substrate protein. SUMO proteins are ubiquitin (Ub)-related modifiers implicated in the regulation of gene transcription, cell cycle, DNA repair and protein localization. The molecular mechanisms by which the sumoylation of target proteins regulates diverse cellular functions remain poorly understood. During my PhD I isolated and characterized SUMO1 and SUMO2 binding motifs. Using Yeast Two Hybrid system, bioinformatics and NMR spectroscopy we defined a common SUMO-interacting motif (SIM) and map its binding surfaces on SUMO1 and SUMO2. This motif forms a β-strand that could bind in parallel or anti-parallel orientation to the β2-strand of SUMO due to the environment of the hydrophobic core. A negative charge imposed by a stretch of neighboring acidic amino acids and/or phosphorylated serine residues determines its specificity in binding to distinct SUMO paralogues and can modulate the spatial orientation of SUMO-SIM interactions. Mutation of the SUMO interacting motif of TTRAP (TRAFS and TNF receptor associated protein) influences both its localization and dynamic behaviour in living cells. Ubiquitin (Ub)-binding domains (UBDs) are key elements in conveying Ub-based cellular signals. UBD-containing proteins interact with ubiquitylated targets and control numerous biological processes including receptor trafficking, DNA repair, virus budding and gene transcription. They themselves undergo UBD-dependent monoubiquitylation, which promotes intramolecular binding of the UBD to the attached Ub and consequently leads to their functional inhibition. During the second part of my PhD I could show that, in contrast to the established ubiquitylation pathway, the presence of UBDs allows the monoubiquitylation of host protein independently of classical E3 ligases. UBDs of different types including UBA, UIM, UBM, NFZ and UBZ, can directly cooperate with E2 Ub-conjugating enzymes to promote monoubiquitylation of their host proteins. Using FRET technology I verified that the E2 enzyme and the substrate directly interact in cells. Moreover, UBD-containing proteins Stam2 and Sts2 promote self-ubiquitylation and not ubiquitylation of other targets or form polyUb chains from free Ub. Our study revealed a yet unappreciated role of E2 enzymes in ubiquitylation reactions of UBD containing proteins.
The siliceous claystone and chert lithologic units of the Triassic-Jurassic chert-clastic sequence are well exposed in the Inuyama, Mt. Kinkazan and Hisuikyo areas of the southeastern Mino Terrane. Twenty-one continuous sections from those areas were investigated in order to establish comprehensive radiolarian biozones and clarify the successive lithologic changes through the Triassic and lowest Jurassic. Twenty new radiolarian zones are established; the lowest two are assemblage zones and the others are defined by the first or last occurrence of index taxa. The definitions are as follows in chronological order: TR 0, Follicucullus Assemblage Zone (early Spathian or older); TR 1, Parentactinia nakatsugawaensis Assemblage Zone (late Spathian); TR 2A, Eptingium nakasekoi Lowest-occurrence Zone (early Anisian); TR 2B, Triassocampe coronata group Lowest-occurrence Zone (early Anisian); TR 2C, Triassocampe deweveri Lowest-occurrence Zone (late Anisian); TR 3A, Spine A2 (possiblly derived from Oertlispongus inaequispinosus) Lowest occurrence Zone (late Anisian) ; TR 3B, Yeharaia elegans group Lowest-occurrence Zone (early Ladinian); TR 4A, Muelleritortis cochleata Lowest-occurrence Zone (late Ladinian); TR 4B, Spongoserrula dehli Lowest-occurrence Zone (late Ladinian to early Carnian); TR 5A, Capnuchosphaera Lowest-occurrence Zone (early Carnian); TR 5B, Poulpus carcharus sp. nov. Lowest-occurrence Zone (early to late Carnian); TR 6A, Capnodoce- Trialatus Concurrentrange Zone (late Carnian to early Norian), TR 6B, Trialatus robustus-Lysemelas olbia gen. et sp. nov. Partial-range Zone (early Norian); TR 7, Lysemelas olbia gen. et sp. nov. Lowest-occurrence Zone (early to late Norian); TR 8A: Praemesosaturnalis multidentatus group Lowest-occurrence Zone (late Norian); TR 8B: Praemesosaturnalis pseudokahleri sp. nov. Lowest-occurrence Zone (late Norian) ; TR 8C: Skirt F (possiblly derived from Haeckelicyrtium takemurai) Lowest-occurrence Zone (late Norian to early Rhaetian); TR 8D: Haeckelicyrtium breviora sp. nov. Taxon-range Zone (early to late Rhaetian) ; JR OA: Haeckelicyrtium breviora sp. nov.-Bipedis horiae sp. nov. Partial-range Zone (Hettangian); and JR OB: Bipedis horiae sp. nov. Lowest-occurrence Zone (Hettangian/Sinemurian) . These zones are correlated to previousy established radiolarian assemblages and zones in Japan and other regions. Age assignment of the zones is also discussed on the basis of the correlation and other available chronological data. The original stratigraphic succession of the Triassic in the studied area, which ranges in age from Early Triassic to Early Jurassic, is more than 100 m in thickness and can be reconstructed in detail. The succession is subdivided into seven units based on lithologic features. Each unit was probably accumulated under a particular sedimentary condition, thus successive changes of paleoceanographic environments during Triassic time can be traced continuously. Nine new genera including Ayrtonius, Blonzella, Braginella, Bulbocampe, Enoplocampe, Lysenzelas, Parvibrachiale, Spongoxystris and Veles, and 47 new species are described herein. A comprehensive list of identified taxa is presented.
