Review of methodological problems of 'Computer cladistics' exemplified with a case study on isopod phylogeny (Crustacea: Isopoda)

Using a computerized phylogenetic analysis of the Isopoda (Crustacea: Peracarida) as source of typical errors and misunderstandings, problems that may occur in computer cladistics are reviewed. It is concluded that in addition to the errors that are possible in a conventional Hennigian analysis some specific methodological problems exist in computer cladistics. It is recommended that the OTU be replaced by the groundpattern concept. Tree statistics are not useful for comparing different competing hypotheses. Arguments ought to concentrate on the hypo-thetico-deductive steps of the analysis, i.e. on character analysis. The use of computers does not add objectivity to character analysis. Single outgroup taxa should not be used in assessing the character states of ingroups. Concerning isopod phylogeny, it is argued here that the tail fan of the Isopoda can probably be derived from the eumalacostracan groundpattern and did not evolve de novo within the Isopoda.     

Hypothetical Ancestors and Rooting in Cladistic Analysis

Most hypothetical ancestors that are used to root trees in cladistic analyses summarize character-state information in one or more outgroup taxa. Nonetheless, hypothetical ancestors also provide a means of rooting trees using the ontogenetic and paleontological methods of polarizing character transformations, and for incorporating the inferences of more than one of these methods into a single analysis. However, the use of one hypothetical ancestor that combines inferences based on outgroup comparison with those based on other methods of polarizing character transformations to root a cladogram is invalid. Inferences regarding plesiomorphic character states based on outgroup comparison apply to the outgroup node, whereas inferences based on either the ontogenetic or paleontological method apply to the ingroup node. These inferences cannot be combined into a single hypothetical construct. A hypothetical ancestor based on outgroup information is included in the data matrix and used to root the resulting network; however, because this ancestor places potentially problematic constraints on the analysis, the use of actual outgroup taxa is preferable in most instances. Correct use of a hypothetical ancestor inferred with the ontogenetic and paleontological methods involves the Lundberg method in which the shortest ingroup network is rooted at the internode to which the hypothetical ancestor attaches most parsimoniously. Because inferences of polarity based on outgroup comparison cannot be combined directly with those based on other polarization methods, the synthesis of information from all three methods in a single tree must involve taxonomic congruence.     

Phylogenetic interpretation of ontogenetic change: sorting out the actual and artefactual in an empirical case study of centrarchid fishes

Hypothesized relationships between ontogenetic and phylogenetic change in morphological characters were empirically tested in centrarchid fishes by comparing observed patterns of character development with patterns of character evolution as inferred from a representative phylogenetic hypothesis. This phylogeny was based on 56–61 morphological characters that were polarized by outgroup comparison. Through these comparisons, evolutionary changes in character ontogeny were categorized in one of eight classes (terminal addition, terminal deletion, terminal substitution, non-terminal addition, non-terminal deletion, non-terminal substitution, ontogenetic reversal and substitution). The relative frequencies of each of these classes provided an empirical basis from which assumptions underlying hypothesized relationships between ontogeny and phylogeny were tested. In order to test hypothesized relationships between ontogeny and phylogeny that involve assumptions about the relative frequencies of terminal change (e.g. the use of ontogeny as a homology criterion), two additional phylogenies were generated in which terminal addition and terminal deletion were maximized and minimized for all characters. Character state change interpreted from these phylogenies thus represents the maxima and minima of the frequency range of terminal addition and terminal deletion for the 8.7 × 10³⁶ trees possible for centrarchids. It was found for these data that terminal change accounts for c. 75% of the character state change. This suggests either that early ontogeny is conserved in evolution or that interpretation and classification of evolutionary changes in ontogeny is biased in part by the way that characters are recognized, delimited and coded. It was found that ontogenetic interpretation is influenced by two levels of homology decision: an initial decision involving delimitation of the character (the ontogenetic sequence), and the subsequent recognition of homologous components of developmental sequences. Recognition of phylogenetic homology among individual components of developmental sequences is necessary for interpretation of evolutionary changes in ontogeny as either terminal or non-terminal. If development is the primary criterion applied in recognizing individual homologies among parts of ontogenetic sequences, the only possible interpretation of phylogenetic differences is that of terminal change. If homologies of the components cannot be ascertained, recognition of the homology of the developmental sequence as a whole will result in the interpretation of evolutionary differences as substitutions. Particularly when the objective of a study is to discover how ontogeny has evolved, criteria in addition to ontogeny must be used to recognize homology. Interpretation is also dependent upon delimitation within an ontogenetic sequence. This is in part a function of the way that an investigator ‘sees’ and codes characters. Binary and multistate characters influence interpretation differently and predictably. The use of ontogeny for determining phylogenetic polarity as previously proposed rests on the assumptions that ancestral ontogenies are conserved and that character evolution occurs predominantly through terminal addition. It was found for these data that terminal addition may comprise a maximum of 51.9% of the total character state change. It is concluded that the ontogenetic criterion is not a reliable indicator of phylogenetic polarity. Process and pattern data are collected simultaneously by those engaged in comparative morphological studies of development. The set of alternative explanatory processes is limited in the process of observing development. These form necessary starting points for the research of developmental biologists. Separating ‘empirical’ results from interpretational influences requires awareness of potential biases in the course of character selection, coding and interpretation. Consideration of the interpretational problems involved in identifying and classifying phylogenetic changes in ontogeny leads to a re-evaluation of the purpose, usefulness and information conveyed by the current classification system. It is recommended that alternative classification schemes be pursued.     

