Robert W. Scotland

Primary Homology Assessment, Characters and Character States☆

We discuss contrasting approaches to cladistic character definition and thus to cladistic data matrix compilation. The conventional approach considers character states as alternate forms of the “same thing” (the character). A review of the challenges to this convention is presented, and their implications evaluated. We argue that the recognition of structures which are alternate forms is a vital stage of primary homology assessment and is equivalent to the conceptualization of a transformational homology. Such a view complies with the demand that characters are independent and that character states are hierarchically related. We identify one justifiable solution to the inapplicable data coding problem (coding for organisms which have red tails, blue tails or no tails), and show that alternative approaches to character definition support spurious solutions which deny the relation of structures which are “the same but different”. We propose that the term character can be defined, in a cladistic context, as the descriptive label referring to a transformational homology evidenced by the similarity criterion.

Independent recruitment of a conserved developmental mechanism during leaf evolution

Vascular plants evolved in the Middle to Late Silurian period, about 420 million years ago. The fossil record indicates that these primitive plants had branched stems with sporangia but no leaves. Leaf-like lateral outgrowths subsequently evolved on at least two independent occasions. In extant plants, these events are represented by microphyllous leaves in lycophytes (clubmosses, spikemosses and quillworts) and megaphyllous leaves in euphyllophytes (ferns, gymnosperms and angiosperms). Our current understanding of how leaves develop is restricted to processes that operate during megaphyll formation. Because microphylls and megaphylls evolved independently, different mechanisms might be required for leaf formation. Here we show that this is not so. Gene expression data from a microphyllous lycophyte, phylogenetic analyses, and a cross-species complementation experiment all show that a common developmental mechanism can underpin both microphyll and megaphyll formation. We propose that this mechanism might have operated originally in the context of primitive plant apices to facilitate bifurcation. Recruitment of this pathway to form leaves occurred independently and in parallel in different plant lineages.

Herbaria are a major frontier for species discovery

Despite the importance of species discovery, the processes including collecting, recognizing, and describing new species are poorly understood. Data are presented for flowering plants, measuring quantitatively the lag between the date a specimen of a new species was collected for the first time and when it was subsequently described and published. The data from our sample of new species published between 1970 and 2010 show that only 16% were described within five years of being collected for the first time. The description of the remaining 84% involved much older specimens, with nearly one-quarter of new species descriptions involving specimens >50 y old. Extrapolation of these results suggest that, of the estimated 70,000 species still to be described, more than half already have been collected and are stored in herbaria. Effort, funding, and research focus should, therefore, be directed as much to examining extant herbarium material as collecting new material in the field.

How many species of seed plants are there

Recent estimates of the number of described species of seed plant have varied by as much as 62%. The underlying methodology of these estimates is characterised and discussed. We present a revised figure for the number of seed plants based on estimating rates of synonymy in a sample of recently monographed taxa. We conclude that some recent figures overestimate the number of described seed plant species by more than 200,000. This discrepancy is explained by an over-reliance on checklists and floristic studies that underestimate synonymy rates.

How Much Data are Needed to Resolve a Difficult Phylogeny? Case Study in Lamiales

Reconstructing phylogeny is a crucial target of contemporary biology, now commonly approached through computerized analysis of genetic sequence data. In angiosperms, despite recent progress at the ordinal level, many relationships between families remain unclear. Here we take a case study from Lamiales, an angiosperm order in which interfamilial relationships have so far proved particularly problematic. We examine the effect of changing one factor-the quantity of sequence data analyzed-on phylogeny reconstruction in this group. We use simulation to estimate a priori the sequence data that would be needed to resolve an accurate, supported phylogeny of Lamiales. We investigate the effect of increasing the length of sequence data analyzed, the rate of substitution in the sequences used, and of combining gene partitions. This method could be a valuable technique for planning systematic investigations in other problematic groups. Our results suggest that increasing sequence length is a better way to improve support, resolution, and accuracy than employing sequences with a faster substitution rate. Indeed, the latter may in some cases have detrimental effects on phylogeny reconstruction. Further molecular sequencing-of at least 10,000 bp-should result in a fully resolved and supported phylogeny of Lamiales, but at present the problematic aspects of this tree model remain.