The heat stress (hs) response is universal to all organisms. As the cell senses increase in temperature, heat stress transcription factors (Hsfs) are activated to upregulate the expression of a number of genes encoding heat stress proteins (Hsp) which act as molecular chaperones to protect cells against heat damages. In higher plants, the phenomenon seems to be unusually complex both at the level of Hsfs and Hsps (e.g., 21 Hsf encoding genes in Arabidopsis and at least 17 in tomato). Upon prolonged hs, another characteristic property of plant cells is the assembly of large cytosolic aggregates called heat stress granules (HSG), which are composed of Hsps, HsfA2, RNA and RNA-binding proteins. The present work was aimed to understand plant hs response using tomato as a model system. To study the function of tomato Hsfs in their native system, we generated transgenic tomato lines altered in expression of HsfA1, HsfA2, and HsfB1. Tomato plants with 10-fold overexpression of HsfA1 (OE plants) were characterised by integration of a single HsfA1 expression cassette, whereas the plants harbouring a tandem inverted repeat (IR) of the cassette showed cosuppression of HsfA1 (CS plants). The lack of HsfA1 expression in CS plants results from posttranscriptional gene silencing connected with the formation of small interfering RNA (siRNA). Under normal growth conditions, major developmental features were similar for wild-type (WT), OE and CS plants. However, in contrast to the former two, CS plants and fruits were extremely sensitive to elevated temperature because hs-induced synthesis of major chaperones and Hsfs was strongly reduced or lacking. Despite the complexity of the plant Hsf family, the function of tomato HsfA1 is unique as master regulator of induced thermotolerance. On the other hand, maintenance of essential chaperones in CS plants during seed development suggests involvement of other Hsfs and/or transcription factor(s). HsfB1 and HsfA2 transgenic tomato plants, unaffected in thermotolerance, further supported the function of HsfA1 as the major factor regulating hs-inducible genes. Hs87 independent phenotypes of plants with altered expression of HsfB1 indicates developmental role of this Hsf. Using transient reporter assays with mesophyll protoplasts from WT tomato, we demonstrated that plasmids encoding Hsfs A1, A2 and A3 were well expressed which could function as activators for reporter gene expression. However, in protoplasts derived from CS plants, plasmids encoding HsfA2 and HsfA3 were normally expressed but even higher amounts of HsfA1 expression plasmids were completely silenced. Therefore, silencing of HsfA1 in CS plants was also reproduced in its mesophyll protoplasts. Lacking thermotolerance in CS protoplasts could be restored after transformation with expression plasmids encoding functionally equivalent HsfA2 or HsfA3 resulting in (i) expression of chaperones, (ii) survival of the cells at otherwise lethal temperature, (iii) thermoprotection of firefly luciferase, and (iv) assembly of heat stress granules (HSGs). The strong silencing caused by an IR in CS plants opened the possibility of a broad use of RNAi for gene knock-down also in the transient system of mesophyll protoplasts. Using this technology, we attempted to dissect essential components of thermotolerance and HSG assembly. We demonstrated the previously reported function of chaperones such as Hsp70 and Hsp101, and could discriminate the in vivo chaperone functions of different isoforms of Hsp20 and Hsp70 proteins. Hsp17-CI, Hsp70 (hs-inducible isoforms), and Hsp101 are absolutely essential chaperones for thermotolerance in plants. Furthermore, the results also show that despite Hsp17-CI and -CII being major components of HSG complexes, they are dispensable for assembly of these complexes. Based on these results, it is proposed that in the transient protoplast system an approach with gene-specific IRs can be used to discriminate functions of closely related isoforms among protein-families and to dissect complex protein networks.