Two issues in archaeological phylogenetics: taxon construction and outgroup selection

Cladistics is widely used in biology and paleobiology to construct phylogenetic hypotheses, but rarely has it been applied outside those disciplines. There is, however, no reason to suppose that cladistics is not applicable to anything that evolves by cladogenesis and produces a nested hierarchy of taxa. This includes cultural phenomena such as languages and tools recovered from archaeological contexts. Two methodological issues assume primacy in attempts to extend cladistics to archaeological materials: the construction of analytical taxa and the selection of appropriate outgroups. In biology the species is the primary taxonomic unit used, irrespective of the debates that have arisen in phylogenetic theory over the nature of species. Also in biology the phylogenetic history of a group of taxa usually is well enough known that an appropriate taxon can be selected as an outgroup. No analytical unit parallel to the species exists in archaeology, and thus taxa have to be constructed specifically for phylogenetic analysis. One method of constructing taxa is paradigmatic classification, which defines classes (taxa) on the basis of co-occurring, unweighted character states. Once classes have been created, a form of occurrence seriation-an archaeological method based on the theory of cultural transmission and heritability-offers an objective basis for selecting an outgroup.     

Is there a direct ontogenetic criterion in systematics?

Much recent literature focuses on whether ontogenetic information can be used as a direct criterion for determining the polarity of character trasformations in systematic analysis. This paper reviews the relevant literature and concludes that the ontogenetic criterion is dependent on parsimony rather than the sequence observed during ontogeny. It is not, therefore, based on the discredited arguments of recapitulation. From the perspective of phyologenetic systematics the ontogenetic criterion is a valid means of polarizing character transformations that represents a special case of a broader methodology involving parsimony. The alternative perspective perspective of patttern cladistics holds that polarity should be contained within the data and not imposed upon it. Thus, ontogeny is not required to polarize characters, but ontogenetic information can generate unequivocal character interpretations in terms of the relative generality of related attributes, and in the sense that absence precedes presence. Furthermore, ontogeny is central to systematics, providing empirical evidence of character transformation, information on the whole life cycle and an escape from systematics being teleologically related to phylogenetic inference and the theory of evolution.     

PARSIMONY, HOMOLOGY AND THE ANALYSIS OF MULTISTATE CHARACTERS

Abstract— The order of states in a transformation series describes an internested set of synapomorphies. States adjacent to each other in the transformation series thus share a degree of homology not found in the other states. Whether the level of homology is relatively apomorphic is determined by rooting the order with outgroup comparison. The analysis of state order is a homology problem and is solved with a two-step process using similarity and congruence with other characters as criteria. Other methods that have been proposed (e.g. transformation series analysis, non-additive analysis, morphocline analysis, ontogenetic analysis) fail to apply both similarity and congruence, and thus cannot be used independently for determining character state order.     

