Diversity and biogeography of a species‐rich ant fauna of the Australian seasonal tropics

Although ants are an ecologically dominant and extensively studied faunal group throughout the tropics, there is a poor understanding of tropical ant diversity and distribution at large spatial scales. Here we use a collection developed from 3 decades of ant surveys to present the first analysis of ant diversity and biogeography of a large tropical region. Our objective was to document the species richness, composition, and biogeographic distributions of the ant fauna of the 400 000 km² “Top End” of Australia's Northern Territory. The known Top End ant fauna comprises 901 native species from 59 genera. The richest genera are Pheidole (90 species), Melophorus (83), Monomorium (83), Camponotus (71), Meranoplus (63), Polyrhachis (57), Rhytidoponera (50), Tetramorium (43), Cerapachys (32), and Iridomyrmex (31). The fauna is the center of diverse radiations within species‐groups of genera such as Meranoplus, Rhytidoponera, and Leptogenys. It also includes IndoMalayan species that have likely bypassed the normal dispersal route into Australia through Cape York Peninsula in North Queensland. Faunistic similarity with other regions of far northern Australia is associated more with rainfall than with geographic proximity. Most (60%) of Top End ant species have not been recorded elsewhere, and, despite uncertainties relating to species delimitation and sampling intensity, this appears to be a credible estimate of the level of endemism. Such exceptionally high endemism can be attributed to the Top End's geographic isolation from other regions of northern Australia with comparably high rainfall.     

Molecular and morphological diversity of Narragansett Bay (RI,USA) Ulva (Ulvales,Chlorophyta) populations

Macroalgal bloom‐forming species occur in coastal systems worldwide. However, due to overlapping morphologies in some taxa, accurate taxonomic assessment and classification of these species can be quite challenging. We investigated the molecular and morphological characteristics of 153 specimens of bloom‐forming Ulva located in and around Narragansett Bay, RI, USA. We analyzed sequences of the nuclear internal transcribed spacer 1 region (ITS1) and the chloroplast‐encoded rbcL; based on the ITS1 data, we grouped the specimens into nine operational taxonomic units (OTUs). Eight of these OTUs have been previously reported to exist, while one is novel. Of the eight OTUs, all shared sequence identity with previously published sequences or differed by less than 1.5% sequence divergence for two molecular markers. Previously, 10 species names were reported for Ulva in Rhode Island (one blade and nine tube‐forming species) based upon morphological classification alone. Of our nine OTUs, three contained blade‐forming specimens (U. lactuca, U. compressa, U. rigida), one OTU had a blade with a tubular stipe, and six contained unbranched and/or branched tubular morphologies (one of these six, U. compressa, had both a blade and a tube morphology). While the three blade‐forming OTUs in Narragansett Bay can frequently be distinguished by careful observations of morphological characteristics, and spatial/temporal distribution, it is much more difficult to distinguish among the tube‐forming specimens based upon morphology or distribution alone. Our data support the molecular species concept for Ulva, and indicate that molecular‐based classifications of Ulva species are critical for proper species identification, and subsequent ecological assessment or mitigation of Ulva blooms.     

A phenetic investigation of Psophocarpus Neck.ex DC.(Leguminosae-Phaseoleae)

Morphological data were gathered from 126 herbarium specimens representing ten species of Psophocarpus. Specimens were scored for 29 vegetative, 46 inflorescence, 16 legume and six seed characters. The data were analysed using cluster analysis and ordination and the results used to produce an overall phenetic classification. The proposed classification includes a tenth species recently distinguished, but undescribed by Verdcourt. Keys are provided to the subgenera and sections, as well as the ten taxa recognized and the classification is discussed in relation to previous classifications of the genus.     

Taxonomic and biostratigraphical re-assessments of Subterraniphyllum Elliott (Corallinales, Rhodophyta)