Plastids are complex plant organelles fulfilling essential physiological functions, such as photosynthesis and amino acid metabolism. The majority of proteins required for these functions are encoded in the nuclear genome and synthesized on cytosolic ribosomes as precursors, which are subsequently translocated across the outer and inner membrane of the organelle. Their targeting to the organelle is ensured by a so called transit peptide, which is specifically recognized by GTP-dependent receptors Toc159 and Toc34 at the cytosolic side of outer envelope. They cooperatively regulate the insertion of the precursor protein into the channel protein Toc75, thereby initiating the translocation process. Toc34 is regarded as the primary receptor, while Toc159 probably provides the driving force for the insertion. Precursor transfer is achieved by the physical interaction between both receptors in the GTP loaded state. One translocon unit, also called the Toc core complex, is formed by four molecules Toc34, four molecules Toc75 and one molecule Toc159. In the GDP-loaded state, Toc34 preferably forms homodimers, whose physiological function was investigated in the presented study. It could be shown that the dissociation of GDP and therefore the nucleotide exchange are inhibited by the homodimeric state of Toc34. Dissociation of the homodimer is induced by the recognition of a precursor protein, which renders the binding of GTP and subsequent interaction with Toc159 possible. Thus, the homodimeric conformation could reflect an inactive state of the translocon, preventing GTP consumption in the absence of a precursor protein. Both homodimerization as well as heterodimerization of the receptor are regulated by phosphorylation, which could be demonstrated by in vitro and in vivo approaches using atToc33 from Arabidopsis thaliana as a model system. Since the phosphorylated form of Toc34 cannot be assembled with the Toc core complex, it can be concluded that the interactions between GTPase domains not only regulate the transfer of precursor proteins, but also warrant the integrity of the translocon.
The chemiosmotic theory suggested by Peter Mitchell (Mitchell, 1961, Nature 191:144-148; see Mitchell, 1979, Science 206:1148-1159 for review) postulated that the energy released upon the oxidation of electron donor substrates is transiently stored as electrochemical proton potential, delta-p across energy-transducing membranes, which acts then as the driving force for the ATP synthesis. Membrane protein complexes can both generate and utilise a transmembrane electrochemical proton potential, either by transmembrane proton transfer or by transmembrane electron transfer coupled to protolytic reactions on opposite sides of the membrane. The dihaem-containing membrane protein complex quinol:fumarate reductase (QFR) from the anaerobic epsilon-proteobacterium Wolinella succinogenes apparently combines both of these mechanisms (Haas et al, 2005, Biochemistry 44:13949-13961; Lancaster et al, 2005, PNAS 102:18860–18865; Mileni et al, 2005, Biochemistry 44:16718-16728; Madej et al, 2006, EMBO J 25:4963-4970). QFR is the terminal enzyme of anaerobic fumarate respiration that allows bacteria to use fumarate as the terminal electron acceptor (Kröger, 1978, Biochim Biophys Acta 505:129-45; Lancaster, 2004, In: Respiration in Archaea and Bacteria Volume 1:57-85). QFR couples the two-electron reduction of fumarate to succinate to the two-electron oxidation of quinol to quinone. QFR contains two haem b groups bound by the transmembrane subunit C, which are termed the ‘proximal haem’, bP, and the ‘distal haem’, bD, according to the relative proximity to the hydrophilic subunits A and B (Lancaster et al, 1999, Nature 402:377-85). The two-electron transfer via the two haem groups has been proposed (Lancaster, 2002, Biochimica et Biophysica Acta 1565:215-231) and demonstrated (Madej et al, 2006, EMBO J 25:4963-4970) to be coupled to a compensatory, parallel transfer of two protons via a transmembrane proton transfer pathway. The two most prominent constituents of the proposed pathway were suggested to be the haem bD ring C propionate and the side chain of amino-acid residue Glu C180, after which the proton transfer pathway was named the ‘E-pathway’ (Lancaster, 2002, Biochimica et Biophysica Acta 565:215-231). The essential role of Glu C180 was supported by site-directed mutagenesis and structural and functional characterization of the enzyme E180Q, where the Glu C180 was replaced with a Gln residue (Lancaster et al, 2005, PNAS 102:18860–18865). Moreover, multiconformer continuum electrostatics (MCCE) calculations (Haas and Lancaster 2004, Biophys J 87:4298-4315) and Fouriertransformed infrared (FTIR) spectroscopy experiments (Haas et al, 2005, Biochemistry 44:13949-13961) indicated the Glu C180 side chain to undergo a combination of a conformational change and protonation upon haem reduction. The contribution of haem bD propionate is less clear, however, a combination of 13C labelling of the haem propionates with redox-induced FTIR experiments (Mileni et al, 2005, Biochemistry 44:16718-16728) and