Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae)

Faille, A., Casale, A. & Ribera, I. (2010). Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae). —Zoologica Scripta, 40, 282–295. Carabid beetles of tribe Trechini (Coleoptera) are one of the main groups of insects that colonized the subterranean environment. Many species of this group have developed similar morphological modifications related to the subterranean life, resulting in a characteristic Aphaenops‐like phenotype that obscures their phylogenetic relationships (depigmented, blind, elongated body and appendages, narrow head and pronotum). We present here the result of a molecular study using a combination of nuclear (small ribosomal unit, large ribosomal unit) and mitochondrial (cox1, cyb, rrnL, trnL, nad1) genes to investigate the phylogenetic placement of the highly modified subterranean genera of the tribe Trechini from the west Mediterranean area (France, Spain, Morocco and Sardinia). Our results confirm the multiple independent origin of troglomorphism among these genera, and reveal a pattern largely determined by geographical proximity. We discuss the validity of some groups proposed on the base of morphological features, and provide estimates of divergence between subterranean genera and other groups of Trechini, including epigean species of the same area. We compare the estimated age for the origin of the main groups resulting from two different calibrations, using one the standard mitochondrial mutation rate (2.3% divergence per Myr) and the other the separation between Sardinia and mainland 33 Ma. Under the first scenario, the main groups of genera would have a late Miocene origin, with a subsequent colonization of north Africa at the Pliocene–Pleistocene boundary. The assumption that the main groups originated through vicariance due to the separation of the Sardinian plate in the Oligocene results in a Messinian origin of the north African subterranean taxa, and a global mitochondrial rate reduced to 1% divergence per Myr.     

Two new Typhloreicheia species from Sardinia and their biogeographical significance (Coleoptera, Carabidae, Scaritinae)

Typhloreicheia monachasp. n. and Typhloreicheia ilianaesp. n. are described from two caves of Central-Eastern Sardinia (Nuoro province): the Bue Marino cave and the Nurra 'e Pradu cave, respectively. Both caves are located in the part of the island where many highly specialised subterranean carabid beetles are localised. Typhloreicheia monacha is apparently related to two other species of the same area, i.e. Typhloreicheia onnisi Casale & Magrini, 2004 and Typhloreicheia elegans (Dodero, 1916); Typhloreicheia ilianae is closely related to Typhloreicheia henroti Jeannel, 1957, known from a cave near Dorgali. Relationships and diagnostic features among these taxa are discussed and illustrated, and a key for identification of the specialised subterranean Typhloreicheia species of Sardinia is provided. The hypothesis of adaptive radiation of Reicheiina species in Sardinia, recently proposed by the senior author of this contribution, is further elaborated in light of new data.     

A molecular phylogeny shows the single origin of the Pyrenean subterranean Trechini ground beetles (Coleoptera: Carabidae)

Trechini ground beetles include some of the most spectacular radiations of cave and endogean Coleoptera, but the origin of the subterranean taxa and their typical morphological adaptations (loss of eyes and wings, depigmentation, elongation of body and appendages) have never been studied in a formal phylogenetic framework. We provide here a molecular phylogeny of the Pyrenean subterranean Trechini based on a combination of mitochondrial (cox1, cyb, rrnL, tRNA-Leu, nad1) and nuclear (SSU, LSU) markers of 102 specimens of 90 species. We found all Pyrenean highly modified subterranean taxa to be monophyletic, to the exclusion of all epigean and all subterranean species from other geographical areas (Cantabrian and Iberian mountains, Alps). Within the Pyrenean subterranean clade the three genera (Geotrechus, Aphaenops and Hydraphaenops) were polyphyletic, indicating multiple origins of their special adaptations to different ways of life (endogean, troglobitic or living in deep fissures). Diversification followed a geographical pattern, with two main clades in the western and central-eastern Pyrenees respectively, and several smaller lineages of more restricted range. Based on a Bayesian relaxed-clock approach, and using as an approximation a standard mitochondrial mutation rate of 2.3% MY, we estimate the origin of the subterranean clade at ca. 10 MY. Cladogenetic events in the Pliocene and Pleistocene were almost exclusively within the same geographical area and involving species of the same morphological type.     

