Can members of the south‐western Gila robusta species complex be distinguished by morphological features?

The goal for this project was to re‐examine key morphological characters hypothesized to differentiate Gila intermedia, Gila robusta and Gila nigra and outline methods better suited for making species designations based on morphology. Using a combination of meristic counts, morphological measurements and geometric morphometrics, morphological dissimilarities were quantified among these three putative species. Traditional meristic counts and morphological measurements (i.e. distances between landmarks) were not useful for species identification. Geometric morphometrics, however, identified differences among species, while also suggesting an effect of geographic location on morphological variation. Using canonical variate analysis for the 441 fish sampled in this study, geometric morphometrics accurately predicted true group membership 100% of the time for G. nigra, 97% of the time for G. intermedia and 91% of the time for G. robusta. These results suggest that geometric morphometric analysis is necessary to identify morphological differences among the three species. Geometric morphometric analysis used in this study can be adopted by management officials as a tool to classify unidentified individuals.     

Dealing with allometry in linear and geometric morphometrics: a taxonomic case study in the Leporinus cylindriformis group (Characiformes: Anostomidae) with description of a new species from Suriname

To achieve maximum efficacy, taxonomic studies that seek to distinguish amongst species must first account for allometric shape variation within species. Two recently developed software packages (SMATR and MorphoJ) offer regression‐based allometric approaches that are notable for their statistical power and ease of use and that may prove highly useful to taxonomists working with linear or geometric morphometric data. We investigate species delimitation of the slender‐bodied fishes in the Leporinus cylindriformis group using these programs and demonstrate the utility of the allometric corrections that they provide. Without allometric correction, many pairs of species are difficult to distinguish on the basis of morphometrics, but once regressions are used to account for marked allometric variation within species, most of the recognized species in this group can be readily distinguished with linear or geometric morphometrics, particularly using variation in the depth of the body. Both approaches returned congruent patterns of separation amongst putative species, but the geometric approach in MorphoJ distinguished amongst four more pairs of species than did the linear approach in SMATR and appears to provide slightly more statistical power. Based on distinctive morphometrics, meristics, and coloration, a highly elongate species of Leporinus from the Suriname, Corantijn, and Coppename rivers of Suriname is described herein as a new species, Leporinus apollo sp. nov. The unique L. cylindriformis holotype from Porto de Moz, Brazil differs in morphology, meristics, and pigmentation from specimens commonly referred to that species from the main basin of the Amazon; the latter specimens may represent an additional undescribed species. The L. cylindriformis holotype itself may represent a rare species or a specimen collected at the edge of its native range. Measurements of the holotype and paratype of Leporinus niceforoi, which were collected in the Amazonian slope of Colombia, differ substantially from similarly pigmented and putatively conspecific specimens from Amazonian portions of Ecuador and Peru. Recently collected specimens from Colombia are needed to determine whether the observed morphometric variation encompassed by the current concept of L. niceforoi indicates a morphocline within a single species, suggests the presence of multiple cryptic species, or results from shrinkage of the types. In all these cases, linear or geometric morphometric data can reliably differentiate amongst species, but only after one accounts for allometric shape variation. The new SMATR and MorphoJ software packages both offer easy and effective approaches to such allometrically informed taxonomy, and may prove useful to any systematist working on taxa that change shape as they grow. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 103–130.     

Employing statistical learning to derive species-level genetic diversity for mammalian species

1. The patterns of genetic diversity in several genomic regions have been used in mammalian systematics for decades. For instance, when studying closely related species, it is generally assumed that the mitochondrial cytochrome b gene (cytb) exhibits significant information that can be used for differentiation between intraspecies and interspecies variation in mammals. Because of sampling limitations, early analyses of this proposition were conducted mainly on rodents and bats.
2. Currently, more than 57000 cytb sequences are available covering all major lineages of mammals, and sequencing of several individuals per species is common practice in molecular systematics. We were thus prompted to carry out a large-scale analysis of the utility of cytb genetic variation as a predictor of whether a pair of sequences came from within-species or between-species comparisons.
3. Using predetermined species-level assignments, we employed standard methods from statistical learning to calculate the cut-off values able to classify genetic distances in either intraspecies or interspecies categories; we then measured the performance of such statistical classifiers to predict the species-level taxonomic rank as defined by experts.
4. Depending on the classifier, our results showed that when adopting cytb distance cut-off values of 7.3% and 5.5% for small mammals (Metatheria, Rodentia, Chiroptera, and Eulipotyphla) and 4.3% and 3% for medium-sized to large mammals (Primates, Carnivora, and Artiodactyla), the frequency of incorrect assignment of within-species divergences to the between-species category (type I error) varied from 7 to 11%.
5. In order to avoid over-splitting by future researchers, we calculated cut-off values using a more conservative evaluation and provided a list of mammalian species that are likely to consist of complexes of cryptic species. We show that our supervised method can provide practical guidelines to improve the performance of unsupervised algorithms for species delimitation. Finally, we discuss limitations of large-scale approaches (e.g. effects of misclassification in databases and the need for case-by-case evaluation of cryptic species complexes) and their consequences for conservation policies.

