Inclusion Of Fossils Research Articles

Fossils Do Not Substantially Improve, and May Even Harm, Estimates of Diversification Rate Heterogeneity.

The fossilized birth-death (FBD) model is a naturally appealing way of directly incorporating fossil information when estimating diversification rates. However, an important yet often overlooked property of the original FBD derivation is that it distinguishes between two types of sampled lineages. Here, we first discuss and demonstrate the impact of severely undersampling, and even not including fossils that represent samples of lineages that also had sampled descendants. We then explore the benefits of including fossils, generally, by implementing and then testing two types of FBD models, including one that converts a fossil set into stratigraphic ranges, in more complex likelihood-based models that assume multiple rate classes across the tree. Under various simulation scenarios, including a scenario that exists far outside the set of models we evaluated, including fossils rarely outperform analyses that exclude them altogether. At best, the inclusion of fossils improves precision but does not influence bias. Similarly, we found that converting the fossil set to stratigraphic ranges, which is one way to remedy the effects of undercounting the number of k-type fossils, results in turnover rates and extinction fraction estimates that are generally underestimated. Although fossils remain essential for understanding diversification through time, in the specific case of understanding diversification given an existing, largely modern tree, they are not especially beneficial. [Fossilized birth-death; fossils; MiSSE; state speciation extinction; stratigraphic ranges; turnover rate.].

Ancient tropical extinctions at high latitudes contributed to the latitudinal diversity gradient.

Global biodiversity currently peaks at the equator and decreases toward the poles. Growing fossil evidence suggest this hump-shaped latitudinal diversity gradient (LDG) has not been persistent through time, with similar diversity across latitudes flattening out the LDG during past greenhouse periods. However, when and how diversity declined at high latitudes to generate the modern LDG remains an open question. Although diversity-loss scenarios have been proposed, they remain mostly undemonstrated. We outline the "asymmetric gradient of extinction and dispersal" framework that contextualizes previous ideas behind the LDG under a time-variable scenario. Using phylogenies and fossils of Testudines, Crocodilia, and Lepidosauria, we find that the hump-shaped LDG could be explained by (1) disproportionate extinctions of high-latitude tropical-adapted clades when climate transitioned from greenhouse to icehouse, and (2) equator-ward biotic dispersals tracking their climatic preferences when tropical biomes became restricted to the equator. Conversely, equivalent diversification rates across latitudes can account for the formation of an ancient flat LDG. The inclusion of fossils in macroevolutionary studies allows revealing time-dependent extinction rates hardly detectable from phylogenies only. This study underscores that the prevailing evolutionary processes generating the LDG during greenhouses differed from those operating during icehouses.

Ambarticus myanmaricus gen. et sp. nov., the first diving beetle from mid-Cretaceous amber of northern Myanmar (Coleoptera, Dytiscidae, Dytiscinae)

Ambarticus myanmaricus gen. et sp. nov. is described from mid-Cretaceous amber of Myanmar based on examination of a single female specimen, and represents the unique taxon inclusion of fossils of the family Dytiscidae (Coleoptera: Adephaga) from Oriental Region. A new tribe, Ambraticini trib. nov. is established. The new taxon is assigned to subfamily Dytiscinae. This description represents one of a limited number of diving beetle taxa described from fossiliferous amber and is currently the oldest known fossil assigned to Dytiscinae. Key morphological characters of the new taxon are illustrated, and compared with tribes of Dytiscinae.

Hunting the wolf: A molecular phylogeny of the wolf spiders (Araneae, Lycosidae)

