Phylogenetic nomenclature is compatible with diverse philosophical perspectives

Pleijel, F. & Härlin, M. (2004). Phylogenetic nomenclature is compatible with diverse philosophical perspectives. — Zoologica Scripta , 33 , 587–591.     

Taxon names as paradigms: the structure of nomenclatural revolutions

In the present paper I argue that the two systems of phylogenetic nomenclature hitherto proposed represent, in a generalized sense, two different philosophies for how science develops and progresses. The phylogenetic system of definition initially proposed by de Queiroz and Gauthier [Syst. Zool. 39 (1990) 307], and later labeled PSD, is typically Popperian in the sense that science progresses toward truth by an accumulation of knowledge. Phylogenetic definitions of taxon names are assumed to adapt automatically to each new hypothesis of phylogeny, thereby reflecting better and better hypotheses. The phylogenetic system of reference proposed by Härlin [Zool. Scr. 27 (1998a) 381], on the other hand, is more Kuhnian, because it is built on the idea that successive hypotheses are incommensurable (and thus not cumulative) and that taxon names might be equalled with low‐level paradigms.     

A cladistic analysis of the cockroach wasps based on morphological data (Hymenoptera: Ampulicidae)

The Ampulicidae are one of the most basal groups within the apoid wasps, the paraphyletic assemblage of wasps previously known as Sphecidae. The morphology and taxonomy of the Ampulicidae are poorly studied, and there is no strict cladistic approach on their phylogeny yet. Here we assemble morphological characters of phylogenetic significance and submit them to parsimony analyses using modern cladistic methods. This produces a well-resolved and firmly supported phylogenetic hypothesis on the generic relationships within the group. Based on our phylogenetic results a revised classification is provided, subdividing the Ampulicidae into two monophyletic subfamilies, Ampulicinae ( Ampulex and Trirogma ) and Dolichurinae, the latter comprising two monophyletic tribes, Dolichurini ( Dolichurus and Paradolichurus ) and Aphelotomini, new tribe ( Aphelotoma and Riekefella ). Based on the resulting cladogram, the historical biogeography and the fossil record of Ampulicidae are briefly discussed.
© The Willi Hennig Society 2009.     

Osteology of Pseudochampsa ischigualastensis gen. et comb. nov. (Archosauriformes: Proterochampsidae) from the Early Late Triassic Ischigualasto Formation of Northwestern Argentina

Proterochampsids are crocodile-like, probably semi-aquatic, quadrupedal archosauriforms characterized by an elongated and dorsoventrally low skull. The group is endemic from the Middle-Late Triassic of South America. The most recently erected proterochampsid species is “Chanaresuchus ischigualastensis”, based on a single, fairly complete skeleton from the early Late Triassic Ischigualasto Formation of northwestern Argentina. We describe here in detail the non-braincase cranial and postcranial anatomy of this species and revisit its taxonomy and phylogenetic relationships. The phylogenetic analysis recovered ‘Chanaresuchus ischigualastensis’ as part of a trichotomy together with Gualosuchus reigi and Chanaresuchus bonapartei. Accordingly, “Chanaresuchus ischigualastensis” can be potentially more closely related to Gualosuchus reigi, or even Rhadinosuchus gracilis, than to Chanaresuchus bonapartei. In addition, after discussing previously claimed synapomorphies of Chanaresuchus, we could not find unambiguous support for the monophyly of the genus. As a result, we propose here the erection of the new genus Pseudochampsa for ‘Chanaresuchus ischigualastensis’, which results in the new combination Pseudochampsa ischigualastensis. The information provided here about the anatomy and taxonomy of Pseudochampsa ischiguaslastensis will be useful for future quantitative analyses focused on the biogeography and macroevolutionary history of proterochampsids.     

An updated megaphylogeny of plants,a tool for generating plant phylogenies and an analysis of phylogenetic community structure

