Macaronesian millipedes (Diplopoda) with emphasis on endemic species swarms on Madeira and the Canary Islands

Endemic species swarms constitute large fractions of the millipede faunas of Madeira (29 species of the Cylindroiulus madeirae group, plus six species of Acipes , out of a total of 60 species) and the Canary Islands (46 species of Dolichoiulus , plus four species of the Glomeris alluaudi-group , out of a total of about 79 species). The poorer faunas of the Azores (22 species) and the Cape Verde Islands (15 species) in contrast only include a few endemics. The Cylindroiulus madeirae group and Dolichoiulus show a high degree of diversity of structure (size, colour, leg length etc.) and habitat (laurisilva, xeric habitats, caves). The C. madeirae group, unlike Dolichoiulus , is strongly concentrated in the laurisilva. In this habitat, microhabitat differentiation is pronounced in both swarms.     

Drivers of diversity in Macaronesian spiders and the role of species extinctions

Aim To identify the biogeographical factors underlying spider species richness in the Macaronesian region and assess the importance of species extinctions in shaping the current diversity. Location The European archipelagos of Macaronesia with an emphasis on the Azores and Canary Islands. Methods Seven variables were tested as predictors of single‐island endemics (SIE), archipelago endemics and indigenous spider species richness in the Azores, Canary Islands and Macaronesia as a whole: island area; geological age; maximum elevation; distance from mainland; distance from the closest island; distance from an older island; and natural forest area remaining per island – a measure of deforestation (the latter only in the Azores). Different mathematical formulations of the general dynamic model of oceanic island biogeography (GDM) were also tested. Results Island area and the proportion of remaining natural forest were the best predictors of species richness in the Azores. In the Canary Islands, area alone did not explain the richness of spiders. However, a hump‐shaped relationship between richness and time was apparent in these islands. The island richness in Macaronesia was correlated with island area, geological age, maximum elevation and distance to mainland. Main conclusions In Macaronesia as a whole, area, island age, the large distance that separates the Azores from the mainland, and the recent disappearance of native habitats with subsequent unrecorded extinctions seem to be the most probable explanations for the current observed richness. In the Canary Islands, the GDM model is strongly supported by many genera that radiated early, reached a peak at intermediate island ages, and have gone extinct on older, eroded islands. In the Azores, the unrecorded extinctions of many species in the oldest, most disturbed islands seem to be one of the main drivers of the current richness patterns. Spiders, the most important terrestrial predators on these islands, may be acting as early indicators for the future disappearance of other insular taxa.     

Interpreting colonization of the Calathus (Coleoptera: Carabidae) on the Canary Islands and Madeira through the application of the parametric bootstrap

The Canary Islands have proven to be an interesting archipelago for the phylogeographic study of colonization and diversification with a number of recent studies reporting evolutionary patterns and processes across a diversity of floral and faunal groups. The Canary Islands differ from the Hawaiian and Galapagos Islands by their close proximity to a continental land mass, being 110 km from the northwestern coast of Africa. This close proximity to a continent obviously increases the potential for colonization, and it can be expected that at the level of the genus some groups will be the result of more than one colonization. In this study we investigate the phylogeography of a group of carabid beetles from the genus Calathus on the Canary Islands and Madeira, located 450 km to the north of the Canaries and 650 km from the continent. The Calathus are well represented on these islands with a total of 29 species, and on the continent there are many more. Mitochondrial cytochrome oxidase I and II sequence data has been used to identify the phylogenetic relationships among the island species and a selection of continental species. Specific hypotheses of monophyly for the island fauna are tested with parametric bootstrap analysis. Data suggest that the Canary Islands have been colonized three times and Madeira twice. Four of these colonizations are of continental origin, but it is possible that one Madeiran clade may be monophyletic with a Canarian clade. The Calathus faunas of Tenerife and Madeira are recent in origin, similar to patterns previously reported for La Gomera, El Hierro, and Gran Canaria.     