MCCE calculations (Haas and Lancaster, 2004, Biophys J 87:4298-4315) support a change in protonation, possibly accompanied by a change in environment upon haem reduction. These experiments and their results strongly support the existence of the ‘E-pathway’ which is transiently open during the reduction of the haem groups and blocked in the oxidized state of the enzyme (Lancaster, 2002b, Biochim Biophys Acta 1565:215-231). All available crystal structures of the QFR, however, are those of the oxidized enzyme. Therefore, it is advantageous to perform simulations of various redox states of the enzyme to determine for instance, how the side-chain of Glu C180 and haem bD ring C propionate behave upon changes of the redox states of the haem groups and why is the ‘E-pathway’ blocked in the oxidized state of the enzyme. Although the distal haem ring C propionate and Glu C180 were identified as the most prominent components of the proton transfer pathway, it was not clear, on the basis of the structure, how proton transfer could occur between them. In addition, two constituents are not enough to span the membrane region and the additional participants in the proton transfer pathway must be identified. Since an atomistic investigation of proton transfer in this system is not yet possible experimentally, I used available theoretical methods such as classical molecular dynamics (MD) simulation (Alder and Wainwright, 1959, J Phys Chem 31:459-466; McCammon et al, 1977, Nature 267:585-590) and Q-HOP molecular dynamics (Q-HOP MD) simulation (Lill and Helms, 2001, J Chem Phys 115:7993-8005) to investigate the postulated mechanism of electron coupled proton transfer in QFR. MD simulations allowed us to move away from static difference pictures obtained from FTIR experiments and MCCE calculations. The advantage of the MD simulations over the experiments and the simulations performed so far is that the time-dependent properties could now be analyzed. The behaviour of various residues and their side-chains and any environmental changes may be directly observed during MD simulations. Although classical MD simulations cannot be used to study proton transfer reactions, they can provide information on formation of configurations that would allow either direct proton transfer between donor and acceptor residues or indirect proton transfer mediated by water molecules. To avoid the static protonation of residues which is inherent in classical MD simulations, Q-HOP MD simulations were performed which explicitly describe proton transfer reactions by allowing the change of the protonation state of residues ‘on the fly’. The structures obtained after classical molecular dynamics simulations ....
The Indian Hill Mynah (Gracula religiosa) was studied in the field in Assam in north-east India. The aims of the study were two-fold: (i) to understand better this bird's exceptional ability in captivity to imitate human speech; and (ii) to provide background understanding to studies of the importance of early auditory experience and of vocal imitation in the development of normal song patterns in birds. First is given a brief description of the distribution, general behaviour, and breeding biology of this arboreal, sexually isomorphic, semi-gregarious species. The remainder of the monograph deals with vocalizations; these were either tape-recorded in the field, or transcribed directly using a written notation developed for the purpose. Any wild adult Mynah of either sex possesses four categories of vocalizations: (i) 'Chip-call'; a loud piercing squeak made in contexts which include alarm. (ii) 'Um-sound'; a soft grunt, acting in close range social contexts, and (like chip-calls) common to all individuals. (iii) 'Whisper-whistles': several soft sounds of types unique to the individual. (iv) 'Calls': several loud noises, of extremely varied patterns. The bulk of the monograph deals with 'calls', as defined thus. Calls were compared quantitatively with one another by a method developed which measured the degree of overlap of one sonogram with a tracing of a second sonogram. Both by this method and by ear, calls were divided into discrete types, without intermediates. Birds of either sex have a repertoire of usually between five and twelve such call types, some of which are produced much more commonly than others. Repertoires tend to be larger in birds which call more frequently, or which have mates with large repertoires. The repertoire of a given bird stays largely constant from year to year in size. composition, and proportions. No bird shares any of its call types with its mate, but it shares several of them with near neighbours of the same sex. There is a progressive change of dialect with distance, such that birds nesting more than about 14 km apart have no call types in common. No general characteristics of call structure could be found which were indicative of the sex of the caller, but in a known locality the call type made immediately reveals the sex of the bird producing it. Call types are learnt by selective imitation of neighbouring individuals during a young bird's first several