Descriptions of the first semi-aphenopsian troglobiotic Patrobini and of a new anophthalmic cave-dwelling Trechini from central Sichuan,China (Coleoptera: Caraboidea)

Summary

The first known troglobiotic species belonging to the ground beetle tribe Patrobini is reported from a limestone cave called Lianhua Dong in Pengzhou, a suburb county-level city in northern Chengdu, Sichuan Province, China. Troglopatrobus zhouchaoi n. gen., n. sp., is an eyeless and large-sized patrobine beetle with extremely elongated body (esp. head and elytra) and appendages. Duvalioblemus (Shublemus) liyuani n. subgen., n. sp., a minute anophthalmic trechine species, is also discovered from the same cave. It is the first cave-dwelling species belonging to this genus already known as endogean from Sichuan and Yunnan provinces.     

Phylogeography and molecular rates of subterranean aquatic Stenasellid Isopods with a peri-Tyrrhenian distribution

The subterranean Isopods belonging to the genus Stenasellus have an interesting disjunct distribution in the peri-Tyrrhenian area with morphologically closely related taxa occurring in Sardinia, Corsica, Tuscany and in the Pyrenees phreatic and interstitial waters. Because the dispersal capacities of these organisms are limited, their distribution has been associated traditionally with the tectonic events leading to the separation of the Sardinia-Corsica microplate from the Pyrenees and its subsequent movement towards the Italian peninsula. We sequenced a fragment of the mtDNA cytochrome oxidase I gene (COI) for multiple populations of the S. racovitzai species-group (Corsica, Sardinia, Tuscany) and S. virei (Pyrenees). We found that multiple phylogenetic analyses always gave the same topology, which is consistent with the genetic relations found using allozyme data, and with the palaeogeography of the area. The molecular data suggest that a combination of vicariance and dispersal events explain most effectively the present distribution pattern of these organisms. We also calculated COI rates and calibrated them against absolute time, taking advantage of the availability of two geologically based time estimates. Rates on all substitutions are similar to those published for other crustaceans for the same COI fragment, including taxonomically and ecologically distant groups. Rates on third codon positions or on transversions are generally lower than those found in other crustaceans.     

Mitochondrial DNA rates and biogeography in European newts (genus Euproctus)

Sequence divergence for segments of three mitochondrial DNA (mtDNA) genes encoding the 12S and 16S ribosomal RNA and cytochrome b was examined in newts belonging to the genus Euproctus (E. asper, E. montanus, E. platycephalus) and in three other species belonging to the same family (Salamandridae), Triturus carnifex, T. vulgaris, and Pleurodeles waltl. The three Euproctus species occur (one species each) in Corsica, Sardinia, and the Pyrenees. This vicariant distribution is believed to have been determined by the disjunction and rotation of the Sardinia-Corsica microplate from the Pyreneean region. Because time estimates are available for the tectonic events that led to the separation of the three landmasses, we used sequence data to estimate rates of evolution for the three gene fragments and investigated whether they conform to the rate-constancy hypothesis. By the Tajima (1993, Genetics 135:599-607) test, we could not detect rate heterogeneities for the ribosomal genes and for transversions in the cytochrome b gene. Assuming that these sites are evolving linearly over time and that cessation of gene flow occurred simultaneously with vicariant events, we compared the time of divergence estimated by molecular distances with the divergence times based on the geological estimates. Because we have two estimates of divergence time from the geological record, the split of Corsica/Sardinia from the Pyrenees and the split of Corsica from Sardinia, we could compare ratios of molecular divergence with the ratio of geological time divergence. The ratios are very similar, indicating that the molecular clock hypothesis cannot be rejected. These geological events also allowed us to calculate absolute rates of evolution for ribosomal and cytochrome b genes and compare them to rates for the same regions in other salamandrids and other vertebrates. Ribosomal mtDNA rates are comparable to those reported for other vertebrates, but cytochrome b rates are 3-7 times lower in salamanders than in other ectotherms. From a phylogenetic perspective, our data suggest that the cladogenic events leading to species formation in Euproctus and Triturus occurred very closely in time, indicating that the two genera may not be monophyletic. A duplication of the cytochrome b gene in T. carnifex was found, and the implications of this finding for mtDNA phylogenetic studies are discussed.     