Lycosids are a diverse family of spiders distributed worldwide. Previous studies recovered some of the deeper splits of the family, but with little support. We present a broad phylogenetic analysis of the Lycosidae including a wide geographic sampling of representatives and all the subfamilies described to date. Additionally, we extend the amount of molecular data used in previous studies (28S, 12S and NADH) through the inclusion of two additional markers, the nuclear H3 and the mitochondrial COI. We estimated the divergence times through the inclusion of fossils as calibration points and used the phylogenetic hypothesis obtained to explore the evolution of particular traits associated with dispersal capabilities. We recovered most of the currently recognized subfamilies with high nodal support. Based on these results, we synonymize Piratinae and Wadicosinae with Zoicinae and Pardosinae, respectively, and revalidate the subfamily Hippasinae. We corroborated that lycosids are a family with a relatively young origin that diversified with the reduction of tropical forests and the advance of open habitats. We show that a gradual accumulation of behavioral traits associated with ambulatory dispersal made Lycosidae the most vagrant subfamily of spiders, with an impressive ability to disperse long distances which helps to explain the worldwide distribution of some very young clades, such as the members of the subfamily Lycosinae.

One (more) step towards a dynamic view of the Latitudinal Diversity Gradient

Biodiversity currently peaks at the equator, decreasing toward the poles. Growing fossil evidence suggest that this hump-shaped latitudinal diversity gradient (LDG) has not been persistent through time, with similar species diversity across latitudes flattening out the LDG during past greenhouse periods. This provides a new starting point for LDG research. Most studies assume the processes shaping the LDG have acted constantly through time and seek to understand why diversity accumulated in the Holarctic at lower levels than at the equator, e.g. as the result of limited dispersal, or higher turnover in Holarctic regions. However, fossil investigations suggest that we need to explain when and why diversity was lost at high latitudes to generate the LDG. Unfortunately, diversity lost scenarios in the Holarctic have been repeatedly proposed but not yet clearly demonstrated. Here, we use diversification approaches for both phylogenies and fossils to study the LDG of Testudines, Crocodilia and Lepidosauria. We show the LDG of these groups has varied over time, with high latitudes serving as a source of tropical diversity but suffering disproportionate extinction during transitional periods to cold climate. We outline the "asymmetric gradient of extinction and dispersal" (AGED) framework, which contextualizes previous ideas behind the LDG under a time-variable scenario. We suggest the current steep LDG may be explained by the extinction of clades adapted to warmer conditions from the new temperate regions formed in the Neogene, together with the equator-ward dispersal of organisms tracking their own climatic preferences, when tropical biomes became restricted to the equator. Conversely, high rates of speciation and pole-ward dispersals can account for the formation of an ancient flat LDG during the Cretaceous-Paleogene greenhouse period. Our results demonstrate that the inclusion of fossils in macroevolutionary studies allows detecting extinction events less detectable in analyses restricted to present-day data only.

Building the monocot tree of death: Progress and challenges emerging from the macrofossil‐rich Zingiberales

Inclusion of fossils in phylogenetic analyses is necessary in order to construct a comprehensive "tree of death" and elucidate evolutionary history of taxa; however, such incorporation of fossils in phylogenetic reconstruction is dependent on the availability and interpretation of extensive morphological data. Here, the Zingiberales, whose familial relationships have been difficult to resolve with high support, are used as a case study to illustrate the importance of including fossil taxa in systematic studies. Eight fossil taxa and 43 extant Zingiberales were coded for 39 morphological seed characters, and these data were concatenated with previously published molecular sequence data for analysis in the program MrBayes. Ensete oregonense is confirmed to be part of Musaceae, and the other seven fossils group with Zingiberaceae. There is strong support for Spirematospermum friedrichii, Spirematospermum sp. 'Goth', S. wetzleri, and Striatornata sanantoniensis in crown Zingiberaceae while "Musa" cardiosperma, Spirematospermum chandlerae, and Tricostatocarpon silvapinedae are best considered stem Zingiberaceae. Inclusion of fossils explains how different topologies from morphological and molecular data sets is due to shared plesiomorphic characters shared by Musaceae, Zingiberaceae, and Costaceae, and most of the fossils. Inclusion of eight fossil taxa expands the Zingiberales tree and helps explain the difficulty in resolving relationships. Inclusion of fossils was possible in part due to a large morphological data set built using nondestructive microcomputed tomography data. Collaboration between paleo- and neobotanists and technology such as microcomputed tomography will help to build the tree of death and ultimately improve our understanding of the evolutionary history of monocots.