A taxonomic and biostratigraphical re‐assessment of Subterraniphyllum Elliott (Corallinales, Rhodophyta) is presented. Results from studies of the type collection and of newly collected material from north‐eastern Italy and northern Slovenia have shown that this taxon is not a geniculate coralline red alga as originally suggested by Elliott and most subsequent authors. Vegetatively, Subterraniphyllum most closely resembles certain living members of the Corallinales; however, the phenetic and phylogenetic relationships of Subterraniphyllum to other Corallinales cannot be determined with greater certainty. The exclusion of Subterraniphyllum from any group of Corallinaceae with genicula is based on unequivocal evidence that branch formation does not involve the occurrence of genicula. Subterraniphyllum seems to be restricted to the Oligocene. Reports of occurrences in Upper Eocene and Lower Miocene sediments cannot be substantiated. Subterraniphyllum, however, cannot be considered a useful stratigraphical marker until further data on its occurrence in well‐dated carbonate sequences are acquired. This study illustrates the problems associated with placing fossil coralline algal specimens into geniculate genera without the preservation of relevant morphological characters. This is especially true in the absence of the careful assessment of fossil material with respect to current taxonomic concepts of geniculate coralline genera, all of which are based on studies of living species. According to the current concepts for geniculate coralline genera, the placing of fossil specimens into geniculate genera without appropriate evidence must be avoided by grouping all potentially geniculate fragments under the informal group ‘Geniculate sensu lato’. Furthermore, for all those many fossil specimens where unequivocal evidence is not present, it is possible to utilize ‘form genera’ based on characters that are normally preserved. This leads to creating a consistent, workable system of applying names to most fossil corallines so that they can be reliably used in relation to stratigraphical and palaeoecological studies.     

Catching the phylogenic history through the ontogenic hourglass: a phylogenomic analysis of Drosophila body segmentation genes

SUMMARY The phylogenetic information content of different developmental stages is a long‐standing issue in the study of development and evolution. We performed phylogenetic analyses of 51 body segmentation genes in 12 species of Drosophila in order to investigate the impact of the mode of evolution of development on phylogeny inference. Previous studies of these genes in Drosophila using pairwise phenetic comparisons at the species group level revealed the presence of an “hourglass model” (HG), wherein mid‐embryonic stages are the most evolutionarily constrained. We utilized two character‐based approaches: taxonomic congruence using the relative consensus fork index (RCFI), in which phylogenies are inferred from each gene separately and compared with a total evidence tree (TET), and partitioned simultaneous analysis using several indices such as branch support (BS) and localized incongruence length difference (LILD) test. We also proposed a new index, the recapitulatory index (R), which divides the number of synapomorphies on the total number of informative characters in a data set. Polynomial adjustment of both BS and R indices showed strong support for the hourglass model regardless of the taxonomic level (species subgroup vs. subgenera), showing less phylogenetic information content for mid‐developmental stages (mainly the zygotic segment polarity stage). Significant LILD scores were randomly distributed among developmental stages revealing the absence of differential selective constraints, but were significantly related to chromosomal location showing physical (linkage) impact on phylogenetic incongruence. RCFI was the most sensitive measure to taxonomic level, having a convex parabola at the species subgroup level in support of the hourglass model and a concave parabola at the subgeneric level in support of the adaptive penetrance model. This time‐dependent discrepancy of best fit developmental model parallels previous conflicting results from the vertebrates. Because of the quasi‐phenetic nature of this index, we argue that the discrepancy is due to the evolutionary rate heterogeneity of developmental genes rather than to fundamental differences among organisms. We suggest that simultaneous character‐based analyses give better macroevolutionary support to the hourglass model of the developmental constraints on genome evolution than pairwise phenetic comparisons.     

Too strict or too loose? Integrative taxonomic assessment of Bombus lapidarius complex (Hymenoptera: Apidae)

The latest progress of the taxonomy is the use of integrative approach for species delimitation based on a multisource dataset. However, the taxonomic decision that should be made when convergence between the different lines of evidence is not observed remains debated. Here, we investigate the consequences of the application of an ‘integration by cumulation’ approach on the taxonomic statuses within the Bombus lapidarius complex when using an integrative taxonomic framework (i.e. nuclear and mitochondrial markers along with reproductive traits) compared with a strict ‘integration by congruence’ method. Our results show similar taxonomic conclusions whatever the decision-making approach used except for one taxon. According to the differentiation observed in our integrative taxonomic framework, recent divergence time and other field observations for this taxon, we assume that a too strict decision-making method could fail to detect recently diverged species. This is exemplified by the new species Bombus bisiculus sp. n. occurring in South Italy and Sicily.     

Cross‐taxon congruence of multiple diversity facets of freshwater assemblages is determined by large‐scale processes across China

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Does the presence of grassy strips and landscape grain affect the spatial distribution of aphids and their carabid predators?

