The historical biogeography of sturgeons (Osteichthyes: Acipenseridae): a synthesis of phylogenetics, palaeontology and palaeogeography

Abstract. The historical biogeography of sturgeons is explored using information from palaeogeography, palaeontology and phylogenetic interrelationships. The integration of information from these diverse sources indicates that sturgeons reached a wide Laurasian distribution in the Cretaceous and Tertiary by freshwater and coastal dispersal routes across land connections and along newly forming continental margins. The fossil record also suggests a considerable degree of morphological stasis and also supports an estuarine habit, and perhaps diadromy, as an old and conserved life history trait. While a ‘centre of origin’ for sturgeons remains elusive, phylogenetic relationships indicate that diversification appears to have been associated with fragmentation of biota, and of landmasses and basins, by late Tertiary geological and climatic phenomena, such as orogeny and unequal glaciation over North America, the desiccation of central Asia and alteration of its drainages, and the formation of discrete Ponto-Caspian basins by the fragmentation of the Paratethys. Amphi-oceanic distributions of certain species (Acipenser medirostris Ayres) and sister taxa (e.g. A. oxyrhynchus Mitchill and A. sturio L.) are explained by coastal dispersal and subsequent vicariance by geological (sea-floor spreading and development of new continental margins) and climatic (Pliocene cooling) changes during the Tertiary. An hypothesis is developed for the relationships of the North American sturgeons and their potential relationships with the Siberian sturgeon A. baeri. Late Tertiary climatic and geological phenomena are hypothesized as mediators of vicariance and subsequent diversification of these acipenserids. It appears that although acipenserids are a geologically old group, the historical biogeography of surviving lineages is best explained by more recent geological and climatic changes.     

Evolutionary morphology of whip spiders: towards a phylogenetic system (Chelicerata: Arachnida: Amblypygi)*

The aim of this study is to present a cladogram and phylogenetic system and to use this to discuss the phylogeny and biogeography of the Amblypygi. A total of 29 morphological structures were studied, their plesiomorphic and apomorphic characters or character states were identified, and the resulting data matrix was analysed. As a result, the ‘old’Charontidae or Pulvillata emerge as a paraphyletic group; the genus Paracharon is the sister group of all other amblypygids, which are now termed Euamblypygi. The ‘new’Charontidae (sensu Quintero: the genera Stygophrynus and Charon) are the sister group of the Phrynida or Apulvillata; together they form the Neoamblypygi. The relationships of the genera of the Charinidae cannot be resolved with the available data. They may be a paraphyletic group. The genus Catageus is a possible candidate for being the sister group of the Neoamblypygi. The new system allows a discussion of the phylogeny and biogeography of whip spiders. It also points to unresolved taxa and thus indicates the questions future research should address.     

Polytomies,signal and noise: revisiting the mitochondrial phylogeny and phylogeography of the Eurasian blindsnake species complex (Typhlopidae,Squamata)

Polytomies are multifurcating nodes on a phylogenetic tree that represent unresolved relationships. Contrary to ‘hard’ polytomies that hide simultaneous splitting events, ‘soft’ polytomies can theoretically be resolved with the addition of phylogenetic signal. This is not always successful, especially when a radiation is old and rapid, because adding more signal will inevitably increase the noise. The Xerotyphlops vermicularis species complex is an example of a largely unresolved old and rapid radiation. This mtDNA phylogeny is revisited by analysing samples representative of all lineages and generating a 3700‐bp final data set, after preliminary tests found two of the analysed markers responsible for long‐branch attraction. The new enhanced data set increased phylogenetic resolution, resulting in a robust time‐calibrated phylogeny and phylogeographic conclusions. Jordan/south Syria populations separated during the Middle Miocene, and the Messinian salinity crisis (MSC) appears responsible for the second diversification wave. A third radiation occurred during the Early Pliocene, resulting in four mtDNA groups west, east and south of the Amanos mountains in east Turkey/north Syria. Finally, three hypotheses regarding the number of potential ‘species’ within the complex are made, with the most moderate ‘four species’ hypothesis fitting very well with the diversity patterns and established systematics of east Mediterranean reptiles.     