Divergence times in demosponges (Porifera): first insights from new mitogenomes and the inclusion of fossils in a birth-death clock model

BackgroundApproximately 80% of all described extant sponge species belong to the class Demospongiae. Yet, despite their diversity and importance, accurate divergence times are still unknown for most demosponge clades. The estimation of demosponge divergence time is key to answering fundamental questions on the origin of Demospongiae, their diversification and historical biogeography. Molecular sequence data alone is not informative on an absolute time scale, and therefore needs to be “calibrated” with additional data such as fossils. Here, we calibrate the molecular data with the fossilized birth-death model, which compared to strict node dating, allows for the inclusion of young and old fossils in the analysis of divergence time. We use desma-bearing sponges, a diverse group of demosponges that form rigid skeletons and have a rich and continuous fossil record dating back to the Cambrian (~500 Ma), to date the demosponge radiation and constrain the timing of key evolutionary events, like the transition from marine to freshwater habitats. To infer a dated phylogeny of Demospongiae we assembled the mitochondrial genomes of six desma-bearing demosponges from reduced-representation genomic libraries. The total dataset included 33 complete demosponge mitochondrial genomes and 30 fossils.ResultsOur study supports a Neoproterozoic origin of Demospongiae. Novel age estimates for the split of freshwater and marine sponges dating back to the Carboniferous and the previously assumed recent (~18 Ma) diversification of freshwater sponges is supported. Moreover, we provide detailed age estimates for a possible diversification of Tetractinellidae (~315 Ma), the Astrophorina (~240 Ma), the Spirophorina (~120 Ma) and the family Corallistidae (~188 Ma) all of which are considered as key groups for dating the Demospongiae due to their extraordinary rich and continuous fossil history.ConclusionThis study provides novel insights into the evolution of Demospongiae. Observed discrepancies of our dated phylogeny with their putative first fossil appearance dates are discussed for selected sponge groups. For instance, a Carboniferous origin of the order Tetractinellida seems to be too late, compared to their first appearance in the fossil record in the Middle Cambrian. This would imply that Paleozoic spicule forms are not homologous to post-Paleozoic forms.

Fossils and living taxa agree on patterns of body mass evolution: a case study with Afrotheria.

Most of life is extinct, so incorporating some fossil evidence into analyses of macroevolution is typically seen as necessary to understand the diversification of life and patterns of morphological evolution. Here we test the effects of inclusion of fossils in a study of the body size evolution of afrotherian mammals, a clade that includes the elephants, sea cows and elephant shrews. We find that the inclusion of fossil tips has little impact on analyses of body mass evolution; from a small ancestral size (approx. 100 g), there is a shift in rate and an increase in mass leading to the larger-bodied Paenungulata and Tubulidentata, regardless of whether fossils are included or excluded from analyses. For Afrotheria, the inclusion of fossils and morphological character data affect phylogenetic topology, but these differences have little impact upon patterns of body mass evolution and these body mass evolutionary patterns are consistent with the fossil record. The largest differences between our analyses result from the evolutionary model, not the addition of fossils. For some clades, extant-only analyses may be reliable to reconstruct body mass evolution, but the addition of fossils and careful model selection is likely to increase confidence and accuracy of reconstructed macroevolutionary patterns.

Phylogeny, divergence times and biogeography of window flies (Scenopinidae) and the therevoid clade (Diptera: Asiloidea)