The phylogenetic interpretation of biological surveys

The ecological attributes of two species may be similar through convergent evolution or common ancestry. The extent of similarity by descent can be evaluated by comparing them with their most closely‐related outgroup in a given phylogeny. I describe a method of nested sister‐group analysis for estimating ecological similarity based on landscape features or on co‐distribution. The phylogeny is dissected into triplets, each comprising two sister taxa and their outgroup. For a triplet at any phylogenetic level, the similarity of sister groups with respect to some given character can be compared with their joint similarity to the outgroup to give a single test of similarity by descent. Each comparison is independent, and the full set of triplets provides a complete accounting of phylogenetic variation at all levels. This procedure was applied to 188 moderately abundant species of dicots in two independent surveys from adjoining districts of midland England, supplemented by physical surveys of landscape attributes obtained from digitized maps of the same districts. The co‐distribution of sister species was consistently more positive than the co‐distribution of random species pairs, demonstrating the existence of a phylogenetic signal at some level. When sister species are compared with their most closely‐related outgroup, however, neither landscape attributes nor co‐distribution showed any overall similarity arising from common ancestry, in the sense that ecological attributes are not generally conserved after lineage splitting. Instead, the distribution of similarity is strikingly similar to random data. The lack of ecological similarity between closely‐related groups was attributed to rapid character change at or shortly after the splitting of lineages, coupled with a lack of correlation between successive lineage splits.     

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.     

The role of fossils in phylogeny reconstruction: Why is it so difficult to integrate paleobiological and neontological evolutionary biology?

Why has it been so difficult to integrate paleontology and “mainstream” evolutionary biology? Two common answers are: (1) the two fields have fundamentally different aims, and (2) the tensions arise out of disciplinary squabbles for funding and prestige. This paper examines the role of fossil data in phylogeny reconstruction in order to assess these two explanations. I argue that while cladistics has provided a framework within which to integrate fossil character data, the stratigraphic (temporal) component of fossil data has been harder to integrate. A close examination of how fossil data have been used in phylogeny reconstruction suggests that neither explanation is adequate. While some of the tensions between the fields may be intellectual “turf wars,” the second explanation downplays the genuine difficulty of combining the distinctive data of the two fields. Furthermore, it is simply not the case that the two fields pursue completely distinct aims. Systematists do disagree about precisely how to represent phylogeny (e.g., minimalist cladograms or trees with varying levels of detail) but given that every tree presupposes a pattern of branching (a cladogram), these aims are not completely distinct. The central problem has been developing methods that allow scientists to incorporate the distinctive bodies of data generated by these two fields. Further case studies will be required to determine if this explanation holds for other areas of interaction between paleontology and neontology.     

Head capsule characters in the Hymenoptera and their phylogenetic implications

The head capsule of a taxon sample of three outgroup and 86 ingroup taxa is examined for characters of possible phylogenetic significance within Hymenoptera. 21 morphological characters are illustrated and scored, and their character evolution explored by mapping them onto a phylogeny recently produced from a large morphological data set. Many of the characters are informative and display unambiguous changes. Most of the character support demonstrated is supportive at the superfamily or family level. In contrast, only few characters corroborate deeper nodes in the phylogeny of Hymenoptera.     

Pattern Cladistics and the ‘Realism–Antirealism Debate’ in the Philosophy of Biology

Despite the amount of work that has been produced on the subject over the years, the ‘transformation of cladistics’ is still a misunderstood episode in the history of comparative biology. Here, I analyze two outstanding, highly contrasting historiographic accounts on the matter, under the perspective of an influential dichotomy in the philosophy of science: the opposition between Scientific Realism and Empiricism. Placing special emphasis on the notion of ‘causal grounding’ of morphological characters (sensu Olivier Rieppel) in modern developmental biology’s (mechanistic) theories, I arrive at the conclusion that a ‘new transformation of cladistics’ is philosophically plausible. This ‘reformed’ understanding of ‘pattern cladistics’ entails retaining the interpretation of cladograms as ‘schemes of synapomorphies’, but in association to construing cladogram nodes as ‘developmental-genetic taxic homologies’, instead of ‘standard Darwinian ancestors’. The reinterpretation of pattern cladistics presented here additionally proposes to take Bas Van Fraassen’s ‘constructive empiricism’ as a philosophical stance that could properly support such analysis of developmental-genetic data for systematic purposes. The latter suggestion is justified through a reappraisal of previous ideas developed by prominent pattern cladists (mainly, Colin Patterson), which concerned a scientifically efficient ‘observable/non-observable distinction’ linked to the conceptual pair ‘ontogeny and phylogeny’. Finally, I argue that a robust articulation of Antirealist alternatives in systematics may provide a rational basis for its disciplinary separation from evolutionary biology, as well as for a critical reconsideration of the proper role of certain Scientific Realist positions, currently popular in comparative biology.     