THE HETEROMORPHS AND AMMONOID EXTINCTION

The heteromorph ammonoids are quoted as a favourite example of degeneration and the decline of a Bauplan‘condemned’ to extinction. With astonishing tenacity this view of the heteromorphs as ‘phylogenetic end-forms’ has embedded itself in the palaeontological literature and is still current. This is contradicted by the most recent investigations, directed especially at the Cretaceous heteromorphs, which necessitate correction of the typolysis concept as well as modification of the most uncontested of the phylogenetic ‘laws’, Dollo's ‘law of irreversibility’. Contrary to the usual textbook hypothesis, the heteromorphs return in several evolutionary lineages to normal coiling of the shell and, in general, to a phylogenetically older type of suture line. At the same time these results encourage fresh reflexion on possible exogenous causes of phylogenetic extinction of the ammonoids. A clear causal connexion exists between this extinction and the far-reaching epirogenic changes in sea level in the late Cretaceous; cosmic explanations are unnecessary. In conclusion it may be added that the precipitate formulation of phylogenetic ‘laws’ and ‘principles’ based on too little basic information has encumbered this branch of palaeontology with a stifling set of prejudices rather than providing it with guide lines crystallized from long experience and observation. It is vitally necessary in the interests of palaeontology that interpretation and observation be separated far more than has been the case in the past.     

Exploring the phylogenetic history of mammal species richness

Aim At broad geographical scales, species richness is a product of three basic processes: speciation, extinction and migration. However, determining which of these processes predominates is a major challenge. Whilst palaeontological studies can provide information on speciation and extinction rates, data are frequently lacking. Here we use a recent dated phylogenetic tree of mammals to explore the relative importance of these three processes in structuring present‐day richness gradients. Location The global terrestrial biosphere. Methods We combine macroecological data with phylogenetic methods more typically used in community ecology to describe the phylogenetic history of regional faunas. Using simulations, we explore two simple phylogenetic metrics, the mean and variance in the pairwise distances between taxa, and describe their relationship to phylogenetic tree topology. We then use these two metrics to characterize the evolutionary relationships among mammal species assemblages across the terrestrial biome. Results We show that the mean and variance in the pairwise distances describe phylogenetic tree topology well, but are less sensitive to phylogenetic uncertainty than more direct measures of tree shape. We find the phylogeny for South American mammals is imbalanced and ‘stemmy’ (long branches towards the root), consistent with recent diversification within evolutionarily disparate lineages. In contrast, the phylogeny for African mammals is balanced and ‘tippy’ (long branches towards the tips), more consistent with the slow accumulation of diversity over long times, reflecting the Old World origin of many mammal clades. Main conclusions We show that phylogeny can accurately capture biogeographical processes operating at broad spatial scales and over long time periods. Our results support inferences from the fossil record – that the New World tropics are a diversity cradle whereas the Old World tropics are a museum of old diversity.     

‘Representative Genes’, is it OK to use a small amount of data to obtain a phylogeny that is at least close to the true tree?

Despite the problems associated with the one‐gene‐constructed phylogenetic relationships, recently it has been shown that merely adding more sequences to the analyses is not enough to resolve all the inconsistencies present in these relationships. In this work, the existence of ‘representative’ genes for groups of species is evaluated in terms of efficiency (costs/benefits), employing the cytochrome c oxidase subunit I (COI) and the penguins' phylogeny as a case study. ‘Representative’ genes are very useful and they could produce good results for first approaches or humble laboratories, especially if the gene is chosen depending on the time scale and objectives of our work.     

The geography of speciation in narrow‐range endemics of the ‘Haenydra’ lineage (Coleoptera,Hydraenidae, Hydraena)