Predicting unknown species numbers using discovery curves

A common approach to estimating the total number of extant species in a taxonomic group is to extrapolate from the temporal pattern of known species descriptions. A formal statistical approach to this problem is provided. The approach is applied to a number of global datasets for birds, ants, mosses, lycophytes, monilophytes (ferns and horsetails), gymnosperms and also to New World grasses and UK flowering plants. Overall, our results suggest that unless the inventory of a group is nearly complete, estimating the total number of species is associated with very large margins of error. The strong influence of unpredictable variations in the discovery process on species accumulation curves makes these data unreliable in estimating total species numbers.

Molecular systematics of Clerodendrum (Lamiaceae): ITS sequences and total evidence

Thirty-three species of Clerodendrum s.l. and five outgroup genera were included in a sequence analysis of internal transcribed spacers of the nuclear ribosomal DNA. The results of the cladistic analysis were compared to and combined with cpDNA restriction site data from a previous study. All molecular data identified four major clades within Clerodendrum s.l. and showed the genus to be polyphyletic. Clerodendrum s.s., minus Konocalyx and Cyclonema, is monophyletic and the genus should be restricted to this group. Cyclonema and Konocalyx form a clade distinct from Clerodendrum s.s., which has been recognized as Rotheca Raf.

What is parallelism

SUMMARY Although parallel and convergent evolution are discussed extensively in technical articles and textbooks, their meaning can be overlapping, imprecise, and contradictory. The meaning of parallel evolution in much of the evolutionary literature grapples with two separate hypotheses in relation to phenotype and genotype, but often these two hypotheses have been inferred from only one hypothesis, and a number of subsidiary but problematic criteria, in relation to the phenotype. However, examples of parallel evolution of genetic traits that underpin or are at least associated with convergent phenotypes are now emerging. Four criteria for distinguishing parallelism from convergence are reviewed. All are found to be incompatible with any single proposition of homoplasy. Therefore, all homoplasy is equivalent to a broad view of convergence. Based on this concept, all phenotypic homoplasy can be described as convergence and all genotypic homoplasy as parallelism, which can be viewed as the equivalent concept of convergence for molecular data. Parallel changes of molecular traits may or may not be associated with convergent phenotypes but if so describe homoplasy at two biological levels—genotype and phenotype. Parallelism is not an alternative to convergence, but rather it entails homoplastic genetics that can be associated with and potentially explain, at the molecular level, how convergent phenotypes evolve.

Deep homology: A view from systematics

Over the past decade, it has been discovered that disparate aspects of morphology – often of distantly related groups of organisms – are regulated by the same genetic regulatory mechanisms. Those discoveries provide a new perspective on morphological evolutionary change. A conceptual framework for exploring these research findings is termed ‘deep homology’. A comparative framework for morphological relations of homology is provided that distinguishes analogy, homoplasy, plesiomorphy and synapomorphy. Four examples – three from plants and one from animals – demonstrate that homologous developmental mechanisms can regulate a range of morphological relations including analogy, homoplasy and examples of uncertain homology. Deep homology is part of a much wider range of phenomena in which biological (genes, regulatory mechanisms, morphological traits) and phylogenetic levels of homology can both be disassociated. Therefore, to understand homology, precise, comparative, independent statements of both biological and phylogenetic levels of homology are necessary.

Widespread mistaken identity in tropical plant collections

Specimens of plants and animals preserved in museums are the primary source of verifiable data on the geographical and temporal distribution of organisms. Museum datasets are increasingly being uploaded to aggregated regional and global databases (e.g. the Global Biodiversity Information Facility; GBIF) for use in a wide range of analyses. Thus, digitisation of natural history collections is providing unprecedented information to facilitate the study of the natural world on a global scale. The digitisation of this information utilises information provided on specimen labels, and assumes they are correctly identified. Here we evaluate the accuracy of names associated with 4,500 specimens of African gingers from 40 herbaria in 21 countries. Our data show that at least 58% of the specimens had the wrong name prior to a recent taxonomic study. A similar pattern of wrongly named specimens is also shown for Dipterocarps and Ipomoea (morning glory). We also examine the number of available plant specimens worldwide. Our data demonstrate that, while the worlds collections have more than doubled since 1970, more than 50% of tropical specimens, on average, are likely to be incorrectly named. This finding has serious implications for the uncritical use of specimen data from natural history collections.