Parasitoids of whiteflies (Hymenoptera: Aphelinidae,Eulophidae, Platygastridae; Hemiptera: Aleyrodidae) from the Macaronesian archipelagos of the Canary Islands,Madeira and the Azores

Data on the whitefly parasitoid species known from the Macaronesian archipelagos of the Canary Islands, Madeira and the Azores are presented, based largely on recently collected material. A total of 26 species are treated, including six new species, six new records for the Canary Islands, two new records for Madeira, and two new records for the Azores. All species are fully described and illustrated. New species described are: Encarsia atlantica Polaszek & Hernández; Encarsia levadicola Polaszek & Hernández; Encarsia melanostoma Polaszek & Hernández; Encarsia noahi Polaszek & Hernández; Euderomphale gomer LaSalle & Hernández; Euderomphale insularis LaSalle & Hernández. A fully illustrated identification key based on females is provided for recognition of whitefly parasitoids in these archipelagos. Data on the known distribution and hosts are provided, as well as references to biology and use in biological control.     

Molecular phylogeny of the green lacewings (Neuroptera: Chrysopidae)

Abstract  The first quantitative analysis of phylogenetic relationships of green lacewings (Chrysopidae) is presented based on DNA sequence data. A single nuclear and two mitochondrial genes are used in the analysis: carbomoylphosphate synthase (CPS) domain of carbamoyl-phosphate synthetase-aspartate transcarbamoylase-dihydroorotase (CAD) (i.e. rudimentary locus), large subunit ribosomal gene (16S) and cytochrome oxidase I (COI). This study represents the first use of the CAD gene to investigate phylogenetic relationships of the lacewings. DNA sequences for 33 chrysopid species from 18 genera, representing all subfamilies and tribes, were compared with outgroups sampled from families Hemerobiidae, Osmylidae and Polystoechotidae. Parsimony analyses of the combined data set recovered all of the previously established subfamilial and tribal groups as monophyletic clades (although relatively weakly supported) except Apochrysinae sensu lato . The enigmatic Nothancyla verreauxi Navás has historically been difficult to place in a subfamily group based on morphological characteristics; molecular data presented herein do not adequately resolve this problem.     

Mitochondrial DNA, ecology and morphology: interpreting the phylogeography of the Nesotes (Coleoptera: Tenebrionidae) of Gran Canaria (Canary Islands)

The genus Nesotes (Coleoptera: Tenebrionidae) is represented in the Canary Islands by 19 endemic species, the majority of which are single island endemics. Nesotes conformis and N. fusculus are described on four and three islands, respectively, but each forms a paraphyletic assemblage between Gran Canaria and the other islands. The other described species for Gran Canaria are N. quadratus, N. lindbergi and N. piliger. Thirty-six individuals representing the five species on Gran Canaria have been sequenced for 675 bp of the mitochondrial DNA (mtDNA) cytochrome oxidase II gene. Neighbour-joining analysis of maximum likelihood distances resulted in five distinct mtDNA lineages for N. quadratus, two of which also include mitotypes of N. conformis. Each of the other three species is found on only one mtDNA lineage. We propose from the molecular data that differentiation in a widespread N. quadratus-type ancestor was followed by morphological adaptation to coastal, pine and laurel forest habitats.     

Characterization of microsatellite loci in Brachyderes rugatus,the Canary Islands pine weevil (Coleoptera: Curculionidae)

The Canary Island pine weevil Brachyderes rugatus (Wollaston) consists of four allopatric subspecies that are thought to have arisen from several historic colonization events within the archipelago. We have isolated and optimized seven microsatellite loci from Brachyderes rugatus calvus from Gran Canaria. Six of these loci are polymorphic within B. rugatus (11–22 alleles per locus; heterozygosity between 0.43 and 0.84). There is no evidence for heterozygote deficit within populations or for linkage disequilibrium between pairs of loci. These molecular markers are likely to prove useful tools for quantifying the genetic variability of bottlenecked island populations.     

