Relative apparent synapomorphy analysis (RASA). I: The statistical measurement of phylogenetic signal

We have developed a new approach to the measurement of phylogenetic signal in character state matrices called relative apparent synapomorphy analysis (RASA). RASA provides a deterministic, statistical measure of natural cladistic hierarchy (phylogenetic signal) in character state matrices. The method works by determining whether a measure of the rate of increase of cladistic similarity among pairs of taxa as a function of phenetic similarity is greater than a null equiprobable rate of increase. Our investigation of the utility and limitations of RASA using simulated and bacteriophage T7 data sets indicates that the method has numerous advantages over existing measures of signal. A first advantage is computational efficiency. A second advantage is that RASA employs known methods of statistical inference, providing measurable sensitivity and power. The performance of RASA is examined under various conditions of branching evolution as the number of characters, character states per character, and mutations per branch length are varied. RASA appears to provide an unbiased and reliable measure of phylogenetic signal, and the general approach promises to be useful in the development of new techniques that should increase the rigor and reliability of phylogenetic estimates.      

A phylogenetic analysis of the land plants

Phylogenetic systematics (cladistics) is a theory of phylogeny reconstruction and classification widely used in zoology. Taxa are grouped hierarchically by the sharing of derived (advanced) characters. The information is expressed in a cladogram, a best estimate of a phylogeny. Plant systematists generally use a phenetic system, grouping taxa on overall similarity which results in many groups being formed, at least in part, on the basis of shared primitive characters.
The methods of phylogenetic systematics are used to create a preliminary cladogram of land plants. The current classification of land plants is criticized for its inclusion of many groups which are not monophyletic.
Objections to the use of phylogenetic systematics in botany, apparent convergences within major groups and frequent hybridization, are shown to be invalid. It is concluded that cladistic analysis presents the best estimate of die natural hierarchy of organisms, and should be adopted by plant systematists in their assessment of plant interrelationships.     

Evolution and character congruence in some western Indian Ocean Phelsuma: Numerical analysis of biochemistry, shape and scalation

The evolution of five island populations of Green gecko, representing inter- and intra-specific divergence, was studied using biochemical data, scalation and shape. The data were numerically analysed using ordination analyses for the phenetic classification and Wagner trees to hypothesize the phylogeny. These studies revealed three phenetic groups corresponding to three mono-phyletic lineages. The numerical analysis of morphological data agreed with the numerical analysis of biochemical data. It is concluded that the classification based on biochemical affinities differed from the previous classification based on conventional analysis of morphology due to methodological and philosophical differences rather than differences between morphological and biochemical evolution.
The ordination analyses were very congruent between data sets (biochemical, shape, scalation, total) and the Wagner trees were generally congruent between data sets. Some Wagner trees based on scalation data were incongruent. The phenetic and cladistic classifications corresponded to each other but differed from the conventional classification. The phylogenetic analysis of the total data set indicated that the three specific lineages showed relatively equal anagenesis. However, anagenic divergence differed markedly between character types. It is suggested that a range of character types be used when studying anagenesis.     

Infrageneric relationships within Pogostemon Desf. (Labiatae)

Cladistic and phenetic analyses of the morphology of 79 species of Pogostemon Desf. sens lato have been undertaken. The cladistic analysis included 135 character state differences encoded as 41 binary or multistate characters. The phenetic analysis included 52 metric variables and 54 binary variables. There is a very good correspondence between the results of cladistic and phenetic analyses based on different data sets. The existence of several clearly marked clades/clusters has been demonstrated. Some groups/clades are similar to those previously taxonomically recognized but the relationships and membership of infra-generic groups have been clarified by these analyses, validating a revised infra-generic classification which has been proposed.     

Methods of systematic analysis: the relative superiority of phylogenetic systematics

The superiority of cladistic methods to both synthetic and phenetic methods is briefly advanced and reviewed. Cladistics creates testable hypotheses of phylogeny that also give a highly informative summary of available data. Thus it best fits the criteria for a method for determining the general reference classification in biology. For protistologists in particular, cladistics is especially useful. Inundated by an abundance of ultrastructural, biochemical, and cell biological information, protistologists could be greatly helped by the informative way in which cladistics orders and summarizes the data. In addition to classifying protist taxa, hypotheses about the evolution of cell organelles and cellular could be scientifically formulated and tested by cladistics . Because cladistic classifications best summarize the data, they would also be best for making predictions about taxa and characters. They would, for the same reason, be the most stable. Widespread adoption of cladistic methods would serve to stabilize the now fluid state of protist taxonomy. It is for all of these reasons that such methods best suit the needs of the evolutionary protistologist .     