AbstractThe evolution of the ‘therevoid’ clade, with an emphasis on window flies (Scenopinidae), is presented by combining DNA sequence data with morphological characters for living and fossil species. The therevoid clade represents a group of four families (Apsilocephalidae, Evocoidae, Scenopinidae and Therevidae) of lower brachyceran Diptera in the superfamily Asiloidea. A comprehensive phylogenetic analysis using parsimony and likelihood methods was undertaken using extensive taxon sampling from all families and subfamilies, and compared with outgroup taxa sampled from the related families Asilidae, Mydidae, Apioceridae and Empididae. Fifty‐nine morphological characters (adult, larval and pupal) were combined with 6.4 kb of DNA sequences for two ribosomal genes (16S and 18S ribosomal DNA) and three protein‐encoding genes [cytochrome oxidase I (COI), triose phosphate isomerase (TPI) and the CPSase region of carbamoyl‐phosphate synthase‐aspartate transcarbamoylase‐dihydroorotase (CAD)]. Results from combined analyses of morphological and molecular data for 78 taxa representing all families of the therevoid clade are presented. Specific hypotheses of the relationship between respective families and subfamilies were tested statistically using four‐cluster likelihood mapping. The therevoid clade is a well‐supported monophyletic group within Asiloidea, with Evocoidae sister to Apsilocephalidae and Therevidae sister to Scenopinidae. Temporal and zoogeographical aspects of therevoid clade evolution were investigated using Bayesian divergence time estimates and Lagrange ancestral range scenarios. The effect of inclusion of fossils as terminal taxa on phylogenetic and divergence time estimation was investigated, with morphological scoring for fossil representatives included in the analyses rather than used simply as minimum age constraints. In each analysis there was either improvement in estimation, or only marginal and localized loss in tree resolution, and with younger estimates of divergence time across the tree. The historical biogeography of the therevoid clade was examined with multiple trans‐Antarctic vicariance events between Australasia and South America evident during the Late Cretaceous to early Palaeogene. Scenopininae is newly subdivided into two tribes, Metatrichini trib.n. and Scenopinini Fallén stat.r.This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4974EBF8‐3117‐4189‐B6DE‐7D5BF9B23E53.

Feeding habits of extant and fossil canids as determined by their skull geometry

AbstractThe canids belong to one of the most prominent families of mammalian carnivores. Feeding adaptations of extant species is well documented by field observations; however, we are still missing palaeoecological insights for many enigmatic fossil specimens. We employ geometric morphometrics to quantify skull size and shape in extant and fossil members of the Canini tribe, inclusive of jackals and wolf‐like taxa. Skull data are tested to identify correlates of dietary adaptations in extant species for predicting adaptations in fossils. Main vectors of shape variation correlate with the relative skull‐palatal length, the position of the upper carnassial tooth and the anterior tip of the secondary palate. Allometry occurs in the palatal shape but size explains only a small fraction (about 4%) of shape variance. Although we quantified only palatal and tooth shape for the inclusion of fragmentary fossils, discriminant function analysis successfully classify extant Canini in dietary groups (small, medium and large prey specialist) with 89% of accuracy. The discriminant functions provide insights into many enigmatic specimens such as Eucyon adoxus (=small prey), fossil jackal‐like from Koobi Fora formation (=small prey) and the Plio‐Pleistocene Old World canid guild (Canis etruscus, C. arnensis and Lycaon falconeri). Clearly, both skull size and shape are excellent predictors of feeding habits in Canini thus also provide information about fossil taxonomic affinities.

Early fossils illuminate character evolution and interrelationships of Lampridiformes (Teleostei, Acanthomorpha)

Lampridiformes is a peculiar clade of pelagic marine acanthomorph (spiny-rayed) teleosts. Its phylogenetic position remains ambiguous, and varies depending on the type of data (morphological or molecular) used to infer interrelationships. Because the extreme morphological specializations of lampridiforms may have overwritten the ancestral features of the group with a bearing on its relationships, the inclusion of fossils that exhibit primitive character state combinations for the group as a whole is vital in establishing its phylogenetic position. Therefore, we present an osteological data set of extant (ten taxa) and fossil (14 taxa) acanthomorphs, including early Late Cretaceous taxa for which a close relationship with extant Lampridiformes has been suggested: †Aipichthyoidea, †Pharmacichthyidae, and †Pycnosteroididae. We find that all three taxa plus Lampridiformes form a clade that we call Lampridomorpha. Under this hypothesis, †Aipichthyoidea is paraphyletic. The inclusion of fossils in the analysis changes the topology, highlighting their critical importance in phylogenetic studies of morphological characters. When fossils are included, Lampridomorpha is sister to Euacanthomorpha (all other extant acanthomorphs), concurring with most previous anatomical studies, but conflicting with most molecular results. Lampridomorpha as a whole was a major component of the earliest acanthomorph faunas, notably in the Cenomanian. © 2014 The Linnean Society of London

Reweaving the tapestry: a supertree of birds.