Aims The aim of this article is 3-fold. First, we present an updated version of a published megaphylogeny of vascular plants that can be used in studies of plant ecology and biogeography. Second, we develop a tool that can be used by botanists and plant ecologists to generate phylogenetic hypotheses in three scenarios. Third, we use a set of regional assemblages of angiosperm trees in North America as a model system to evaluate the effect of differences in phylogenies generated using the three scenarios on the quantification of phylogenetic properties and the relationship between measures of phylogenetic properties and environment.Methods The taxonomy and nomenclature of plant species in the megaphylogeny were standardized according to The Plant List (version 1.1). A tool for generating phylogenies was created using the R language. The robustness of derived phylogenies was evaluated using correlation and regression analyses.Important findings An updated megaphylogeny of vascular plants (PhytoPhylo) and a tool for reconstructing phylogenies of seed plants (S.PhyloMaker) were generated. Our study shows that phylogenies generated by S.PhyloMaker using the PhytoPhylo megaphylogeny as a backbone are nearly as good as phylogeny resolved at the species level when using derived phylogenies to quantify phylogenetic properties (e.g. phylogenetic diversity and phylogenetic relatedness) of biological assemblages, and that S.PhyloMaker-generated phylogenies are robust for studies of community ecology and biogeography, particularly those seeking for patterns of phylogenetic properties along environmental gradients.     

Taxonomic names and phylogenetic trees

This paper addresses the issue of philosophy of names within the context of biological taxonomy, more specifically how names refer. By contrasting two philosophies of names, one that is based on the idea that names can be defined and one that they cannot be defined, I point out some advantages of the latter within phylogenetic systematics. Due to the changing nature of phylogenetic hypotheses, the former approach tends to rob taxonomy from its unique communicative value since a name that is defined refers to whatever fits the definition. This is particularly troublesome should the hypothesis of phylogenetic relationship change. I argue that, should we decide to accept a new phylogenetic hypothesis, it is also likely that our view of what to name may change. A system where names only refer acknowledge this, and accordingly leaves it open whether to keep a name (and accept the way it refers in the new hypothesis) or discard a name and introduce new names for the parts of the tree that we find scientifically interesting. One of the main differences between a phylogenetic system of definition (PSD) and a phylogenetic system of reference (PSR) is that the former is governed by laws of language while the latter by communicative needs of taxonomists. Thus, a PSR tends to give primacy to phylogenetic trees rather than phylogenetic definitions of names should our views of which phylogenetic hypothesis to accept change. © 1998 The Norwegian Academy of Sciences and Letters     

Species boundaries and global biogeography of the Alexandrium tamarense complex (Dinophyceae)1

Alexandrium catenella (Whedon et Kof.) Balech, A. tamarense (M. Lebour) Balech, and A. fundyense Balech comprise the A. tamarense complex, dinoflagellates responsible for paralytic shellfish poisoning worldwide. The relationships among these morphologically defined species are poorly understood, as are the reasons for increases in range and bloom occurrence observed over several decades. This study combines existing data with new ribosomal DNA sequences from strains originating from the six temperate continents to reconstruct the biogeography of the complex and explore the origins of new populations. The morphospecies are examined under the criteria of phylogenetic, biological, and morphological species concepts and do not to satisfy the requirements of any definition. It is recommended that use of the morphospecies appellations within this complex be discontinued as they imply erroneous relationships among morphological variants. Instead, five groups (probably cryptic species) are identified within the complex that are supported on the basis of large genetic distances, 100% bootstrap values, toxicity, and mating compatibility. Every isolate of three of the groups that has been tested is nontoxic, whereas every isolate of the remaining two groups is toxic. These phylogenetic groups were previously identified within the A. tamarense complex and given geographic designations that reflected the origins of known isolates. For at least two groups, the geographically based names are not indicative of the range occupied by members of each group. Therefore, we recommend a simple group‐numbering scheme for use until the taxonomy of this group is reevaluated and new species are proposed.     

A comparative analysis of island floras challenges taxonomy‐based biogeographical models of speciation

Speciation on islands, and particularly the divergence of species in situ, has long been debated. Here, we present one of the first, complete assessments of the geographic modes of speciation for the flora of a small oceanic island. Cocos Island (Costa Rica) is pristine; it is located 550 km off the Pacific coast of Central America. It harbors 189 native plant species, 33 of which are endemic. Using phylogenetic data from insular and mainland congeneric species, we show that all of the endemic species are derived from independent colonization events rather than in situ speciation. This is in sharp contrast to the results of a study carried out in a comparable system, Lord Howe Island (Australia), where as much as 8.2% of the plant species were the product of sympatric speciation. Differences in physiography and age between the islands may be responsible for the contrasting patterns of speciation observed. Importantly, comparing phylogenetic assessments of the modes of speciation with taxonomy‐based measures shows that widely used island biogeography approaches overestimate rates of in situ speciation.     