Aim We test whether species of western Mediterranean aquatic Coleoptera of the ‘Haenydra’ lineage (Hydraenidae, Hydraena) originated through: (1) successive periods of dispersal and speciation, (2) range fragmentation by random vicariance, or (3) range fragmentation by geographic isolation owing to a general reduction of population density. Location Europe. Methods To discriminate between scenarios we use contrasting predictions of the relationship between phylogenetic and geographic distance. The phylogeny was based on 3 kb of four mitochondrial and two nuclear gene fragments of about half of the known species of ‘Haenydra’, including most western Mediterranean taxa. Divergences were estimated using a molecular clock. The relationship between phylogenetic and geographic distance was tested using bivariate plots, Mantel tests and comparison of the observed phylogeny with the one minimizing geographic distances between species, as measured using Euclidean minimum spanning trees (EMSTs). Results The monophyly of ‘Haenydra’ was strongly supported, although its phylogenetic placement was not resolved. ‘Haenydra’ was estimated to be of late Miocene age, with most species originating during the Pleistocene. In two clades (Hydraena tatii and Hydraena emarginata clades) there was a significant association between geographic and phylogenetic distance, and the reconstructed phylogeny was identical to that obtained through the EMST, demonstrating a strong non‐randomness of the geographic distribution of the species. In two other clades (Hydraena iberica and Hydraena bitruncata clades) there was no association between geographic and phylogenetic distance, and the observed phylogeny was not the one minimizing geographic distances. In one of the clades this seems to be due to a secondary, recent range expansion of one species (H. iberica), which erased the geographic signal of their distributions. Main conclusions We show that it is possible to obtain strong evidence of stasis of the geographic ranges of narrow‐range endemic species through the study of their phylogenetic relationships and current distributions. In at least two of the studied clades, current species seem to have originated through the fragmentation of a more widely distributed species, without further range movements. A process of range expansion and fragmentation may have occurred repeatedly within the ‘Haenydra’ lineage, contributing to the accumulation of narrow‐range endemics in Mediterranean Pleistocene refugia.     

Phylogenetic interrelationships of the barbets (Aves: Capitonidae) and toucans (Aves: Ramphastidae) based on morphology with comparisons to DNA-DNA hybridization

A phylogenetic analysis of the interrelationships of the barbets (Capitonidae) and the toucans (Aves: Ramphastidae, Superfamily Ramphastoidea) is presented. Thirty-two morphological characters from the literature and independent osteological observations were analysed. Character polarity was determined by outgroup comparison to the Picidae, Indicatoridae, Galbulidae, Bucconidae and Coraciiformes. Four alternative phylogenetic hypotheses were compared: (1) the overall most parsimonious morphological phylogeny, (2) the most parsimonious morphological phylogeny in which the capitonids and ramphastids were hypothesized as monophyletic sister groups, and (3) and (4) the most parsimonious hypotheses for the evolution of the morphological characters within two proposed DNA-DNA hybridization phylogenies of the ramphastoids. The analysis focused on the higher level relationships of ramphastids and capitonids and interrelationships among capitonid genera. Two cladistic analyses were performed using 26 phylogenetically informative characters, and the PAUP and CONTREE computer alogorithms. The most parsimonious morphological phylogeny required fewer character changes and had a lower consistency index than any of the alternative hypotheses but congruence between the most parsimonious phylogeny and the second, revised DNA-DNA hybridization hypothesis was very high. Based on these results the monophyly of the Capitonidae is rejected. The ramphastids and the Neotropical capitonids form a well corroborated clade within the pantropical ramphastoid radiation. Neither the African, Asian nor New World capitonids is monophyletic. The genus Trachyphonus is the sister group to all other capitonids and ramphastids. The sister group to the ramphastids is the genus Semnornis. The interrelationships of the Old World capitonids excluding Trachyphonus are not completely resolved by these morphological data but one of the alternative phylogenetic resolutions is presented as a preliminary hypothesis. The clades in this resolved phylogeny are diagnosed and the palaeontology and biogeography of the ramphastoids arc-reviewed in light of this new evidence. A phylogenetic classification is proposed in which the Capitonidae is rejected and the capitonids and ramphastids are placed in seven subfamilies of the Ramphastidae.     

A mitochondrial phylogeny of the rainforest skink genus Saproscincus, Wells and Wellington (1984)

The phylogenetic relationships and historical biogeography of 10 currently described rainforest skinks in the genus Saproscincus were investigated using mitochondrial protein-coding ND4 and ribosomal RNA 16S genes. A robust phylogeny is inferred using both maximum likelihood and Bayesian analysis, with all inter-specific nodes strongly supported when datasets are combined. The phylogeny supports the recognition of two major lineages (northern and southern), each of which comprises two divergent clades. Both northern and southern lineages have comparably divergent representatives in mid-east Queensland (MEQ), providing further molecular evidence for the importance of two major biogeographic breaks, the St. Lawrence gap and Burdekin gap separating MEQ from southern and northern counterparts respectively. Vicariance associated with the fragmentation and contraction of temperate rainforest during the mid-late Miocene epoch underpins the deep divergence between morphologically conservative lineages in at least three instances. In contrast, one species, Saproscincus oriarus, shows very low sequence divergence but distinct morphological and ecological differentiation from its allopatric sister clade within Saproscincus mustelinus. These results suggest that while vicariance has played a prominent role in diversification and historical biogeography of Saproscincus, divergent selection may also be important.     