     

Inter- and intra-researcher variation in measurement of morphometrics on Steller sea lions (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>)

Substantial effort has been made to increase the accuracy of photogrammetric measurement techniques, especially on marine mammals. However, accuracy of the physical measurements is rarely considered when calculating the photogrammetric accuracy. We examined inter- and intra-researcher variation in the measurement of physical morphometrics of Steller sea lions (Eumetopias jubatus) to assess their potential effects on photogrammetric accuracy and body mass predictions based on photogrammetrically derived morphometrics. Significant differences were found inter-researcher in median standard length and axillary girth and intra-researcher when measuring axillary girth of the same animals. Photogrammetrically derived estimates of these parameters are comparable to physical measurements when taking into account the inherent inter- and intra-researcher variation.     

Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.     

Using Delaunay triangulation to sample whole‐specimen color from digital images

1. Color variation is one of the most obvious examples of variation in nature, but biologically meaningful quantification and interpretation of variation in color and complex patterns are challenging. Many current methods for assessing variation in color patterns classify color patterns using categorical measures and provide aggregate measures that ignore spatial pattern, or both, losing potentially important aspects of color pattern.
2. Here, we present Colormesh, a novel method for analyzing complex color patterns that offers unique capabilities. Our approach is based on unsupervised color quantification combined with geometric morphometrics to identify regions of putative spatial homology across samples, from histology sections to whole organisms. Colormesh quantifies color at individual sampling points across the whole sample.
3. We demonstrate the utility of Colormesh using digital images of Trinidadian guppies (Poecilia reticulata), for which the evolution of color has been frequently studied. Guppies have repeatedly evolved in response to ecological differences between up‐ and downstream locations in Trinidadian rivers, resulting in extensive parallel evolution of many phenotypes. Previous studies have, for example, compared the area and quantity of discrete color (e.g., area of orange, number of black spots) between these up‐ and downstream locations neglecting spatial placement of these areas. Using the Colormesh pipeline, we show that patterns of whole‐animal color variation do not match expectations suggested by previous work.
4. Colormesh can be deployed to address a much wider range of questions about color pattern variation than previous approaches. Colormesh is thus especially suited for analyses that seek to identify the biologically important aspects of color pattern when there are multiple competing hypotheses or even no a priori hypotheses at all.

     

Perception and knowledge of the Sirex woodwasp and other forest pests in South Africa

• 1 Sirex noctilio F. (Siricidae: Hymenoptera) is one of the most serious invasive pests of pine. In South Africa, there has been a national effort to control S. noctilio, including a campaign to increase awareness of the pest amongst the local forestry community.
• 2 We considered the impacts that the arrival of the pest and the awareness campaign have had on perceptions and knowledge of S. noctilio, as well as other forestry pests, amongst members of the forestry community.
• 3 For data collection, a survey questionnaire was developed and used in telephone interviews.
• 4 The results obtained in the present study showed that the Sirex awareness campaign had increased knowledge of forestry pests in general but basic knowledge regarding the identification and symptoms of specific pests, such as S. noctilio, was poor. This will negatively influence monitoring efficacy.
• 5 Traditional paper‐based media and personal contact contributed most to enhanced awareness. Electronic media were less effective and improvement would require a more focused effort. It was of concern that private farmers and contractors, as well as non‐European first language speakers, were less well informed about forestry pests.
• 6 Clearly, a fragmented landscape in terms of ownership and language, presents challenges for effective communication of forestry pest threats.

     

Breakdown of species boundaries in Mandevilla: floral morphological intermediacy,novel fragrances and asymmetric pollen flow

• Phenotypic intermediacy is an indicator of putative hybrid origin and has provided the main clues to discovering hybrid plants in nature. Mandevilla pentlandiana and M. laxa (Apocynaceae) are sister species with clear differences in floral phenotype and associated pollinator guilds: diurnal Hymenoptera and nocturnal hawkmoths, respectively. The presence of individuals with intermediate phenotypes in a wild population raises questions about the roles of visual and olfactory signals (i.e. corolla morphology and floral fragrances) as barriers to interbreeding, and how the breakdown of floral isolation occurs.
• We examined phenotypic variation in a mixed Mandevilla population, analysing the chemical composition of floral fragrances, characterising floral shape through geometric morphometrics and assessing individual grouping through taxonomically relevant traits and an unsupervised learning algorithm. We quantified the visitation frequencies of floral visitors and tracked their foraging movements using pollen analogues.
• The presence of morphologically intermediate individuals and pollen analogue movement suggested extensive hybridisation between M. laxa and M. pentlandiana, along with asymmetrical rates of backcrossing between these putative hybrids and M. laxa. Floral volatiles from putative hybrid individuals showed a transgressive phenotype, with additional compounds not emitted by either parental species.
• Our results suggest the presence of a hybrid swarm between sympatric M. pentlandiana and M. laxa and indicate that initial hybridisation events between these parental species are rare, but once they occur, visits between putative hybrids and M. laxa are common and facilitate continued introgression.