Our knowledge of the avian tree of life remains uncertain, particularly at deeper levels due to the rapid diversification early in their evolutionary history. They are the most abundant land vertebrate on the planet and have been of great historical interest to systematists. Birds are also economically and ecologically important and as a result are intensively studied, yet despite their importance and interest to humans around 13% of taxa currently on the endangered species list perhaps as a result of human activity. Despite all this no comprehensive phylogeny that includes both extinct and extant species currently exists. Here we present a species-level supertree, constructed using the Matrix Representation with Parsimony method, of Aves containing approximately two thirds of all species from nearly 1000 source phylogenies with a broad taxonomic coverage. The source data for the tree were collected and processed according to a strict protocol to ensure robust and accurate data handling. The resulting tree topology is largely consistent with molecular hypotheses of avian phylogeny. We identify areas that are in broad agreement with current views on avian systematics and also those that require further work. We also highlight the need for leaf-based support measures to enable the identification of rogue taxa in supertrees. This is a first attempt at a supertree of both extinct and extant birds, it is not intended to be utilised in an overhaul of avian systematics or as a basis for taxonomic re-classification but provides a strong basis on which to base further studies on macroevolution, conservation, biodiversity, comparative biology and character evolution, in particular the inclusion of fossils will allow the study of bird evolution and diversification throughout deep time.

Combining fossil and molecular data to date the diversification of New World Primates

Recent methodological advances in molecular dating associated with the growing availability of sequence data have prompted the study of the evolution of New World Anthropoidea in recent years. Motivated by questions regarding historical biogeography or the mode of evolution, these works aimed to obtain a clearer scenario of Platyrrhini origins and diversification. Although some consensus was found, disputed issues, especially those relating to the evolutionary affinities of fossil taxa, remain. The use of fossil taxa for divergence time analysis is traditionally restricted to the provision of calibration priors. However, new analytical approaches have been developed that incorporate fossils as terminals and, thus, directly assign ages to the fossil tips. In this study, we conducted a combined analysis of molecular and morphological data, including fossils, to derive the timescale of New World anthropoids. Differently from previous studies that conducted total-evidence analysis of molecules and morphology, our approach investigated the morphological clock alone. Our results corroborate the hypothesis that living platyrrhines diversified in the last 20Ma and that Miocene Patagonian fossils compose an independent evolutionary radiation that diversified in the late Oligocene. When compared to the node ages inferred from the molecular timescale, the inclusion of fossils augmented the precision of the estimates for nodes constrained by the fossil tips. We show that morphological data can be analysed using the same methodological framework applied in relaxed molecular clock studies.

Convergence and divergence in the evolution of cat skulls: temporal and spatial patterns of morphological diversity.

BackgroundStudies of biological shape evolution are greatly enhanced when framed in a phylogenetic perspective. Inclusion of fossils amplifies the scope of macroevolutionary research, offers a deep-time perspective on tempo and mode of radiations, and elucidates life-trait changes. We explore the evolution of skull shape in felids (cats) through morphometric analyses of linear variables, phylogenetic comparative methods, and a new cladistic study of saber-toothed cats.Methodology/Principal FindingsA new phylogenetic analysis supports the monophyly of saber-toothed cats (Machairodontinae) exclusive of Felinae and some basal felids, but does not support the monophyly of various saber-toothed tribes and genera. We quantified skull shape variation in 34 extant and 18 extinct species using size-adjusted linear variables. These distinguish taxonomic group membership with high accuracy. Patterns of morphospace occupation are consistent with previous analyses, for example, in showing a size gradient along the primary axis of shape variation and a separation between large and small-medium cats. By combining the new phylogeny with a molecular tree of extant Felinae, we built a chronophylomorphospace (a phylogeny superimposed onto a two-dimensional morphospace through time). The evolutionary history of cats was characterized by two major episodes of morphological divergence, one marking the separation between saber-toothed and modern cats, the other marking the split between large and small-medium cats.Conclusions/SignificanceAncestors of large cats in the ‘Panthera’ lineage tend to occupy, at a much later stage, morphospace regions previously occupied by saber-toothed cats. The latter radiated out into new morphospace regions peripheral to those of extant large cats. The separation between large and small-medium cats was marked by considerable morphologically divergent trajectories early in feline evolution. A chronophylomorphospace has wider applications in reconstructing temporal transitions across two-dimensional trait spaces, can be used in ecophenotypical and functional diversity studies, and may reveal novel patterns of morphospace occupation.