• 1 We investigated, over the course of 2 years, the spatial distribution and abundance of two species of aphid, Metopolophium dirhodum and Sitobion avenae, and predatory species of carabid. This was undertaken in 24 wheat fields in ‘coarse‐grain’ and ‘fine‐grain’ landscapes in western France. A greater percentage of the latter landscape was covered by hedgerows and grassland and the total area covered by fields and the average size of the fields were smaller.
• 2 The effects on aphid abundance of the distance from field margins, the presence of grassy strips and carabid abundance were determined in both landscapes.
• 3 Both aphid species were more abundant in the ‘fine‐grain’ landscape, which may have been a result of the higher density of semi‐natural elements. In both types of landscape, the total numbers of aphids were negatively correlated with the distance from the field margin. This may have been because aphids were dispersing from overwintering sites in field margins. The abundance of M. dirhodum was strongly negatively correlated with the presence of grassy strips in the ‘coarse‐grain’ landscape, although there were no such significant correlations for either of the aphid species in the ‘fine‐grain’ landscape.
• 4 Aphid and carabid abundances were negatively correlated in the ‘fine‐grain’ and positively in ‘coarse‐grain’ landscape.
• 5 The results obtained in the present study emphasize the importance of semi‐natural areas in agricultural landscapes in shaping the spatial distribution of aphids and carabid beetles, their natural enemies, at different spatial scales.

     

Scale and trends in species richness: considerations for monitoring biological diversity for political purposes

Switzerland's governmental ‘Biodiversity Monitoring’ program is designed to produce factual information on the dynamics of biodiversity within the country for governmental agencies, politicians, and the general public. Monitoring a complex issue like biodiversity in order to give relevant and accurate messages to the general public and politicians within a politically relevant timescale and at moderate cost means focusing on few elements. Because relevant human impacts on biodiversity operate differently at different spatial scales, we need at least three different indicators to observe changes over time in local (‘within‐habitat’), landscape (‘habitat‐mosaic’), and macro‐scale (‘regional’) diversity. To keep things as simple as possible, we use species richness as an indicator for all three levels of diversity, just defining three different spatial scales (10 m², 1 km², regions, respectively). Each indicator is based on a number of taxonomic groups which have been selected mainly on the basis of costs and the availability of appropriate methods.     

The canopy beetle faunas of Gondwanan element trees in Chilean temperate rain forests

Aim To describe the coleopteran fauna occurring in canopies of temperate Gondwanan tree species in terms of their diversity and guild and taxonomic structures, and to test the proposition that this structuring reflects the Gondwanan origins of this fauna. Location The Andes and the coastal cordillera of temperate Chile. Methods Canopy fogging was used to sample beetles from 29 trees. The samples were statistically described using S_chao and the Simpson diversity index D. Cluster analyses and multi‐dimensional scaling (MDS) were performed. The taxonomic and guild structures of the Chilean coleopteran fauna were compared quantitatively with those found in other parts of the world using homogeneity chi‐square and t‐tests. Results A collection of 25,497 beetle specimens was obtained primarily from Nothofagus dombeyi, Nothofagus obliqua and Araucaria araucana. The specimens collected were distributed between 485 morphospecies and included 107 putative, new generic‐level taxa and 223 apparently undescribed species. Estimates of the size of the canopy beetle fauna showed that 600+ species were likely to be present. The communities found on a tree species differed markedly between years. MDS plots showed less community divergence between tree species for predators than for phytophages and xylophages. The guild structure was similar to that found on Australian ‘Gondwanan’ trees but differed significantly from the community structures found on ‘Laurasian’ tropical and temperate trees in supporting fewer phytophages and saprophages, but more xylophages. The predator guild showed a different pattern, with tropical faunas differing from those of more temperate regions, irrespective of hemisphere, as did the distribution of superfamilies. Main conclusions The beetle fauna found in the canopies of N. dombeyi, N. obliqua and A. araucana was large (600+ species), with about half of the species undescribed. S_chao was found to vary with sample size and to give lower estimates of S than species attenuation curves, raising the possibility that the two methods are estimating the sizes of different statistical communities. It is possible that the attenuation curve is estimating the number of species to be found on a particular tree species, while S_chao is estimating the ‘carrying capacity’ for beetle species of individual trees, and this varies from tree to tree. Care also needs to be taken in experimental design when monitoring temporal changes in forest insect communities given the difference in communities found between years in this study. The proportions of phytophages, saprophages and xylophages resemble those of a ‘Gondwanan’ rain forest from Australia and differ significantly from those of tropical and temperate ‘Laurasian’ forests.     