When phylogenetics met biogeography: Willi Hennig,Lars Brundin and the roots of phylogenetic and cladistic biogeography

Willi Hennig's (Beitr. Ent. 1960, 10, 15) Die Dipteren-Fauna von Neuseeland als systematisches und tiergeographisches Problem applied a phylogenetic approach to examine the distributional patterns exhibited by the Diptera of New Zealand. Hennig showed how phylogenetic trees may be used to infer dispersal, based on the progression and deviation rules, and also discussed the existence of vicariance patterns. The most important author who applied Hennig's phylogenetic biogeography was Lars Brundin, when analysing the phylogenetic relationships of two taxa of Chironomidae (Diptera) and using them to examine the biogeographic relationships of Australia, New Zealand, South America and South Africa. The relevance of Brundin's contribution was noted by several authors, as it began the cladistic or vicariance approach to biogeography, that implies the discovery of vicariance events shared by different monophyletic groups. Both phylogenetic and cladistic biogeography have a place in contemporary biogeography, the former for analysing taxon biogeography and the latter when addressing Earth or biota biogeography. The recent use of the term “phylogenetic biogeography” to refer to a posteriori methods of cladistic biogeography is erroneous and should be avoided.     

Integrating phylogeny,environment and space to explore variation in macroecological traits of Viperidae and Elapidae (Squamata: Serpentes)

We used eigenvector mapping in space and phylogeny to investigate the relationships among space, phylogeny and environment on body size and range size variation across two groups of venomous snakes – Viperidae and Elapidae – from the New World. Data on species geographic range sizes, maximum body sizes and phylogenetic relationships were compiled from the available literature. The distributional data were also used to calculate the latitudinal and longitudinal midpoint and the environmental centroids for each species. The eigenvectors extracted from the pair wise spatial and phylogenetic distance matrices were integrated with environmental variables into a method of variation partitioning where the variation in each trait was quantitatively attributed to ‘pure’ and/or shared effects of phylogeny, environment and space. Our results showed that variation in body size was predominantly determined by phylogeny in both groups of snakes. For Viperidae, we found that pure ‘effects’ of phylogeny were the strongest, indicating that most of the body size evolution that was phylogenetically determined in this group occurred independently of environment and geographical proximity. Regarding range sizes, pure phylogenetic influences were very low in both groups, whereas the largest single fraction of explained variation corresponded to overlapped influences of the three sets of predictors, especially for Elapidae. Along with this, we found evidence that niche conservatism is an important processes underlying variation in body size and range size in both groups of snakes.     

Molecular phylogenetics of the hummingbird genus Coeligena

Advances in the understanding of biological radiations along tropical mountains depend on the knowledge of phylogenetic relationships among species. Here we present a species-level molecular phylogeny based on a multilocus dataset for the Andean hummingbird genus Coeligena. We compare this phylogeny to previous hypotheses of evolutionary relationships and use it as a framework to understand patterns in the evolution of sexual dichromatism and in the biogeography of speciation within the Andes. Previous phylogenetic hypotheses based mostly on similarities in coloration conflicted with our molecular phylogeny, emphasizing the unreliability of color characters for phylogenetic inference. Two major clades, one monochromatic and the other dichromatic, were found in Coeligena. Closely related species were either allopatric or parapatric on opposite mountain slopes. No sister lineages replaced each other along an elevational gradient. Our results indicate the importance of geographic isolation for speciation in this group and the potential interaction between isolation and sexual selection to promote diversification.     

Phylogeny of the freshwater crabs of the Western Ghats (Brachyura,Gecarcinucidae)

The Western Ghats mountain range in India is a biodiversity hotspot for a variety of organisms including a large number of endemic freshwater crab species and genera of the family Gecarcinucidae. The phylogenetic relationships of these taxa, however, have remained poorly understood. Here, we present a phylogeny that includes 90% of peninsular Indian genera based on mitochondrial 16S rRNA and nuclear histone H3 gene sequences. The subfamily Gecarcinucinae was found to be paraphyletic with members of two other subfamilies, Liotelphusinae and Parathelphusinae, nesting within. We identify a well‐supported clade consisting of north Indian species and one clade comprising mostly south Indian species that inhabit the southern ‘sky islands’ of the Western Ghats. Relationships of early diverging genera, however, were resolved with low support. This study also includes newly sampled material from an isolated mountain plateau in the northern part of the Western Ghats, representing a new species of Gubernatoriana, which we describe here as Gubernatoriana basalticola sp. n. The new species is immediately distinguished from its congeners and the related genera Ghatiana and Inglethelphusa by its carapace and cheliped morphology, which are unique among Indian freshwater crabs. This study highlights the urgent need for continued faunistic studies to assess the true diversity of gecarcinucid crabs on the Indian subcontinent, to fully understand the basal phylogenetic relationships within the freshwater crab family Gecarcinucidae, and to evaluate the conservation threat status and biogeography of the montane freshwater crabs of the Western Ghats.     