     

Ecological clustering within a diverse minnow assemblage according to morphological,dietary and isotopic data

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Detecting shrub encroachment in seminatural grasslands using UAS LiDAR

1. Shrub encroachment in seminatural grasslands threatens local biodiversity unless management is applied to reduce shrub density. Dense vegetation of Cytisus scoparius homogenizes the landscape negatively affecting local plant diversity. Detecting structural change (e.g., biomass) is essential for assessing negative impacts of encroachment. Hence, exploring new monitoring tools to achieve this task is important for effectively capturing change and evaluating management activities.
2. This study combines traditional field‐based measurements with novel Light Detection and Ranging (LiDAR) observations from an Unmanned Aircraft System (UAS). We investigate the accuracy of mapping C. scoparius in three dimensions (3D) and of structural change metrics (i.e., biomass) derived from ultrahigh‐density point cloud data (>1,000 pts/m²). Presence–absence of 12 shrub or tree genera was recorded across a 6.7 ha seminatural grassland area in Denmark. Furthermore, 10 individuals of C. scoparius were harvested for biomass measurements. With a UAS LiDAR system, we collected ultrahigh‐density spatial data across the area in October 2017 (leaf‐on) and April 2018 (leaf‐off). We utilized a 3D point‐based classification to distinguish shrub genera based on their structural appearance (i.e., density, light penetration, and surface roughness).
3. From the identified C. scoparius individuals, we related different volume metrics (mean, max, and range) to measured biomass and quantified spatial variation in biomass change from 2017 to 2018. We obtained overall classification accuracies above 86% from point clouds of both seasons. Maximum volume explained 77.4% of the variation in biomass.
4. The spatial patterns revealed landscape‐scale variation in biomass change between autumn 2017 and spring 2018, with a notable decrease in some areas. Further studies are needed to disentangle the causes of the observed decrease, for example, recent winter grazing and/or frost events.
5. Synthesis and applications: We present a workflow for processing ultrahigh‐density spatial data obtained from a UAS LiDAR system to detect change in C. scoparius. We demonstrate that UAS LiDAR is a promising tool to map and monitor grassland shrub dynamics at the landscape scale with the accuracy needed for effective nature management. It is a new tool for standardized and nonbiased evaluation of management activities initiated to prevent shrub encroachment.

     

Morphometric comparison of the cephalic region of cultured Acipenser baerii (Brandt, 1869), Acipenser naccarii (Bonaparte, 1836) and their hybrid*

Geometric morphometric techniques were used in the characterization of morphological differences of the cranial region in cultured Acipenser baerii and A. naccarii and their hybrid. Species identification in sturgeons sometimes is difficult, because of the morphological plasticity of individual variability and the small differences between taxa. Identification is also complicated by human intervention and manipulation (e.g. aquaculture, hybridization, pollution). This study indicates how the differences between two species and their hybrids can be quantified and accurately visualized via geometric morphometrics. This technique may serve as a useful methodology to supplement analytical studies on morphometrics and systematics in Acipenseridae.     

Assessing the taxonomic status of dingoes Canis familiaris dingo for conservation

• 1 The conservation status of the dingo Canis familiaris dingo is threatened by hybridization with the domestic dog C. familiaris familiaris. A practical method that can estimate the different levels of hybridization in the field is urgently required so that animals below a specific threshold of dingo ancestry (e.g. 1/4 or 1/2 dingoes) can reliably be identified and removed from dingo populations.
• 2 Skull morphology has been traditionally used to assess dingo purity, but this method does not discriminate between the different levels of dingo ancestry in hybrids. Furthermore, measurements can only be reliably taken from the skulls of dead animals.
• 3 Methods based on the analysis of variation in DNA are able to discriminate between the different levels of hybridization, but the validity of this method has been questioned because the materials currently used as a reference for dingoes are from captive animals of unproven genetic purity. The use of pre‐European materials would improve the accuracy of this method, but suitable material has not been found in sufficient quantity to develop a reliable reference population. Furthermore, current methods based on DNA are impractical for the field‐based discrimination of hybrids because samples require laboratory analysis.
• 4 Coat colour has also been used to estimate the extent of hybridization and is possibly the most practical method to apply in the field. However, this method may not be as powerful as genetic or morphological analyses because some hybrids (e.g. Australian cattle dog × dingo) are similar to dingoes in coat colour and body form. This problem may be alleviated by using additional visual characteristics such as the presence/absence of ticking and white markings.