Phylogenetic signal and functional significance of incisor enamel microstructure in Arvicola (Rodentia, Arvicolinae)

Many genera of arvicolid rodents (voles) contain species that spend a considerable amount of time underground. Incisors provide a considerable effort in the digging process and a difference in the enamel microstructure of incisors is expected between species that spend most of their life above ground and those who have greater specializations for a subterranean habitat. The ratio between the thicknesses of the two layers composing the enamel of the incisors has proved to be an effective proxy to infer fossorial activity of extinct forms of Arvicola but this ratio exhibited no phylogenetic signal (although lifestyle was in a greater extent related to phylogeny). We were able to infer ancestral lifestyles (of extinct populations and of nodes of the phylogeny) from enamel morphologies. Inclusion of fossils did not solve the uncertainties associated with the lifestyle for the last common ancestor of extant Arvicola species and it did not change the inferred lifestyle of several other, less inclusive, clades.

THE AGE OF THE ANGIOSPERMS: A MOLECULAR TIMESCALE WITHOUT A CLOCK

The age of the angiosperms has long been of interest to botanists and evolutionary biologists. Many early efforts to date the age of the angiosperms and evolutionary divergences within the angiosperm clade using a molecular clock have yielded age estimates that are grossly inconsistent with the fossil record. We investigated the age of angiosperms using Bayesian relaxed clock (BRC) and penalized likelihood (PL) approaches. Both of these methods allow the incorporation of multiple fossil constraints into the optimization procedure. The BRC method allows a range of values for among-lineage rate of substitution, from a nearly clocklike behavior to a condition in which each branch is allowed an optimal substitution rate, and also accounts for variation in molecular evolution across multiple genes. A topology derived from an analysis of genes from all three plant genomes for 71 taxa was used as a backbone. The effects on age estimates of different genes, single-gene versus concatenated datasets, and the inclusion and assumptions of fossils as age constraints were examined. In addition, the influence of prior distributions on estimates of divergence times was also explored. These results indicate that widely divergent age estimates can result from the different methods (198-139 million years ago), different sources of data (275-122 million years ago), and the inclusion of temporal constraints to topologies. Most dates, however, are between 180-140 million years ago, suggesting a Middle Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil angiosperms by about 45-5 million years. Nonetheless, these dates are consistent with other recent studies that have used methods that relax the assumption of a strict molecular clock and also agree with the hypothesis that the angiosperms may be somewhat older than the fossil record indicates.

THE AGE OF THE ANGIOSPERMS: A MOLECULAR TIMESCALE WITHOUT A CLOCK

The age of the angiosperms has long been of interest to botanists and evolutionary biologists. Many early efforts to date the age of the angiosperms and evolutionary divergences within the angiosperm clade using a molecular clock have yielded age estimates that are grossly inconsistent with the fossil record. We investigated the age of angiosperms using Bayesian relaxed clock (BRC) and penalized likelihood (PL) approaches. Both of these methods allow the incorporation of multiple fossil constraints into the optimization procedure. The BRC method allows a range of values for among-lineage rate of substitution, from a nearly clocklike behavior to a condition in which each branch is allowed an optimal substitution rate, and also accounts for variation in molecular evolution across multiple genes. A topology derived from an analysis of genes from all three plant genomes for 71 taxa was used as a backbone. The effects on age estimates of different genes, single-gene versus concatenated datasets, and the inclusion and assumptions of fossils as age constraints were examined. In addition, the influence of prior distributions on estimates of divergence times was also explored. These results indicate that widely divergent age estimates can result from the different methods (198–139 million years ago), different sources of data (275–122 million years ago), and the inclusion of temporal constraints to topologies. Most dates, however, are between 180–140 million years ago, suggesting a Middle Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil angiosperms by about 45–5 million years. Nonetheless, these dates are consistent with other recent studies that have used methods that relax the assumption of a strict molecular clock and also agree with the hypothesis that the angiosperms may be somewhat older than the fossil record indicates.