Historical and Ecological Factors in the Evolution, Adaptive Radiation, and Biogeography of Freshwater Mollusks

SYNOPSIS. There are some 36 families that are wholly freshwateror with representative species in freshwater. There are virtuallyno phylogenetic analyses for these families. Zoogeographic analysesof freshwater molluscan faunas are hindered by a lack of significantsystematic studies of these faunas. Such studies are essentialif one hopes to develop hypotheses about phylogeny or biogeography. It is clear from a phylogenetic analysis of the Pomatiopsidaethat phylogenetic, vicariance, dispersal, and ecological factorsall have significant effects on the patterns of distributionof this family. At one stage in history vicariance may be adominant factor while during another stage of history, dispersalmay be a dominant factor. At every stage, ecological considerationsare necessary to understand the phenotypes seen and the spatialrelationships among taxa. In examining the distribution patternsof dominant freshwater families with regard to their biological,ecological, and overall phylogenetic relationships it is evidentthat ecology plays a major role along with dispersal and vicariance.Clearly a synthesis is needed in biogeographical studies thatincorporates vicariance dispersal, ecology, and geology-paleontology.     

Molecular phylogeny and biogeography of African diploid barbs, ‘Barbus’, and allies in Africa and Asia (Teleostei: Cypriniformes)

African diploid barbs (‘Barbus’, Clypeobarbus, Barboides, etc.) are a group of small cyprinids with a body size less than 20 cm and widely distributed in drainages across Africa. These species constitute a significant component of African freshwater fish fauna. This study is the first to focus on the molecular systematics and biogeography of African diploid barbs ‘Barbus’ and its African and Asian allies using both mitochondrial and nuclear genes. We test for monophyly of groups, determine interspecific relationships and estimate the time of divergence of 52 species of ‘Barbus’ and allies using two mitochondrial and four nuclear genes. Resulting trees demonstrate that ‘Barbus’ and allies (Systomus, Barboides, Clypeobarbus and African tetraploid barbs) form a strongly supported clade; however, ‘Barbus’ is not resolved as monophyletic. Divergence time analyses identify the separation between Systomus and ‘Barbus’ plus African allies may have occurred around 26 MYA. In addition to the phylogenetic results, these findings highlight the need for more thorough taxonomic and systematic studies on ‘Barbus’ and allies using morphological and additional molecular data and greater taxon sampling, including the type species of the genus Enteromius, ‘Barbus’ potamogalis.     

Panbiogeography: Its origin,metamorphosis and decline

From the viewpoint of 2007, one can trace the history of an interesting and contentious trend in biogeography and evolution that began with Croizat’s concept of panbiogeography in 1958. After a quiescent period of about 16 years, some young biologists in New York and in New Zealand read Croizat’s books and became enthusiastic supporters of his ideas. In New York, in the early 1970s, panbiogeography was combined with a part of Hennig’s phylogenetic method to create vicariance biogeography. In 1986, the name of the latter was changed to cladistic biogeography. In the meantime, Croizat’s followers in New Zealand sought to maintain panbiogeography in its original form without reference to phylogeny. This idea reached its peak of popularity in 1989–1990 and then began to fade. In comparison, cladistic biogeography became much more widespread, especially when its followers began publishing laudatory books and papers. Its decline became noticeable after the turn of the century as the dispersal counterrevolution began to have its effect. It served a useful purpose by engaging the interest of young biologists who otherwise may not have become aware of biogeography.     