Methods of systematic analysis: The relative superiority of phylogenetic systematics

The superiority of cladistic methods to both synthetic and phenetic methods is briefly advanced and reviewed. Cladistics creates testable hypotheses of phylogeny that also give a highly informative summary of available data. Thus it best fits the criteria for a method for determining the general reference classification in biology.For protistologists in particular, cladistics is especially useful. Inundated by an abundance of ultrastructural, biochemical, and cell biological information, protistologists could be greatly helped by the informative way in which cladistics orders and summarizes the data. In addition to classifying protist taxa, hypotheses about the evolution of cell organelles and cellular could be scientifically formulated and tested by cladistics. Because cladistic classifications best summarize the data, they would also be best for making predictions about taxa and characters. They would, for the same reason, be the most stable. Widespread adoption of cladistic methods would serve to stabilize the now fluid state of protist taxonomy. It is for all of these reasons that such methods best suit the needs of the evolutionary protistologist.     

Classification of the family Plectonemertidae (Nemertea): a phenetic comparison

A phenetic classification based on overall morphological similarity between the species in the family Plectonemertidae (genera Plectonemertes, Campbellonemertes, Potamonemertes, Leptonemertes, Katechonemertes, Argonemertes, Anliponemertes, and Acteonemertes ) was undertaken and the result compared with a cladistic and an evolutionary classification. Similarity between species was computed by Gower's general coefficient of similarity and various techniques were used to find patterns in the similarity matrix: single-linkage, average-linkage, and complete-linkage clustering, together with principal coordinate analysis. Although the explicit aim of phenetics is not to estimate the phylogeny, the classification based on overall similarity still portrays phylogeny better than an intuitive assessment of morphological similarity, as judged by the cladistic analysis. The classification does not support the previously proposed hypothesis that the two freshwater genera Campbellonemertes and Potamonemertes have descended from a terrestrial ancestor.     

Cladistic and phenetic studies of genera of Afrotropical Cryptobiina (Coleoptera,Staphylinidae, Paederinae) II. The phenetic approach

The phenetics of 12 genera of Afrotropical Cryptobiina have been analysed by multivariate statistics. Three analyses (principal component analysis, canonical discriminant analysis and cluster analysis) have been used to determine the relative positions of the taxa in a morphospace. These analyses have allowed us to confirm the generic divisions based on alpha-systematic and phylogenetic grounds. While phenetic systematics and cladistic systematics have often been opposed, this study shows that both approaches are, in the end, complementary. Phenetical analysis allow, in addition to a quantitative definition of the genera, the isolation of ethological or ecological characters, while the cladistic approch gives to them precision and a phylogenetic sense. The three types of analyses point out the importance in Cryptobiina of the size, length of the scapus and width of the neck, all three characters which were also designated as significant characters by the phylogenetic analysis.     

Quantitative taxonomic analyses of Actinidia (Actinidiaceae) in China based on micromorphological characters of foliar trichomes

There are various arguments on classification of the genus Actinidia Lindl., a genus with approximately 63 species, 59 of which have been found in China. The paper investigated the characteristics of foliar trichomes of 35 taxa from China under optical microscope, including size, celluar structure, distribution and density. According to their micromorphological characteristics, foliar trichomes can be classified into the following six categories: 1) single-cell hairs; 2) uniseriate hairs, including linear, bulbous, twisted, straight-walled, and bent-walled hairs; 3) multiseriate hairs, including twisted, straight-walled and gradually sharpening, straight-walled and suddenly sharpening, bent-walled and gradually sharpening, and suddenly sharpening hairs; 4) multiseriate thick hairs, including pillar hairs, gradually sharpening thick hairs, and suddenly sharpening thick hairs; 5) stellate hairs, including parenchyma-stellate and sclerenchyma-stellate (normal state and special states such as rosulate, peltate-stellate, and overlopping-stellate) hairs; and 6) dichotomous hairs. On the basis of the micromorphological characteristics of foliar trichomes in Actinidia, with Clematoclethra lasioclada as an outgroup, both the quantitative cladistic analysis and phenetic analysis were performed using Wagner method and UPGMA clustering method respectively to reconstruct the phylogeny of Actinidia in China. The phylogenetic tree generated by cladistic analysis suggests that the sect. Leiocarpae be a monophyletic group, but other three sections, i.e., sect. Maculatae, sect. Strigosae and sect. Stellatae, be non-monophyletic groups. The results obtained from the phenetic analysis reflect relationships among the taxa of Actinidia in China, especially a close relationship between A. chinensis and A. deliciosa, and a relatively remote relationship between A. callosa var. henryi and A. callosa var. discolor. In conclusion, the micromorphological characters of foliar trichomes and the methods of quantitative taxonomic analysis are of key importance tostudies on phylogenetic and phenetic relationships of Actinidia.     