ONTOGENY AND CHARACTER PHYLOGENY

Abstract— The doctrine of recapitulation was long ago exchanged for a view that ontogeny is orderly and recapitulates ancestral ontogenies. Only recently has the pattern of ontogeny, as a means for determining individual character phylogenies (polarities), been explicitly explored by Gareth Nelson and other cladists. Constraints on outgroup comparisons are suggested to apply to Nelson's rule equally, and the supposed distinction between "direct" and "indirect" methods is suggested to be nonexistent. Nelson's rule is concluded to measure character adjacency directly, but polarity indirectly. Nelson's (ontogenetic) rule is compared to the Outgroup Rule of Watrous and Wheeler, based on 60 postembryonic larval characters of three species of slime-mold beetles of the genus Agathidium ( onisemdes Palisot de Beauvois, pulchrum LeConte, aristerium Wheeler), and reference to larvae of Amsotoma basalts (Le Conte) as an outgroup. Polarities were hypothesized based on each rule and cladograms constructed with the microcomputer programs Henning86 (J. S. Farris) and CLADOS (K. C. Nixon). With levels of analytical error, or homoplasy, measured as the consistency index, the results were compared and the least homoplastic solutions preferred. It is concluded that the outgroup rule and Nelson's rule arc of about the same efficacy as criteria for polarity, and that each is ultimately justified on the basis of parsimony. The following hypothesis of relationships is accepted for the species studied: ( Anisotoma basalts + ( Agathidium oniscoides + ( Agathidium pulchrum + Agathidium arislerium ))).     

ONTOGENY AND PHYLOGENETIC SYSTEMATICS

Abstract— Dedifferentiation, paedomorphosis, and the insertion and deletion of developmental stages make it impossible to deduce the genealogical hierarchy from only ontogenetic transformation series. Like the outgroup criterion, ontogenetic character precedence is not theory-neutral and to use it to deduce genealogy requires certain assumptions.
If scientists are going to use logically unbeatable theories about the world, they might as well give up natural science and take up religion (Lewontin, 1972: 181).     

Problems with the use of cladistic analysis in palaeoanthropology

Cladistic analysis is a popular method for reconstructing evolutionary relationships on the human lineage. However, it has limitations and hidden assumptions that are often not considered by palaeoanthropologists. Some researchers who are opposed to its use regard cladistics as the preferred method for taxonomic «splitters» and claim it has lead to a revitalisation of typology. Typology remains a part of human evolutionary studies, regardless of the acceptance or use of cladistics. The assumption/preference for «splitting» over «lumping» in cladistics (alpha) taxonomy and the general failure to evaluate (post-hoc) such taxonomies have served to reinforce this assertion.

Researchers have also adopted a number of practices that are logically untenable or introduce considerable error. The evolutionary trend of human encephalisation, apparently isometric with body size, and concurrent reduction in the gut and masticatory apparatus, suggests continuous cladistic characters are biased by problems of body size.

The method suffers a logical weakness, or circularity, leading to bias when characters with multiple states are used. Coding of such characters can only be done using prior criteria, and this is usually done using an existing phylogenetic scheme. Another problem with coding character states is the handling of variation within species. While this form of variation is usually ignored by palaeoanthropologists, when characters are recognised as varying, their treatment as a separate state adds considerable error to cladograms.

The genetic proximity of humans, chimpanzees and gorillas has important implications for cladistic analyses. It is argued that chimpanzees and gorillas should be treated as ingroup taxa and an alternative outgroup such as orangutans should be used, or an (hypothetical) ancestral body plan developed. Making chimpanzees and gorillas ingroup taxa would considerably enhance the biological utility of anthropological cladograms.

All published human cladograms fail to meet standard quality criteria indicating that none of them may be considered reliable. The continuing uncertainty over the number and composition of fossil human species is the largest single source of error for cladistics and human phylogenetic reconstruction.     