Molecular phylogeny of the nutmeg shells (Neogastropoda, Cancellariidae)

Cancellariidae, or nutmeg shells, is a family of marine gastropods that feed on the body fluids and the egg cases of marine animals. The 300 or so living species are distributed worldwide, mostly on soft bottoms, from intertidal to depths of about 1000 m. Although they are a key group for the understanding of neogastropod evolution, they are still poorly known in terms of anatomy, ecology and systematics. This paper reports the first mitochondrial multi-gene phylogenetic hypothesis for the group. Data were collected for 50 morphospecies, representative of 22 genera belonging to the three currently recognized subfamilies. Sequences from three genes (12S, 16S and COI) were analyzed with Maximum Likelihood analysis and Bayesian Inference, both as single gene datasets and in two partitioned concatenated alignment. Largely consistent topologies were obtained and discussed with respect to the traditional subfamilial arrangements. The obtained phylogenetic trees were also used to produce Robinson-Foulds supertrees. Our results confirmed the monophyly of the subfamily Plesiotritoninae, while Admetinae and Cancellariinae, as currently conceived, were retrieved as polyphyletic. Based on our findings we propose changes to the systematic arrangement of these subfamilies. At a lower taxonomic rank, our results highlighted the rampant homoplasy of many characters traditionally used to segregate genera, and thus the need of a critical re-evaluation of the contents of many genera (e.g. Nipponaphera, Merica, Sydaphera, Bivetia), the monophyly of which was not recovered.     

Reconciling gene trees with organism history: the mtDNA phylogeography of three Nesotes species (Coleoptera: Tenebrionidae) on the western Canary Islands

The processes of island colonization and speciation are investigated through mtDNA studies on Canary Island beetles. The genus Nesotes (Coleoptera: Tenebrionidae) is represented by 19 endemic species on the Canary Islands, the majority of which are single island endemics. Nesotes conformis is the most widespread, occurring on Gran Canaria, Tenerife, La Palma and El Hierro. Nesotes conformis forms a paraphyletic assemblage, with a split between Gran Canaria and the other three islands. Nesotes conformis of the western Canary Islands cluster with Nesotes altivagans and Nesotes elliptipennis from Tenerife. Fifty‐two individuals from this western islands species complex have been sequenced for 675 base pairs of the mtDNA cytochrome oxidase II gene, representing Tenerife, La Palma and El Hierro. A neighbour joining analysis of maximum likelihood distances resulted in three distinct mtDNA lineages for N. conformis, two of which also include mitotypes of N. altivagans and N. elliptipennis. Through application of parametric bootstrap tests, we are able to reject hypotheses of monophyly for both N. conformis and N. altivagans. Nesotes altivagans and N. elliptipennis are poorly separated morphologically and mtDNA sequence data adds support to this being one species with a highly variable morphology. We propose that N. altivagans/N. elliptipennis is recently derived from two ancestral mtDNA lineages within N. conformis from the Teno region of Tenerife. We further propose colonization of the younger islands of La Palma and El Hierro by N. conformis from a mitochondrial lineage within the Teno massif (colonization; diversification; mitochondrial DNA; Canary Islands; Coleoptera).     

Molecular systematics and evolution of the Ptinidae (Coleoptera: Bostrichoidea) and related families

The Ptinidae (Coleoptera: Bostrichoidea) are a cosmopolitan, ecologically diverse, but poorly known group of Coleoptera and, excluding a few economic pests, species are rarely encountered. This first broad phylogenetic study of the Ptinidae s.l. (i.e. including both the spider beetles and anobiids) examines relationships based on DNA sequence data from two mitochondrial genes (16S and COI) and one nuclear gene (28S), using out‐group taxa from both the Bostrichidae and Dermestidae. Topologies varied depending on the genes used and whether data were analysed with either parsimony or Bayesian methods. Generally the two mitochondrial genes supported relationships near the tips of the phylogeny, whereas the nuclear gene supported the basal relationships. The monophyly of the Ptinidae was not inferred by all of the gene combinations and analysis methods, although the combined Ptinidae and Bostrichidae have a single origin in all cases. Alternative relationships include the Ptinidae s.s. (i.e. Ptininae and Gibbiinae) as sister to the anobiids (i.e. the nine remaining subfamilies of Ptinidae s.l.) + Bostrichidae, or the Bostrichidae as sister to the Ptinidae s.s.+ anobiids. Most of the larger subfamilies within the Ptinidae are not monophyletic. Further analysis with more taxa and more genes will be required to clarify and decide upon the best hypothesis of relationships found within the clades of the Bostrichidae and Ptinidae. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 88–108.     