Mapping cladograms into morphospaces

In cladistic analyses, taxa are grouped hierarchically into clades according to shared apomorphic character states to construct cladograms; cladograms are interpretable as phylogenetic hypotheses. In morphological space analyses, organism forms are represented as points in morphospaces; point proximities in morphospaces represent similarities that might be attributable to phenetic convergence and, consequently, may correspond inaccurately with hypothesized evolutionary relationships. A method for synthesizing phylogenetic results that are interpreted from cladistic analyses with phenetic results that are obtained from morphological space analyses is presented here; in particular, points that represent forms typifying taxa in morphospace are assigned as terminal nodes for appropriate cladograms that are mapped into morphospaces by positioning nonterminal nodes and orienting internodes according to a geometric algorithm. Nonterminal nodes may be interpreted as ancestors in phylogenetic hypotheses and occupy positions that represent particular organism forms in morphospaces. By mapping cladograms into morphospaces, therefore, evolutionary morphologists can reconstruct ancestral morphologies and test historical transformation hypotheses.     

Paraphyly,ancestors, and the goals of taxonomy: A botanical defense of cladism

Cronquist (1987) criticizes cladism for its rejection of paraphyletic groups, which he would retain if he feels they are “conceptually useful.” We argue that paraphyletic higher taxa are artificial classes created by taxonomists who wish to emphasize particular characters or phenetic “gaps,” and that formal recognition of such taxa conveys a misleading picture of common ancestry and character evolution. In our view, classifications should accurately reflect the nested hierarchy of monophyletic groups that is the natural outcome of the evolutionary process. Such systems facilitate the study of evolution and provide an efficient summary of character distributions. Paraphyletic groups, such as “prokaryotes,” “green algae,” “bryophytes,” and “gymnosperms,” should be abandoned, as continued recognition of such groups will only serve to retard progress in understanding evolution. Contrary to Cronquist’s (1987) assertions, cladistic theory is not at odds with standard views on speciation and the existence of ancestors. Groups of interbreeding organisms can continue to exist after giving rise to descendant species, and there are several ways in which such groups, whether extant or extinct, can be incorporated into cladistic classification. In contrast, paraphyletic higher taxa are neither cohesive (integrated by gene flow) nor whole, do not serve as ancestors, and are unacceptable in the phylogenetic system. Fossils may be of great value in assessing phylogenetic relationships and are readily accommodated in cladistic classification. Cladistic studies are helping to answer major questions about plant evolution, and we anticipate increased efforts to develop a truly phylogenetic system.     

Cladistic and phenetic analysis of allozyme data for nine species of sea anemones of the family Actiniidae (Cnidaria: Anthozoa)

The aim of this study was to infer from allozyme data the phylogenetic relationships of nine species of actiniid sea anemones, and also use these data to assess the various methods (phenetic and cladistic) available for phylogenetic analysis. Starch gel electrophoresis was used to obtain genetic data from 13 gene loci. The anemone Metridium senile, from the family Metridiidae, was used as an outgroup. For the phenetic analysis a matrix of pairwise unbiased genetic distances was computed and, from this, dendrograms were produced both by the Wagner distance and the UPGMA methods. For the cladistic analyses three different approaches were used: the first was to treat the allele as a binary character; this was investigated using a Wagner parsimony algorithm. Another approach used was to consider the locus as an unordered character, using the alleles as states. Finally, we used the locus as an ordered multistate character, where mutation, fixation and elimination of each allele were treated as evolutionary novelties, and the heterozygotes were used as cues for the construction of transformation series. The trees produced by the phenetic and cladistic methods were highly congruent. This result suggests that allozymes can be used to produce phylogenetic hypotheses at higher taxonomic levels than those at which they are more usually employed. The Solé difference between the various trees was the relative positions of Bunodosoma caissarum and Bunodactis verrucosa in relation to the two species of Urticina. This difference was probably due to a high rate of anagenic change in B. verrucosa, which distorted the UPGMA dendrogram. The genera Actinia and Urticina appeared monophyletic in all of the trees produced. Also, the sea anemones with specialized column structures such as verrucae and vesicles (U.felina, U. eques, B. verrucosa, B. caissarum) formed a monophyletic cluster, a result compatible with the suggestion that these structures may have appeared only once in the evolutionary history of the Actiniidae.     