A remarkably modified species of the tribe Platynini (Coleoptera,Carabidae) from a limestone cave in Jiangxi Province,eastern China

Morimotoidius zhushandong sp. n. is described and illustrated from a limestone cave called Zhushan Dong II in Wanzhai Xian (=County) of western Jiangxi Province, eastern China. This species is the most modified species within the tribe Platynini in China by having very slender body and appendages, extremely elongated head, and especially, narrowed and barrel-liked pronotum which is as wide as head. However, the above derived characters are autapomorphies to adapt the subterranean environment. M. zhushandong sp. n. must be a troglobite though it has well pigmented body and flat eyes.     

Molecular phylogeography of Dolichopoda cave crickets (Orthoptera, Rhaphidophoridae): a scenario suggested by mitochondrial DNA

This study focuses on the phylogenetic relationships among a number of West-Mediterranean cave crickets species belonging to Dolichopoda; primarily a Mediterranean genus, distributed from eastern Pyrenees to Caucasus. In this paper, 11 Dolichopoda species from the French Pyrenees (D. linderi), the island of Corsica (D. bormansi and D. cyrnensis), and northern, central, and southern Italy (D. ligustica, D. schiavazzii, D. aegilion, D. baccettii, D. laetitiae, D. geniculata, D. capreensis, and D. palpata) were studied. Two more species, one from the Caucasus, D. euxina, and one from Greece, D. remyi, were also included in the analyses, together with more distant species within the same family to be used as outgroups. Fifteen hundred base pairs of mitochondrial DNA, corresponding to the small subunit of the ribosomal RNA (16S rRNA) and to the subunit I of the cytochrome oxidase I (COI), were sequenced in order to clarify the phylogenetic relationships and biogeography of this group of Mediterranean cave crickets. The molecular data are congruent with a phylogeographic pattern; with the geographically close species also the most related ones. Based on mtDNA divergence, the present-day distribution of genetic diversity seems to have been impacted by climatic events due to glacial and interglacial cycles that have characterized the Pleistocene era.     

Understanding the biogeography of a group of earthworms in the Mediterranean basin--the phylogenetic puzzle of Hormogastridae (Clitellata: Oligochaeta)

Traditional earthworm taxonomy is hindered due to their anatomical simplicity and the plasticity of the characteristics often used for diagnosing species. Making phylogenetic inferences based on these characters is more than difficult. In this study we use molecular tools to unravel the phylogeny of the clitellate family Hormogastridae. The family includes species of large to mid-sized earthworms distributed almost exclusively in the western Mediterranean region where they play an important ecological role. We analyzed individuals from 46 locations spanning the Iberian Peninsula to Corsica and Sardinia, representing the four described genera in the family and 20 species. Molecular markers include mitochondrial regions of the cytochrome c oxidase subunit I gene (COI), 16S rRNA and tRNAs for Leu, Ala, and Ser, two nuclear ribosomal genes (nearly complete 18S rRNA and a fragment of 28S rRNA) and two nuclear protein-encoding genes (histones H3 and H4). Analyses of the data using different approaches corroborates monophyly of Hormogastridae, but the genus Hormogaster is paraphyletic and Hormogaster pretiosa appears polyphyletic, stressing the need for taxonomic revisionary work in the family. The genus Vignysa could represent an early offshoot in the family, although the relationships with other genera are uncertain. The genus Hemigastrodrilus is related to the Hormogaster elisae complex and both are found in the Atlantic drainage of the Iberian Peninsula and France. From a biogeographic perspective Corsica and Sardinia include members of two separate hormogastrid lineages. The species located in Corsica and Northern Sardinia are related to Vignysa, whereas Hormogaster pretiosa pretiosa, from Southern Sardinia, is closely related to the Hormogaster species from the NE Iberian Peninsula. A molecular dating of the tree using the separation of the Sardinian microplate as a calibration point (at 33 MY) and assuming a model of vicariance indicates that the diversification of Hormogastridae may be ancient, ranging from 97 to 67 Ma.     