months. A call type common in the repertoire of one bird tends also to be common in the repertoire of a neighbour, except at the edge of the limited range of that call type. Which particular call type a calling bird selects from among those in its repertoire is discussed. Few call types could be related to non-auditory contexts. A bird is likely to repeat the call type last made, and also tends to standardize the order in which it produces its different call types; this standard order tends to be the same as that of its neighbours. A birdtends also to reply at once and to standardize the call type it makes in immediate reply to a particular call type of its mate; again, neighbouring pairs of birds tend to use the same standardized call and reply types. The length of the interval between a particular call and its reply tends to be constant in a given pair of birds, and approximately the same in neighbouring pairs. These are all further aspects of extensive but selective vocal imitation by Mynahs of adult birds; other species are not imitated. Information on calling when in contact with other pairs came mainly from playback experiments, when single calls were presented to nesting pairs of Mynahs. Response strength was measured by the incidence of flight, number of subsequent vocalizations, latency of response, and proportion of playbacks ignored. When presented with playbacks of calls of familiar types (of neighbours) and of unknown types (of strangers), birds responded more strongly to the familiar than to the unknown call types. They did, however, respond somewhat to the unknown call types, which were of patterns never previously heard by them, presumably recognizing these as being Mynah calls by their sound quality. Mynahs responded as strongly to playbacks of neighbours' calls which were not in their own repertoire as to playbacks of neighbours' calls which were. A bird tends to match at once the call last heard (either from a tape recorder or from a wild neighbour), itself producing the same call type at once, if it possesses it in its own repertoire. That call type, and others associated with it, also occurs more frequently thereafter. Thus calls heard affect calls made, and vice-versa since other individuals nearby behave similarly. A change of nearest neighbours in successive years was shown to affect one pair's repertoire proportions. Further playback experiments showed that Mynahs were able to distinguish between a single call made by their neighbours and a single call of the same call type made by their mates. Small but consistent differences were found in the sonograms of such calls of the same type made by different birds. The structure of a single call type may change gradually with distance. The development of vocalizations with age is briefly described. In the final discussion sections, the ways in which, and the extent to which, Mynahs are able to determine the species, home locality, sex and individual identity of other Mynahs are outlined. There follow consideration, and comparison with other species: (i) of various aspects of repertoires; (ii) of the distribution of call types among different individuals; (iii) of the dynamic aspects of calling, and a scheme is proposed which accounts for the selection for utterance of a particular call type from the repertoire; and (iv) of the organization and coordination of calling. The lack of imitation of other species in the wild is discussed, and contrasted with the several ways in which wild Mynahs imitate one another in various aspects of their calling.
Carnian (Upper Triassic) fishes from Polzberg bei Lunz have been known since 1886 but no comprehensive account has been published. Eleven species are described nine of which, Saurichthys calcaratus, Polzbergia brochatus, Peltoplellrus dinlmptus, Habroichthys gregarius, Nannolepis elegans, Phaidrosoma lunzensis, Elpistoichthys pectinatus, E. striolatus and Pholidophoretes salvus are new, and two others, Thoracopterus niederristi Bronn and Gigantopterus telleri Abel, previously little-known. New supraspecific taxa defined are: the order Polzbergiiformes, the family Thoracopteridae and the genera Polzbergia, Nannolepis, Phaidrosoma, Elpistoichthys and Pholidophoretes. Habroichthys. Thoracopterus, Gigantopterus and Nannolepis show an unusual skull-roof pattern and are included in the re-defined order Luganoiiformes. Two new ichthyokentemids considerably extend the known time-range of this family. The genus Pholidophoretes is intermediate between the Archaeomenidae Goodrich 1909, and the Pholidophoridae sensu stricto Nybelin 1966. The Polzberg assemblage was probably mainly marine with a small freshwater contribution; it shows less similarity to the Besano and Raibl assemblages than these do to each other. The Luganoiiformes are probably, but not certainly, monophyletic; relationships within the order are analyzed and a cladogram constructed. The Platysiagiformes, Peltopleuriformes, Luganoiiformes and Cephaloxeniformes could all have been derived from a common ancestor at the Perleidus level and are probably offshoots of the perleidid radiation.