The role of parsimony,outgroup analysis,and theory of evolution in phylogenetic systematics

It is argued that both the principle of parsimony and the theory of evolution, especially that of natural selection, are essential analytical tools in phylogenetic systematics, whereas the widely used outgroup analysis is completely useless and may even be misleading. In any systematic analysis, two types of patterns of characters and character states must be discriminated which are referred to as completely and incompletely resolved. In the former, all known species are presented in which the characters and their states studied occur, whereas in the latter this is not the case. Dependent on its structure, a pattern of characters and their states may be explained by either a unique or by various conflicting, equally most parsimonious hypotheses of relationships. The so-called permutation method is introduced which facilitates finding the conflicting, equally most parsimonious hypotheses of relationships. The utility of the principle of parsimony is limited by the uncertainty as to whether its application in systematics must refer to the minimum number of steps needed to explain a pattern of characterts and their states most parsimoniously or to the minimum number of evolutionary events assumed to have caused these steps. Although these numbers may differ, the former is usually preferred for simplicity. The types of outgroup analysis are shown to exist which are termed parsimony analysis based on test samples and cladistic type of outgroup analysis. Essentially, the former is used for analysing incompletely resolved patterns of characters and their states, the latter for analysing completely resolved ones. Both types are shown to be completely useless for rejecting even one of various conflicting, equally most parsimonious hypotheses of relationships. According to contemporary knowledge, this task can be accomplished only by employing the theory of evolution (including the theory of natural selection). But even then, many phylogenetic-systematic problems will remain unsolved. In such cases, arbitrary algorithms like those offered by phenetics can at best offer pseudosolutions to open problems. Despite its limitations, phylogenetic systematics is superior to any kind of aphylogenetic systematics (transformed cladistics included) in approaching a (not: the) “general reference system” of organisms.     

The phylogeny of Hildoceratidae (Cephalopoda,Ammonitida) resolved by an integrated coding scheme of the conch

Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework.     

Models and reality: Doctrine and practicality in classification

Phenetic classification corresponds to no biological model and lacks a sound philosophical basis. Cladistics (ignoring meaningless “transformed cladistics”) assumes divergent evolution and, usually, that best estimates of phylogeny are obtained by parsimony principles, both questionable assumptions at times. It is better than phenetics since more-or-less testable hypotheses are generated, but pitfalls are many, in data selection and interpretation (as to homology), and in commensurability of units and direction of change. Above all we learn: homoplasy is rife in nature. Much bad cladistics has been done. If it is to reflect phylogeny, classification cannot be artificially stabilized, but is its only aim to express (hypothesized) cladistic patterns? And can it do that with any degree of overall assurance? Biologists are legitimately interested in defining grades as well as clades. Recognition of an unequivocal clade-grade frequently leaves a paraphyletic grade residue that cannot itself be unequivocally resolved. This is a real problem that requires attention in formal taxonomy and in applying cladistics. Primarily morphological cladistics will be increasingly supplanted by molecular (nucleotide-sequence) cladistics. The role of evolutionary taxonomy will change accordingly.     

Systematics of Bitheca, a new genus of New World Sphaeroceridae (Diptera)

Abstract. The genus Bitheca is described for a distinctive clade of small Limosininae whose range extends from Canada south to Ecuador. The following twelve species are described as new: agarica, boleta, caballa, dispar, ejuncida, fundata, grossa, horrida, involuta, jubilata, kappa and lambda. A key to the species of Bitheca is provided and their phylogeny is discussed. Pterogramma steyskali Deeming and Minilimosina masoni Marshall are moved into the genus Bitheca. A cladogram is provided and its derivation explained in detail, with a discussion of the use of character weighting in cladistics.     

Combining ontogenetic and evolutionary scales of morphological disparity: a study of early Jurassic ammonites

Two major research themes in Evolutionary Developmental Biology and in Paleobiology, respectively, have each become central for the analysis and interpretation of morphological changes in evolution: the study of ontogeny/phylogeny connections, mainly within the widespread and controversial framework of heterochrony; and the study of morphological disparity, the morphological signal of biodiversity, describing secular changes in morphospace occupation during the history of any given clade. Although enriching in their respective fields, these two themes have remained rather isolated to date, despite the potential value of integrating them as some recent studies begin to suggest. Here, we explore the recent notion of developmental morphospace-morphospace carrying ontogenetic information-as a potential tool for bridging the gap between disparity dynamics and developmental dynamics. We elaborate this approach with a case study of Early Jurassic ammonite family Hildoceratidae (Mollusca, Cephalopoda). Morphometric analyses of the shell shape of 20 species spanning the morphological spectrum of the family are used to quantify and contrast juvenile and adult disparity levels. Adult disparity is significantly greater than juvenile disparity at the family level; yet, some subclades also display different patterns. In addition, comparisons of ontogenetic trajectories underline the prevalence of heterochrony-based evolutionary modifications within subfamilies (via ontogenetic scaling); they also point to the probable existence of pervasive developmental constraints structuring inhomogeneous morphospace occupation.