Dispersal and diversification: macroevolutionary implications of the MacArthur–Wilson model, illustrated by Simulium (Inseliellum) Rubstov (Diptera: Simuliidae)

Aim Provide an empirical test of the ‘radiation zone’ hypothesis of the MacArthur–Wilson theory of island biogeography using the taxon‐pulse hypothesis of Erwin and Brooks Parsimony Analysis (BPA) on Simulium (Inseliellum) Rubstov. Location Micronesia, Cook Islands, Austral Islands, Society Islands, Marquesas Islands, Fiji and New Caledonia. Methods Primary and secondary BPA of the phylogeny of Inseliellum. Results Primary BPA showed that 15% of the taxon area cladogram contained area reticulations. Secondary BPA (invoking the area duplication convention) generated a clear sequence of dispersal for Inseliellum. The sequence follows a Micronesia – Cook Islands – Marquesas Islands – Society Islands dispersal, with a separate dispersal from the Cook Islands to the Austral Islands less than 1 Ma. A radiation in the island of Tahiti (Society Islands) produced numerous dispersals from Tahiti to other islands within the Society Islands system. Islands close to Tahiti (source island) have been colonized from Tahiti more often than islands far from Tahiti, but a higher proportion of those species colonizing distant islands have become distinct species. Main conclusions The dispersal sequence of Inseliellum exhibits both old to young island dispersal and young to old island dispersal. This is due to habitat availability on each island. Inseliellum is a model system in exemplifying the ‘radiation zone’ hypothesis of MacArthur and Wilson. As well, islands close to the source are colonized more often that those far from the source, but colonization of islands far away from the source results in a higher proportion of speciation events than for islands close to the source. The diversification of Inseliellum corresponds to a taxon‐pulse radiation, with a centre of diversification on Tahiti resulting from its large area and abundant freshwater habitats. This study illustrates the utility of BPA in identifying complex scenarios that can be used to test theories about the complementary roles of ecology and phylogeny in historical biogeography.     

Phylogeny of the water strider genus Gerris Fabricius (Heteroptera: Gerridae) based on COI mtDNA, EF-1α nuclear DNA and morphology

Phylogenetic relationships between water striders (Heteroptera: Gerridae) of genus Gerris Fabricius were examined using molecular and morphological characters. The molecular dataset was 820 bp DNA from the 3′ half of the mitochondrial gene encoding cytochrome oxidase subunit I and 515 bp DNA from the nuclear gene encoding elongation factor 1 alpha. The morphological dataset was a slightly modified version of a previously published dataset. Representatives from all eight recognized species groups of Gerris, as well as six species from three related genera, including Gigantometra gigas, Limnoporus esakii, L. rufoscutellatus, Aquarius najas, A. conformis and A. paludum, were included. Unweighted parsimony analyses of the COI sequences gave a topology with strong support for only those nodes that were already recognized as closely related based on morphological characters. Similar analyses of EF‐1α gave a cladogram with a topology quite different from that based on morphology and COI. Unweighted parsimony analyses of the ‘total evidence’ dataset largely supports the traditional view of Gerris phylogeny. Finally, the implications of the reconstructed phylogeny in relation to biogeography and ecological phylogenetics of Gerris is discussed.     

The lemur revolution starts now: The genomic coming of age for a non-model organism

Morris Goodman was a revolutionary. Together with a mere handful of like-minded scientists, Morris established himself as a leader in the molecular phylogenetic revolution of the 1960s. The effects of this revolution are most evident in this journal, which he founded in 1992. Happily for lemur biologists, one of Morris Goodman’s primary interests was in reconstructing the phylogeny of the primates, including the tooth-combed Lorisifomes of Africa and Asia, and the Lemuriformes of Madagascar (collectively referred to as the suborder Strepsirrhini). This paper traces the development of molecular phylogenetic and evolutionary genetic trends and methods over the 50-year expanse of Morris Goodman’s career, particularly as they apply to our understanding of lemuriform phylogeny, biogeography, and biology. Notably, this perspective reveals that the lemuriform genome is sufficiently rich in phylogenetic signal such that the very earliest molecular phylogenetic studies – many of which were conducted by Goodman himself – have been validated by contemporary studies that have exploited advanced computational methods applied to phylogenomic scale data; studies that were beyond imagining in the earliest days of phylogeny reconstruction. Nonetheless, the frontier still beckons. New technologies for gathering and analyzing genomic data will allow investigators to build upon what can now be considered a nearly-known phylogeny of the Lemuriformes in order to ask innovative questions about the evolutionary mechanisms that generate and maintain the extraordinary breadth and depth of biological diversity within this remarkable clade of primates.