Phylogenetic relationships of horned lizards (Phrynosoma) based on nuclear and mitochondrial data: evidence for a misleading mitochondrial gene tree

It has proven remarkably difficult to obtain a well-resolved and strongly supported phylogeny for horned lizards (Phrynosoma) because of incongruence between morphological and mitochondrial DNA sequence data. We infer the phylogenetic relationships among all 17 extant Phrynosoma species using >5.1 kb of mtDNA (12S rRNA, 16S rRNA, ND1, ND2, ND4, Cyt b, and associated tRNA genes), and >2.2kb from three nuclear genes (RAG-1, BDNF, and GAPD) for most taxa. We conduct separate and combined phylogenetic analyses of these data using maximum parsimony, maximum likelihood, and Bayesian methods. The phylogenetic relationships inferred from the mtDNA data are congruent with previous mtDNA analyses based on fewer characters and provide strong support for most branches. However, we detected strong incongruence between the mtDNA and nuclear data using comparisons of branch support and Shimodaira-Hasegawa tests, with the (P. platyrhinos+P. goodei) clade identified as the primary source of this conflict. Our analysis of a P. mcalliixP. goodei hybrid suggests that this incongruence is caused by reticulation via introgressive hybridization. Our preferred phylogeny based on an analysis of the combined data (excluding the introgressed mtDNA data) provides a new framework for interpreting character evolution and biogeography within Phrynosoma. In the context of this improved phylogeny we propose a phylogenetic taxonomy highlighting four clades: (1) Tapaja, containing the viviparous short-horned lizards P. ditmarsi, P. hernandesi, P. douglasii, and P. orbiculare; (2) Anota, containing species with prominent cranial horns (P. solare, P. mcallii, and the P. coronatum group); (3) Doliosaurus, containing three species lacking antipredator blood-squirting (P. modestum, P. platyrhinos, and P. goodei); and (4) Brevicauda, containing two viviparous species with extremely short tails that lack blood-squirting (P. braconnieri and P. taurus).     

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.     

Heavily exploited but poorly known: systematics and biogeography of commercially harvested pythons (Python curtus group) in Southeast Asia

More than 100000 blood pythons (brongersmai) and short-tailed pythons (curtus and breitensteini) are taken from Borneo and Sumatra each year for the commercial leather trade. Traditionally, all have been treated as a single polytypic species (Python curtus) , with three subspecies differing in colour, size and geographic distribution. Analyses of DNA sequences and morphological data clarify the phylogenetic relationships, taxonomy and biogeography of this group. The lineage is monophyletic, and each of the three subspecies differs from the other two both morphologically and genetically. Given the morphological and genetic distinctiveness of each taxon, we here elevate the three subspecies to full species status. Python brongersmai is the most distinctive in terms of colour (of the three, only brongersmai has colour-morphs that are red or orange), size (it grows to 2.6 m, vs. approx. 2.0 m for the other taxa), and scalation (e.g. brongersmai has <166 ventral scales, vs. <166 in the other taxa and has two supralabials over each orbit, vs. one supralabial for the other two taxa). In terms of cytochrome b mitochondrial DNA sequence data, brongersmai is almost as distant genetically from the short-tailed pythons (8.9% divergence) as is the reticulated python (P. reticulatus: 10.3% divergence). The other two taxa (P. breitensteini from Kalimantan and P. curtus from western and southern Sumatra) are closely related (3% divergence), despite their disjunct distribution (separated by P. brongersmai). Sea-level fluctuations provide a plausible biogeographic scenario to explain phylogenetic divergence within this lineage. Given the distinctiveness of the component taxa, and the ease with which even dried skins can be identified to species level (based on ventral counts), the managers of this important commercial resource should no longer treat the P. curtus group as a single biological taxon.     

Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia

Sundberg, P., Gibson, R. & Olsson, U. (2003). Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia. — Zoologica Scripta, 32, 279–296.
Based on 18S rDNA nucleotide sequences and morphological characters, we reconstruct the phylogeny for a group of palaeonemerteans estimated to be monophyletic. Two new palaeonemertean species from Queensland and the Great Barrier Reef, Australia are included in the phylogenetic analysis. The results confirm that one of the species, Cephalothrix queenslandica sp. n., is part of the Cephalothrix–Cephalotrichella–Procephalothrix group. These genera are redefined phylogenetically under the name Cephalothrix based on the cladistic analysis. The other species, Balionemertes australiensis gen. et sp. n., is placed in a new genus which forms a sister taxon to Cephalothrix . The morphology of both new species is described in detail.     