Classification of the family Plectonemertidae (Nemertea): a phenetic comparison

A phenetic classification based on overall morphological similarity between the species in the family Plectonemertidae (genera Plectonemertes, Campbellonemertes, Potamonemertes, Leptonemertes, Katechonemertes, Argonemertes, Anliponemertes, and Acteonemertes) was undertaken and the result compared with a cladistic and an evolutionary classification. Similarity between species was computed by Gower's general coefficient of similarity and various techniques were used to find patterns in the similarity matrix: single-linkage, average-linkage, and complete-linkage clustering, together with principal coordinate analysis. Although the explicit aim of phenetics is not to estimate the phylogeny, the classification based on overall similarity still portrays phylogeny better than an intuitive assessment of morphological similarity, as judged by the cladistic analysis. The classification does not support the previously proposed hypothesis that the two freshwater genera Campbellonemertes and Potamonemertes have descended from a terrestrial ancestor.     

Incongruence between cladistic and taxonomic systems

Cladistic and taxonomic treatments of the same plant group usually exhibit a mixture of congruences and incongruences. The question arises in the case of the incongruences as to which version is right and which is wrong. Many cladists believe that cladistics is a superior approach and gives the best results. There are several conceptual and methodological differences between cladistics and taxonomy that cause incongruence. One important conceptual difference is the use of different criteria for grouping: order of branching vs. similarity and difference (clades vs. taxa). Another is the policy regarding paraphyletic groups: to ban them in cladistics but ignore the ban in taxonomy. These two differences automatically lead to some incongruences. One approach is not right and the other wrong; each is operating by its own standards. However, when cladists apply the paraphyly rule to a taxonomic system and conclude that it needs revision to eliminate paraphyly, as cladists often do, they are judging the taxonomic system by a wrong standard. Several differences between the two schools in the use and handling of characters can also cause incongruence. First consider phenetic characters. Taxonomy uses a very wide range of these, whereas phenetic cladistics sets restrictions on the selection of characters, which deprive it of potentially useful evidence. Taxonomic systems generally rest on a broader empirical foundation than phenetic cladistic systems. Next, consider molecular cladistics, which is the leader in the use of DNA evidence. Two sources of incongruence between molecular cladistics and taxonomic systems can come into play here. First, the molecular evidence used in cladistics comes mainly from cytoplasmic organelles, whereas taxonomic systems are based on characters that are determined mainly by the chromosomal genome. More generally, the database in a molecular cladogram is, in itself, too narrow to serve as a foundation for an organismic classification. In cases of incongruence, the molecular evidence can be a reliable indicator of taxonomic relationships sometimes, misleading other times, and may afford no clear basis for a systematic decision. In this situation, it is helpful, indeed necessary, to integrate the molecular evidence with the phenetic evidence and bring more characters to bear on the question.     

A phenetic and cladistic study of spermatozoal ultrastructure in the Oligochaeta (Annelids)

Spermatozoal ultrastructure of nine oligochaete families has been examined for congruence with phylogenetic and taxonomic systems for the Oligochaeta based on general morphology, particularly the holomorphological hennigian analysis of Jamieson (1978a, 1980, 1983). Estimation of congruence has been made following phenetic and cladistic (phylogenetic) analysis. Correspondence, in phenograms and phylograms, of sperm types with taxonomic and phylogenetic groupings previously recognized is generally good. Departure from this rule in the similarity of the phreodrilid sperm to that of the Lumbricina suggests a corresponding alteration of fertilization biology in the phreodrilids. The results indicate that the Haplotaxidae lie at the base of the opisthopores though they do not unequivocally contraindicate acceptance of a Haplotaxis like form as a stem form of the Haplotaxida (opisthopores and Haplotaxidae) and Tubificida. An even more basal position for prosopores, now represented by the Lumbriculida, cannot yet be dismissed.     