A keystone ant species promotes seed dispersal in a “diffuse” mutualism

In order to understand the dynamics of co-evolution it is important to consider spatial variation in interaction dynamics. We examined the relative importance of ant activity, diversity and species identity in an ant seed dispersal mutualism at local, regional and continental scales. We also studied the determinants of seed dispersal rates and dispersal distances at eight sites in the Eneabba sandplain (29.63 S, 115.22 E), western Australia to understand local variation in seed dispersal rate and distance. To test the generality of the conclusions derived from the eight local sites, we established 16 sites along a 1650-km transect in western Australia, covering 11° of latitude and a six-fold increase in rainfall, at which we sampled the ant assemblage, estimated ant species richness and ant activity and observed the removal rate of myrmecochorous seeds. We also assessed the importance of ant species identity at a continental scale via a review of studies carried out throughout Australia which examined ant seed dispersal. Among the eight sandplain shrubland sites, ant species identity, in particular the presence of one genus, Rhytidoponera, was associated with the most dispersal and above average dispersal distances. At the landscape scale, Rhytidoponera presence was the most important determinant of seed removal rate, while seed removal rate was negatively correlated with ant species richness and latitude. Most ant seed removal studies carried out throughout Australia reinforce our observations that Rhytidoponera species were particularly important seed dispersers. It is suggested that superficially diffuse mutualisms may depend greatly on the identity of particular partners. Even at large biogeographic scales, temporal and spatial variation in what are considered to be diffuse mutualisms may often be linked to variation in the abundance of particular partners, and be only weakly – or negatively – associated with the diversity of partners.     

Combining geometric morphometrics with pattern recognition for the investigation of species complexes

The parasitoid Hymenoptera contain a large number of species complexes that are as yet unresolved by traditional taxonomic studies. Evolutionary studies as well as biological control programmes often require further investigations which cannot rely on simple qualitative morphological characters; in many cases, particularly with dried specimens housed in museums, molecular approaches cannot be used. Recent developments in geometric morphometrics and statistical exploratory approaches open new perspectives for the objective evaluation of morphological characters in this taxonomic context. In this study, geometric morphometrics and pattern recognition approaches were applied to the wing shape and venation of two closely related braconid species considered to differ by subtle qualitative morphological head characters. Exploratory analyses such as kernel density estimates and Gaussian mixture analyses were used to explore the structure of the data in the multivariate morphometric space. Discrimination techniques (linear discriminant functions and neural networks combined with cross‐validations) were used to estimate the taxonomic value of qualitative characters. Gaussian mixtures highlighted the existence of two non‐overlapping groups. A good congruence was found between one of the two groups and the a priori defined Bassus tumidulus. The misclassification rate was higher for B. tegularis specimens, which also appeared morphometrically heterogeneous. Discrimination between the two a priori defined species was incomplete with misclassification rates higher than, or equal to, 6%. In most cases, the lack of congruence between species and morphometrically defined subgroups could be related to specimens that exhibited ambiguous qualitative character states. In summary, if two entities are present, they still need to be defined morphologically, while B. tegularis heterogeneity calls for further investigation of specimens of known origin and hosts. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 89–98.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Perspectives: Towards a language for mapping relationships among taxonomic concepts

Abstract

Taxonomic concepts (sensu Berendsohn) embody the underlying meanings of scientific names as stated in a particular publication, thus offering a new way to resolve semantic ambiguities that result from multiple revisions of a taxonomic name. This paper presents a comprehensive and powerful language for representing the relationships among taxonomic concepts. The language features terms and symbols for concept relationships within a single taxonomic hierarchy, or between two related but independently published hierarchies. Taxonomic concepts pertaining to a single hierarchy are characterised by parent/child relationships, whereas those pertaining to two independent hierarchies may have the following basic relationships: congruence, inclusion (non‐symmetrical, relative to the side of comparison), overlap, and exclusion. The relationships are asserted by specialists who have the option to add or subtract concepts on one or both sides of a relationship equation in order to reconcile differences between non‐congruent taxonomic perspectives. The terms ‘and’, ‘or’ and ‘not’ are available, respectively, to connect multiple simultaneously or alternatively valid relationship assessments, or to explicitly negate the validity of a relationship. The language also permits the decomposition of a relationship according to the intensional (property referencing) and ostensive (member pointing) aspects of the compared taxonomic concepts. Adopting the concept relationship language will facilitate a more precise documentation of similarities and differences in multiple succeeding taxonomic perspectives, thereby preparing the stage for an ontology‐based integration of taxonomic and related biological information.