     

Systematics as science: A response to Cronquist

Our reply to the commentary on cladistics presented by Cronquist (1987) is aimed at four issues:

1. the application of scientific principles in systematics;
2. the recognition that the analysis of pattern is a vital precursor to any consideration of evolutionary process. A priori judgements of evolutionary process are unnecessary for the generation of informative systematic hypotheses which are chosen for their ability to explain the patterns of character distributions rather than for compatibility with any particular preconceived ideas about evolution;
3. that phenetic concepts such as overall similarity, grades, gaps, and degree of divergence, if included in methods of phylogenetic inference, will give erroneous results. Paraphyletic and polyphyletic groups must, consequently, be rejected from systematics since they have no rational empirical basis for recognition;
4. the fact that many of the problems of phylogenetic analysis attributed by Cronquist to cladistics are common to all systematic methods but that these can be dealt with by the application of such principles as parsimony, synapomorphy, and strict monophyly.

     

Estimating red deer Cervus elaphus populations: an analysis of variation and cost-effectiveness of counting methods

• 1 Different counting methods are currently used to estimate red deer populations in the open range in Scotland, but there are few data available to compare variation in estimates, or relative cost‐effectiveness.
• 2 While it is impossible to determine the accuracy of counts (as real numbers are unknown), variation within and between different methods can be measured by repeat counts of the same area within as short a period as possible.
• 3 This study aimed to quantify the variation observed from repeat counts using each of four methods (ground, helicopter, infrared helicopter and dung‐counting methods) at one of three study sites in late winters 2003, 2004 and 2005. Additional data from digital camera images of groups from counts in other areas of Scotland were also used to assess the accuracy of visual counts.
• 4 Coefficients of variation (CVs) within any method of between 5% and 16% were recorded, consistent with previous comparisons for red deer open range counts in Scotland. CVs were lowest for ground and helicopter counts. The infrequency of optimal conditions was likely to limit the applicability of infrared counts in Scotland.
• 5 In terms of cost‐effectiveness, helicopter counting was the least labour‐intensive, with costs of other techniques depending on the availability of existing manpower as an overhead cost.
• 6 It is concluded that helicopter counts are most likely to minimize errors while maximizing cost‐efficiency. Accuracy can be improved by the use of digital photography for counting larger deer groups. Estimates are likely to be improved further by increasing the frequency of counts and using the same methods, counters and routes for repeat counts.

     

Traditional morphometrics in plant systematics and its role in palm systematics

A review is given of the role of traditional morphometrics in plant systematics. The three most commonly used techniques of data analysis – Cluster Analysis, Principal Component Analysis and Discriminant Analysis – are discussed. The kinds of data that can be taken from palm specimens and the problems of using specimens as data sources are outlined. Published systematic studies of palms using traditional morphometrics are reviewed. More recent studies indicate that: hybrid zones between species may be common; infraspecific diversity is greater than previously suspected; there may be more than double the currently accepted number of species; and our current knowledge of morphological variation in palms is superficial. A procedure for scientific systematics is given, which incorporates traditional morphometric methods.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 103–111.     

Mimetic colour pattern evolution in the highly polymorphic Bombus trifasciatus (Hymenoptera: Apidae) species complex and its comimics

Müllerian mimetic systems have uncovered some of the dynamic processes by which natural selection can drive the radiation of convergent and divergent phenotypes. We examined evolution involving Müllerian mimicry in bumble bees by documenting the distribution and evolution of colour patterns amongst three colour‐polymorphic lineages –Bombus trifasciatus Smith, Bombus haemorrhoidalis Smith, and Bombus breviceps Smith – that mimic each other across ～14 colour groups in South‐East Asia. Using mitochondrial DNA sequence data, we estimated relationships within each lineage to infer the processes that gave rise to the colour diversity and develop hypotheses on species recognition. We expanded on our assessment of species delineation in the B. trifasciatus lineage using three nuclear gene fragments and morphometrics. Comparison of colour patterns amongst georeferenced specimens showed considerable variation in the degree and geographical range of mimicry amongst mimicry groups. Phylogenetic estimates show high rates of colour pattern evolution, with colour variation often exceeding variation within the fast‐evolving mitochondrial genes. The molecular data, and to some degree the morphometric data, support unique histories for several taxa recognized previously within the B. trifasciatus lineage, which may include several species. Early vicariant events within the B. trifasciatus lineage are likely to have occurred ～2.2 Mya in the mountains of south‐west China. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 805–826.