中国猕猴桃属植物叶表皮毛策形态特征及数量分类分析

选取国产猕猴桃３５个分类群的代表植株,应用光学显微对其新鲜叶表皮毛的微形态特征,形体大小、细胞结构、分布和密度等多态性状和数量性状进行了观察和测量,该属植物的叶表皮毛微形态特征可归纳为６个类型：１）单细胞毛;２）单列多细胞毛,每形单列毛、泡状单列毛、扭曲毛、直壁单列毛、曲壁单列毛;３）多列渐尖毛和急尖毛,包括多列曲壁渐尖毛和急尖毛,多列直壁渐尖毛和急尖毛;４）多列粗毛,包括柱状毛,多列渐尖粗毛,多     

A Review on Cladistics

The theoretical bases and approaches of cladistics and some specific problems that, directly or indirectly, rely on cladistic analysis for their revolution, are outlined and discussed. Seven sections comprise this paper: a ) the philosophical foundation of cladistics; b) the theoretical tenets of cladistics; c) the operational procedure of cladisties; d) three schools of classification; e) cladistics and biogeography; f) cladistics and hybrid recognition; and g) is cladistic systematics a scientific theory ? Considerations of scientific methodology involve philosophical questions. From this point, Popper'falsificationism serves a good foundation. Popper emphasizes that all scientific knowledge is hypothetical-deductive, consisting of general statements (theories) that can never be confirmed or verified but only falsified. The theories, that can be tested most effectively, are preferable. Cladistics, aiming at generating accurately expressed and strictly testable systematic hypotheses, is well compatible with this requirement. The principles central to the cladistic theory and methodology are: the Principle of Synapomorphy; the Principle of Strict Monophyly; and the Principle of Strict Parsimony. The first requires forming nested groups by nesting statements about shared evolutionary novelties (synapomorphy) postulated from observed similarities and is the primary one. The second is mainly methodological, subject to modification and compromise. The principle of strict parsimony specifies the most preferable hypothesis (namely the one exhibiting the most congruence in the synapomorphy pattern). The operational procedure that might be followed in formulating and testing hypotheses of the synapomorphy pattern (the cladogram itself) consists of five steps. The erections of monophyletic groups, to a greater or lesser extent, rely on the hypothesis of the previous systematic studies and is the starting point for cladistic analysis. Character analysis, which focuses on character distribution and determination of the polarities, decides the reconstructed phylogeny. A detailed discussion on the methodological principles for identifying transformation sequence is presented. Many algorithms have been designated to infer the cladogram, and are basically of parsimony techniques and Compatibility techiques. The thus yielded cladograms, with their expected pattern of congruent synapomorphies, are tests of a particular hypothesis of synapomorphy and reciprocally synapomorphies are tests of cladistic hypothesis (cladogram). Such reciprocity is a strong stimulus to profound understanding on phylogenetic process and phyletic relationships. The cladogram and the Linnaean classification have the identical logic structure and the set-membership of the two can be made isomorphic. There are three principal approaches to biological classification : cladistics, phenetics and evolutionary classification. Cladistics is the determination of the branching pattern of evolution, and in the context of classification, the development of nested sets based on cladograms. Phenetics is the classification by overall similarities, without regard to evolutionary considerations. Evolutionary classification attempts to consider all meaningful aspects of phylogeny and to use these for making a classification. The last approach has been done intuitively, without explicit methods. An enumeration of their differences and a discussion on their relative merits are presented. Three theoretical approaches have been proposed for interpreting biogeographical history: the phylogenetic theory of biogeography, classical evolutionary biogeography and vicariance biogeography. The former two show some similarities in that they usually look upon biogeography in terms of centers of origin and dispersal from the centers. But the first puts a strong emphasis on the construction of hypotheses about the phylogenetic relationships of the organisms in question and the subsequent inference of their geographic relationships; the second advocates a theory which does not have a precise deductive link with phylogenetic construction and often results in wildly narratative-type hypotheses. The vicariance approach de-emphasizes the concepts of centers of origin and dispersal and attempts to analyse distribution patterns in terms of subdivision (vicariance) of ancestral biotas. The development of the theory of plate tectonics and its universal acceptance enormously stimulate biogeographers to look at the world's continents and oceans from a mobilist point, which, along with the establishment of the rigorous tool of the phylogenetic analysis (cladistics), profoundly reshapes the above three theories. Hybridization and polyploidy are outstanding features of many plant groups. But hybridization, or reticulate evolution, is inconsistent with the basic concepts of cladistics which is an ever-branching pattern. Cladists have suggested several approaches. One of them analyses all the taxa by a standard cladistic procedure and closely examines the cladograms for polytomies and character conflicts that may indicate possible hybrids. Such generated hypothesis of hybridization can be corroborated or falsified by other forms of data, such as distribution, polyploidy, karyotype and pollen fertility. There are three criteria to justify a theory to be scientific: a) whether it is a theory composed of hypotheses strictly falsifiable; b) whether it has predictive effect; and c) whether it has a explanatory value. Cladistic systematics aims at generating cladograms, which are hypotheses of the nested pattern of synapomorphy, phylogenetic process and phyletic relationships, susceptible to testing by postulated synapomorphies. The predictive effect of systematics relies on the acceptance of hypotheses of congruence about the correlation of characters, which has been well founded. For non-systematic biologists, phylogenetic classification can be used as axiom to form a preliminary and fundamental explanation.     