Phylogeny of colletid bees (Hymenoptera: Colletidae) inferred from four nuclear genes

Colletidae comprise approximately 2500 species of bees primarily distributed in the southern continents (only two colletid genera are widely distributed: Colletes and Hylaeus). Previously published studies have failed to resolve phylogenetic relationships on a worldwide basis and this has been a major barrier to the progress of research regarding systematics and evolution of colletid bees. For this study, data from four nuclear gene loci: elongation factor-1alpha (F2 copy), opsin, wingless, and 28S rRNA were analyzed for 122 species of colletid bees, representing all subfamilies and tribes currently recognized; 22 species belonging to three other bee families were used as outgroups. Bayesian, maximum likelihood, and parsimony methods were employed to investigate the phylogenetic relationships within Colletidae and resulted in highly congruent and well-resolved trees. The phylogenetic results show that Colletidae are monophyletic and that all traditionally recognized subfamilies (except Paracolletinae) are also strongly supported as monophyletic. Our phylogenetic hypothesis provides a framework within which broad questions related to the taxonomy, biogeography, morphology, evolution, and ecology of colletid bees can be addressed.     

Establishing a Framework for a Natural Area Taxonomy

The identification of areas of endemism is essential in building an area classification, but plays little role in how natural areas are discovered. Rather area monophyly, derived from cladistics, is essential in the discovery of natural area classifications or area taxonomy. We propose Area Taxonomy to be a new sub-discipline of historical biogeography, one that can be revised and debated, and which has its own area nomenclature. Separately to area taxonomy, we outline how natural areas may be discovered by transcribing the concepts of homology and monophyly from biological systematics to historical biogeography, in the form of area homologues, area homologies and area monophyly.     

The phylogenetic position of some Philippine "babblers" spans the muscicapoid and sylvioid bird radiations

The Philippines is characterized by a high rate of endemism among its terrestrial vertebrates, including enigmatic genera with uncertain affinities. In a recent comprehensive study of the avian family of Timaliidae (babblers), it was shown that three putative babbler genera endemic to the Philippines (Leonardina, Robsonius, and Micromacronus) are distant relatives of Timaliidae. With additional DNA sequences from new samples and data from Genbank, we attempt to determine the phylogenetic affinities of these three genera and examine the resulting implications for biogeography and avian endemism in the Philippines. Well-supported phylogenies recover the three genera in three different families spanning the sylvioid and muscicapoid radiations of passerine birds. Leonardina groups with Muscicapidae and is most closely related to other isolated montane endemic species in Southeast Asia. Robsonius appears to be an early offshoot of Locustellidae. Micromacronus belongs in Cisticolidae, but its position in the family is unresolved. Contrary to implications based on traditional taxonomy, the Philippine archipelago appears to have played a minor role in the diversification of babblers.     

Using DNA-barcoding for sorting out protist species complexes: A case study of the Nebela tincta–collaris–bohemica group (Amoebozoa; Arcellinida,Hyalospheniidae)

Species identification by means of morphology is often problematic in protists. Nebela tincta–collaris–bohemica (Arcellinida) is a species complex of small to medium-sized (ca.100 μm) testate amoebae common in peat bogs and forest soils. The taxonomic validity of characters used to define species within this group is debated and causes confusion in studies of biogeography, and applications in palaeoecology.We examined the relationship between morphological and genetic diversity within this species complex by combined analyses of light microscopy imaging and Cytochrome Oxidase Subunit 1(COI) sequences obtained from the same individual amoeba cells. Our goals were (1) to clarify the taxonomy and the phylogenetic relationships within this group, and (2) to evaluate if individual genotypes corresponded to specific morphotypes and the extent of phenotypic plasticity.We show here that small variations in test morphology that have been often overlooked by traditional taxonomy correspond to distinct haplotypes. We therefore revise the taxonomy of the group. We redefine Nebela tincta (Leidy) Kosakyan et Lara and N. collaris (Ehrenberg 1848) Kosakyan et Gomaa, change N. tincta var. rotunda Penard to N. rotunda (Penard 1890), describe three new species: N. guttata n. sp. Kosakyan et Lara, N. pechorensis n. sp. Kosakyan et Mitchell, and N. aliciae n. sp. Mitchell et Lara.     