Molecular phylogeny of rootworms and related galerucine beetles (Coleoptera: Chrysomelidae)

The Galerucinae (Coleoptera: Chrysomelidae) sensu stricto (true galerucines) comprise a large assemblage of diverse phytophagous beetles containing over 5000 described species. Together with their sister taxon, the flea beetles, which differ from true galerucines by having the hind femora usually modified for jumping, the Galerucinae sensu lato comprises over 13 000 described species and is the largest natural group within the Chrysomelidae. Unlike the flea beetles, for which robust hierarchical classification schemes have not been erected, an existing taxonomic structure exists for the true galerucines, based mostly on the works of the late John Wilcox. In the most recent taxonomic list of the Galerucinae sensu stricto, five tribes were established comprising 29 sections housing 488 genera. The majority of the diversity within these tribes is found within the tribe Luperini, in which two genera, Monolepta and Diabrotica, are known to contain over 500 described species. Here, we extend the work from previous phylogenetic studies of the Galerucinae by analysing four amplicons from three gene regions (18S and 28S rRNA; COI) representing 249 taxa, providing the largest phylogenetic analysis of this taxon to date. Using two seven‐state RNA models, we combine five maximum likelihood models (RNA + DNA for the rRNAs; three separate DNA models for the COI codon positions) for these partitions and analyse the data under likelihood using Bayesian inference. The results of these two analyses are compared with those from equally weighted parsimony. Instead of choosing the results from one optimality criterion over another, either based on statistical support, tree topology or philosophical predisposition, we elect to draw attention to the similar results produced by all three analyses, illustrating the robustness of the data to these different analytical methods. In general, the results from all three analyses are consistent with each other and previous molecular phylogenetic reconstructions for Galerucinae, except that increased taxon sampling for several groups, namely the tribes Hylaspini and Oidini, has improved the phylogenetic position of these taxa. As with previous analyses, under‐sampled taxa, such as the Old World Metacyclini and all sections of the subtribe Luperina, continue to be unstable, with the few taxa representing these groups fluctuating in their positions based on the implemented optimality criterion. Nonetheless, we report here the most comprehensive phylogenetic estimation for the Galerucinae to date.     

Convergent Adaptations to Cave Life in the Rhamdia Laticauda Catfish Group (Pimelodidae, Teleostei)

The nocturnal Mexican catfish Rhamdia laticauda (Pimelodidae, Teleostei) is the surface sister species of a number of cave species. Comparison between two of them, R. zongolicensis and R. reddelli, reveals an intermediate state of reduction of the biologically functionless eyes, melanin pigmentation and the negative phototactic behavior. The surface species is perfectly adapted to life in darkness. Therefore only few constructive adaptations are developed in the cave species. For survival under conditions of low food supply in the caves the barbels are elongated to improve the senses of taste and touch and more fat can be deposited in the cave fish tissue. Due to convergent evolution the two cave species are morphologically much alike and show only minor diagnostic meristic differences. From geological data it can be concluded that the two cave species started troglobitic evolution at the end of Pleistocene.     

The thoracic morphology of cave-dwelling and free-living ground beetles from China (Coleoptera,Carabidae, Trechinae)

External and internal thoracic structures of two carabid species (Trechini) were examined and documented with different techniques. The study has a main focus on the eyeless cave-dwelling specialist Sinaphaenops wangorum, but detailed information is also provided for a species occurring in cave entrances. The phylogenetic background of the structural features of the thoracic skeletomuscular system was addressed. The thoracic morphology of the examined species was compared to conditions observed in previously studied carabids and non-related subterranean leiodids (Staphylinoidea) in order to identify cave adaptations. Main thoracic character complexes linked with cavernicolous habits in Trechini are elongation of the pro- and mesothorax and the legs, and a complete and irreversible reduction of the flight apparatus. The lost flight capacity is linked with a far reaching modification of skeletal elements of the metathorax including a strongly shortened and simplified metanotum, a shortened metaventrite, and completely reduced wings and sclerites of the wing base. The elongate prothorax together with the long and slender head and elongated legs distinctly increases the activity range in the subterranean lightless environment, which likely facilitates foraging of the carnivorous beetles. Some of the observed features like wing loss and elongation of the anterior thorax and legs are also found in some cave-dwelling Leiodidae (Leptodirini), whereas some other subterranean members of the staphylinoid family have a compact body and legs of normal length. In contrast to the predaceous Trechini, Leptodirini are scavengers.     