Phylogeography and molecular phylogeny of Macaronesian island Tarphius (Coleoptera: Zopheridae): why are there so few species in the Azores?

Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra‐island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra‐island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra‐island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species‐rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature.     

Molecular phylogeny and character evolution in the Western Palaearctic Helicidae s.l. (Gastropoda: Stylommatophora)

In this study, we present a molecular phylogeny for the west Palaearctic Helicidae sensu lato based on sequence data from two mitochondrial (COI, 16S rDNA) and two nuclear (ITS-1, 18S rDNA) genes. Maximum likelihood analysis and Bayesian inference revealed well supported monophyletic clades partly conflicting traditional classifications. Based on these results, we propose the following system. The Western Palaearctic Helicidae s.l. consist of two families, Helicidae and Hygromiidae. Within the Helicidae, three well supported subfamilies can be recognised: the Helicinae, Ariantinae, and Helicodontinae. The Hygromiidae consist of three clades: the Hygromiinae, the Helicellinae, and a yet unnamed clade comprising the genera Sphincterochila and Cochlicella. We then used the phylogeny to study the evolution of anatomical, and ecological characters traditionally used for systematic classification. In the Helicidae s.l., two independent evolutionary transitions to life in xeric environments occurred, which allowed the occupation of new niches with a subsequent radiation of the Helicellinae-Cochlicella/Sphincterochila clade and the Helicinae. Whereas, the multiplication of the Glandulae mucosae is a synapomorphy of the Hygromiidae, the lovedart sac apparatus is present in all groups and thus, the trait cannot provide a synapomorphy for either families or subfamilies. Additionally, we evaluated the use of structural molecular genetic characters for taxonomic assessment. The presence of an unique loop region of the 16S rDNA gene and a short tandem repeat in the ITS-1 region provide independent evidence for the monophyly of these major two groups, and can be used for preliminary classification.     

Molecular phylogeny and zoogeography of the freshwater crayfish genus Cherax Erichson (Decapoda: Parastacidae) in Australia

The evolutionary history and biogeography of freshwater-dependent taxa in Australia is of intrinsic interest given the present-day aridity of this continent. Cherax is the most widespread and one of the most species-rich of Australia's nine freshwater crayfish genera. The phylogenetic relationships amongst 19 of the 23 Australian Cherax were established from mitochondrial DNA sequences representing the 12S rRNA and 16S rRNA gene regions. The relationships among species support an initial east–west separation, followed by a north–south divergence in eastern Australia. Molecular clock estimations suggest that these divergences date back to the Miocene. The phylogenetic relationships support endemic speciation within geographical regions and indicate that long-distance dispersal has not led to recent speciation as previously hypothesized. This new evolutionary scenario is consistent with the climatic history of Australia and the evolutionary history of other similarly distributed freshwater-dependent organisms in Australia.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 553–563.     