Entomopathogenic Nematodes as a Model System for Advancing the Frontiers of Ecology

Entomopathogenic nematodes (EPNs) in the families Heterorhabditidae and Steinernematidae have a mutualistic–symbiotic association with enteric γ-Proteobacteria (Steinernema–Xenorhabdus and Heterorhabditis–Photorhabdus), which confer high virulence against insects. EPNs have been studied intensively because of their role as a natural mortality factor for soil-dwelling arthropods and their potential as biological control agents for belowground insect pests. For many decades, research on EPNs focused on the taxonomy, phylogeny, biogeography, genetics, physiology, biochemistry and ecology, as well as commercial production and application technologies. More recently, EPNs and their bacterial symbionts are being viewed as a model system for advancing research in other disciplines such as soil ecology, symbiosis and evolutionary biology. Integration of existing information, particularly the accumulating information on their biology, into increasingly detailed population models is critical to improving our ability to exploit and manage EPNs as a biological control agent and to understand ecological processes in a changing world. Here, we summarize some recent advances in phylogeny, systematics, biogeography, community ecology and population dynamics models of EPNs, and describe how this research is advancing frontiers in ecology.     

Acknowledging uncertainty in evolutionary reconstructions of ecological niches

Reconstructing ecological niche evolution can provide insight into the biogeography and diversification of evolving lineages. However, comparative phylogenetic methods may infer the history of ecological niche evolution inaccurately because (a) species' niches are often poorly characterized; and (b) phylogenetic comparative methods rely on niche summary statistics rather than full estimates of species' environmental tolerances. Here, we propose a new framework for coding ecological niches and reconstructing their evolution that explicitly acknowledges and incorporates the uncertainty introduced by incomplete niche characterization. Then, we modify existing ancestral state inference methods to leverage full estimates of environmental tolerances. We provide a worked empirical example of our method, investigating ecological niche evolution in the New World orioles (Aves: Passeriformes: Icterus spp.). Temperature and precipitation tolerances were generally broad and conserved among orioles, with niche reduction and specialization limited to a few terminal branches. Tools for performing these reconstructions are available in a new R package called nichevol.     

Discriminating ecological processes affecting different dimensions of α‐ and β‐diversity in desert plant communities

Understanding the spatial distribution of plant diversity and its drivers are major challenges in biogeography and conservation biology. Integrating multiple facets of biodiversity (e.g., taxonomic, phylogenetic, and functional biodiversity) may advance our understanding on how community assembly processes drive the distribution of biodiversity. In this study, plant communities in 60 sampling plots in desert ecosystems were investigated. The effects of local environment and spatial factors on the species, functional, and phylogenetic α‐ and β‐diversity (including turnover and nestedness components) of desert plant communities were investigated. The results showed that functional and phylogenetic α‐diversity were negatively correlated with species richness, and were significantly positively correlated with each other. Environmental filtering mainly influenced species richness and Rao quadratic entropy; phylogenetic α‐diversity was mainly influenced by dispersal limitation. Species and phylogenetic β‐diversity were mainly consisted of turnover component. The functional β‐diversity and its turnover component were mainly influenced by environmental factors, while dispersal limitation dominantly effected species and phylogenetic β‐diversity and their turnover component of species and phylogenetic β‐diversity. Soil organic carbon and soil pH significantly influenced different dimensions of α‐diversity, and soil moisture, salinity, organic carbon, and total nitrogen significantly influenced different dimensions of α‐ and β‐diversity and their components. Overall, it appeared that the relative influence of environmental and spatial factors on taxonomic, functional, and phylogenetic diversity differed at the α and β scales. Quantifying α‐ and β‐diversity at different biodiversity dimensions can help researchers to more accurately assess patterns of diversity and community assembly.     

Soil fungal communities of grasslands are environmentally structured at a regional scale in the Alps

Studying patterns of species distributions along elevation gradients is frequently used to identify the primary factors that determine the distribution, diversity and assembly of species. However, despite their crucial role in ecosystem functioning, our understanding of the distribution of below‐ground fungi is still limited, calling for more comprehensive studies of fungal biogeography along environmental gradients at various scales (from regional to global). Here, we investigated the richness of taxa of soil fungi and their phylogenetic diversity across a wide range of grassland types along a 2800 m elevation gradient at a large number of sites (213), stratified across a region of the Western Swiss Alps (700 km²). We used 454 pyrosequencing to obtain fungal sequences that were clustered into operational taxonomic units (OTUs). The OTU diversity–area relationship revealed uneven distribution of fungal taxa across the study area (i.e. not all taxa are everywhere) and fine‐scale spatial clustering. Fungal richness and phylogenetic diversity were found to be higher in lower temperatures and higher moisture conditions. Climatic and soil characteristics as well as plant community composition were related to OTU alpha, beta and phylogenetic diversity, with distinct fungal lineages suggesting distinct ecological tolerances. Soil fungi, thus, show lineage‐specific biogeographic patterns, even at a regional scale, and follow environmental determinism, mediated by interactions with plants.