Methods of classifying nemerteans: an assessment

Phenetic, cladistic and phyletic methods of classifying animals are discussed with particular reference to nemerteans. It is concluded that phenetic (numerical) taxonomy is particularly inapplicable to any group of invertebrates for which well defined character differences are relatively few, whilst both the phenetic and cladistic methods fail through their fundamental assumption that convergent evolution is a rare occurrence. Terrestrial and freshwater nemerteans especially demonstrate convergent evolution in many ways; cladistic classifications proposed for these animals are therefore untenable. Convergence is shown to be a common occurrence in other nemerteans also. It is concluded that because the traditional phyletic approach does not implicitly assume that resemblances between organisms are more likely to be due to common ancestry than to convergence, it is far more likely to reveal true evolutionary relationships between taxa.     

Do major species concepts support one, two or more species within Cryptococcus neoformans?

Cryptococcus neoformans, the agent of cryptococcosis, had been considered a homogeneous species until 1949 when the existence of four serotypes was revealed based on the antigenic properties of its polysaccharide capsule. Such heterogeneity of the species, however, remained obscure until the two morphologically distinct teleomorphs of C. neoformans were discovered during the mid 1970s. The teleomorph Filobasidiella neoformans was found to be produced by strains of serotype A and D, and Filobasidiella bacillispora was found to be produced by strains of serotype B and C. Ensuing studies revealed numerous differences between the anamorphs of the two Filobasidiella species with regard to their ecology, epidemiology, pathobiology, biochemistry and genetics. At present, the etiologic agent of cryptococcosis is classified into two species, C. neoformans (serotypes A and D) and Cryptococcus gattii (serotypes B and C). Intraspecific genetic diversity has also been revealed as more genotyping methods have been applied for each serotype. As a result, the number of scientifically valid species within C. neoformans has become a controversial issue because of the differing opinions among taxonomists as to the appropriate definition of a species. There are three major species concepts that govern classification of organisms: phenetic (morphologic, phenotypic), biologic (interbreeding) and cladistic (evolutionary, phylogenetic). Classification of the two C. neoformans species has been based on the phenetic as well as the biologic species concept, which is also supported by the cladistic species concept. In this paper, we review and attest to the validity of the current two-species system in light of the three major species concepts.     

Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera)

Abstract. The higher classification of Tenebrionidae is analysed using numerical phenetic, numerical cladistic and traditional Hennigian methods. In all, eighty characters are examined for about 335 taxa; definitive analyses are made on combinations of eighteen to seventy characters for thirty-three OTUs. At lower levels of relationship (genera and closely related tribes) phenetic and cladistic classifications are shown to be congruent, but at higher levels (tribes and subfamilies) there is marked discordance with phenetic results being more stable. A consensus classification is more similar to the Hennigian cladogram than is any single computer generated cladogram. Two main tribal groups – the Lagrioid and Tenebrionoid groups – are suggested which differ in defensive glands, female anatomy, wing and mouthpart morphology, larval characters and other features. The Tenebrionoid group consists of three main subdivisions – the tenebrionine, coelometopine and diaperine lineages. Changes in classificatory position are recommended for eighty-seven genera and tribes (listed in Appendix E) and implied for numerous others.