Molecular phylogeny and DNA barcoding confirm cryptic species in the African freshwater oyster Etheria elliptica Lamarck, 1807 (Bivalvia: Etheriidae)

Recent molecular approaches to taxonomy have led to a steady increase in the identification of cryptic species. Within the Etheriidae, the species Etheria elliptica (freshwater oyster) is widespread and common and exists in most of the major African drainages. Within the African freshwater ecosystems, there are major threats to biodiversity and cryptic species complicate conservation strategies; unknown species exist and no conservation status has been assigned. Our objective here was to determine if E. elliptica from several locations in the Congo drainage are correctly classified as representing a single species. We analysed the genetic diversity at two mitochondrial loci (COI and 16S) and two nuclear loci (H3 and 28S), and estimated evolutionary relationships using phylogenetic and DNA barcoding techniques. Bayesian inference yielded three cryptic species of Etheria, and mismatch analysis revealed discrete differences between the cryptic species. We identified three cryptic species within these collections, and evidence indicates that the third species may resolve further with more sampling. In conclusion, the taxonomic history of E. elliptica makes finding cryptic species unsurprising. However, molecular studies such as this may finally help to resolve the number of species within this genus.     

Molecular and morphometric assessment of the taxonomic status of Ectemnorhinus weevil species (Coleoptera: Curculionidae, Entiminae) from the sub-Antarctic Prince Edward Islands

There are long‐standing controversies on the taxonomic status of Ectemnorhinus weevil species occurring on the sub‐Antarctic Prince Edward Islands. Since the two islands that constitute the Prince Edward Islands archipelago (PEIA), Marion Island (MI) and Prince Edward Island (PEI), differ in terms of alien invasive species such as the introduced house mouse Mus musculus and conservation management strategies, it is important to consider inter‐island dynamics when investigating inter‐specific relationships. Using a combined molecular phylogenetic and morphometric approach, we attempted to resolve the taxonomic status of the PEIA Ectemnorhinus weevil species. A COI gene phylogeny was inferred following the genetic characterization of 52 Ectemnorhinus weevils from both islands, and morphometric assessment using a set of 15 linear, external measurements was used to differentiate between the two currently recognized species, Ectemnorhinus similis and Ectemnorhinus marioni. Analyses revealed the presence of two genetically and morphometrically distinct species on PEI, whilst evidence for a single species, comprising diverse genetically discrete populations was found on MI. Based on these results, the species unique to PEI has been designated Ectemnorhinus kuscheli n. sp. whilst we confirm the synonymy between E. similis and E. marioni, the two species originally described from MI. E. kuscheli appears to be restricted to PEI, whereas E. similis occurs on both MI and PEI.     

Towards the phylogeny of the Curculionoidea (Coleoptera): Reconstructions from mitochondrial and nuclear ribosomal DNA sequences

With about 60,000 described species, Curculionoidea represent the most species-rich superfamily in the animal kingdom. The immense diversity apparently creates difficulties in the reconstruction of the phylogenetic relationships. Independent morphological studies have led to very different classifications. This study is based on molecular data from two independent molecular sources, the 16S and 18S rDNA. Sensitivity analyses were conducted for the sequence alignment (gap costs were varied) as well as the phylogenetic reconstruction algorithms and some of their parameters. The higher-level relationships reconstructed within Curculionoidea are sensitive to alignment and reconstruction method. Nemonychidae or Oxycorynidae+Belidae were found to be sister to all remaining Curculionoidea in many analyses. The 16S rDNA sequence data (obtained from 157 species) corroborate many tribes and genera as monophyletic. It is observed that the phylogenetic reconstruction of genera with specific genetic features such as polyploidy and parthenogenetic reproduction is difficult in weevils. The curculionid subfamily Lixinae appears monophyletic. A new monophylum consisting of Entiminae, Hyperinae, Cyclominae, Myllorhinus plus possibly the Cossoninae is distinguished and we call it Entiminae s.l. For most other subfamilies and families homoplasy concealed the phylogenetic signal (due to saturation of the 16S sequences), or the species sampling was insufficient, although our sampling scheme was rather broad. We observed that although data from one source can easily be misleading (16S) or hardly informative (18S), the combination of the two independent data sets can result in useful information for such a speciose group of organisms. Our study represents the most thorough analysis of molecular sequence data of the Curculionoidea to date and although the phylogenetic results appear less stable than expected, they reflect the information content of these sequence data realistically and thus contribute to the total knowledge about the phylogeny of the Curculionoidea.