The Importance of Even Highly Incomplete Fossil Taxa in Reconstructing the Phylogenetic Relationships of the Tetraodontiformes (Acanthomorpha: Pisces)

The use of fossils in the phylogenetics of extant clades traditionally has been a contentious issue. Fossils usually are relatively incomplete, and their use commonly leads to an increase in the number of equally most parsimonious trees and a decrease in the resolution of phylogenies. Fossils alone, however, provide certain kinds of information about the biological history of a clade, and computer simulations have shown that even highly incomplete material can, under certain circumstances, increase the accuracy of a phylogeny, rather than decrease it.Because empirical data are still scarce on the effects of the inclusion of fossils on phylogenetic reconstructions, we attempted to investigate this problem by using a relatively well-known group of acanthomorph fishes, the Tetraodontiformes (triggerfishes, pufferfishes, and ocean sunfishes), for which robust phylogenies using extant taxa already exist and that has a well-studied fossil record. Adding incomplete fossil taxa of tetraodontiforms usually increases the number of equally most parsimonious trees and often decreases the resolution of consensus trees. However, adding fossil taxa may help to correctly establish relationships among lineages that have experienced high degrees of morphological diversification by allowing for a reinterpretation of homologous and homoplastic features, increasing the resolution rather than decreasing it. Furthermore, taxa that were scored for 25% or more of their characters did not cause a significant loss of resolution, while providing unique biological information.

Taxonomy and fossils: a critical appraisal.

Many compendia at the species, genus and family levels document the fossil record, but these are not standardized, nor usually critical in content, and few are available on the World Wide Web. The sampling of the available record is good for organisms with fossilizable parts, but preservational constraints on the entire morphology, life history and geographical distribution lead to difficulties in recognizing and naming species. We recommend abandoning some of the palaeontological species concepts such as chronospecies and stratospecies, and we advocate species recognition based on unique combinations of characters. The compilation of species lists is extremely time consuming, and given the inherent problems we suggest that compilation of generic lists is a more achievable goal because genera are recognized by definitive morphological characters. In calculating taxon duration, care must be taken to distinguish between mono-, para- and polyphyletic groups, the first being the only reliable unit for use in calculating diversity curves. We support the inclusion of fossils into classifications based on Recent organisms, but we recognize some of the problems this may pose for standard Linnaean classifications. Web-based taxonomy is the way forward, having the advantages of speed and currency of information dissemination, universal access with links to primary literature and increasingly sophisticated imagery. These advantages over conventional outlets will only be realized with careful Web design and a commitment to maintenance.

Relationships of Endemic African Mammals and Their Fossil Relatives Based on Morphological and Molecular Evidence

Analyses of anatomical and DNA sequence data run on a parallel supercomputer that include fossil taxa support the inclusion of tenrecs and golden moles in the Afrotheria, an endemic African clade of placental mammals. According to weighting schemes of morphological and molecular data that maximize congruence, extinct members of the afrotherian crown group include embrithopods, Plesiorycteropus, desmostylians, and the “condylarths” Hyopsodus, Meniscotherium, and possibly Phenacodus. By influencing the optimization of anatomical characters, molecular data have a large influence on the relationships of several extinct taxa. The inclusion of fossils and morphological data increases support for an elephant-sea cow clade within Paenungulata and identifies ancient, northern elements of a clade whose living members in contrast suggest an historically Gondwanan distribution. In addition, maximally congruent topologies support the position of Afrotheria as well-nested, not basal, within Placentalia. This pattern does not accord with the recent hypothesis that the divergence of placental mammals co-occurred with the tectonic separation of Africa and South America.