Using museum collections to explore predator–prey relationships in snail-eating beetles (Carabidae: Cychrini)

The vast majority of species interactions in nature go unnoticed because they occur under circumstances unamenable to observation. This is unfortunate, as our understanding of trophic ecology is predicated on our ability to quantify the nature and magnitude of species interactions. Here, we use specimens from museums and private collections to estimate prey breadth and morphological patterns between predator and prey pairs of the malacophagous Cychrini beetles collected alongside their snail prey. We identified each pair, measured a series of morphological traits on each, and tested for relationships between the morphological characteristics of beetles and the snails they were found eating. Of 38 specimen pairs, we identified 12 species of Cychrini beetles from two genera (Scaphinotus and Cychrus) eating 22 species of snail prey from 12 genera and ranging from 1 to 9 species of snail prey per beetle species. We found 29 unique predator–prey species pairs. Irrespective of species identity, we found that female beetles were discovered eating larger snails compared to male beetles. We detected two trends in which larger beetles were found eating snails with relatively larger aperture openings, and beetles with more slender body shapes (longer, thinner mandibles, heads, and pronota) were found eating snails whose shells had relatively smaller aperture openings. This suggests that, even within the carabid tribe Cychrini, variation in the cychrine body form may influence prey availability. This study provides the most comprehensive list to date of predator–prey pairs in this understudied group of beetles and also demonstrates the utility of museum collections for documenting cryptic species interactions.     

Phylogenetic relationships of Danio within the order Cypriniformes: a framework for comparative and evolutionary studies of a model species

The evolutionary relationships of species of Danio and the monophyly and phylogenetic placement of the genus within the family Cyprinidae and subfamily Rasborinae provide fundamentally important phyloinformatics necessary for direct evaluations of an array of pertinent questions in modern comparative biology. Although the genus Danio is not one of the most diverse within the family, Danio rerio is one of the most important model species in biology. Many investigations have used this species or presumed close relatives to address specific questions that have lasting impact on the hypothesis and theory of development in vertebrates. Largely lacking from this approach has been a holistic picture of the exact phylogenetic or evolutionary relationships of this species and its close relatives. One thing that has been learned over the previous century is that many organismal attributes (e.g., developmental pathways, ecologies, behaviors, speciation) are historically constrained and their origins and functions are best explained via a phylogenetic approach. Herein, we provide a molecular evaluation of the phylogenetic placement of the model species Danio rerio within the genus Danio and among hypothesized closely related species and genera. Our analysis is derived from data using two nuclear genes (RAG1, rhodopsin) and five mitochondrial genes (ND4, ND4L, ND5, COI, cyt b) evaluated using parsimony, maximum likelihood, and Bayesian analyses. The family Cyprinidae is resolved as monophyletic but the subfamily Rasborinae (priority over Danioinae) is an unnatural assemblage. Danio is identified as a monophyletic group sister to a clade inclusive of the genera Chela, Microrasbora, Devario, and Inlecypris, not Devario nor Esomus as hypothesized in previous studies. Danio rerio is sister to D. kyathit among the species of Danio evaluated in this analysis. Microrasbora and Rasbora are non-monophyletic assemblages; however, Boraras is monophyletic.     

EVOLUTION OF ADULT MORPHOLOGY AND LIFE-HISTORY CHARACTERS IN CAVERNICOLOUS PTOMAPHAGUS BEETLES

Morphological and life-history characters were determined for a series of six increasingly cavernicolous species of eastern United States Ptomaphagus beetles. Contrary to expectations, dimensions of only some adult structural characters uniformly covary as an overall measure of evolutionary adaptation for cave life. Adult reproductive characters that show adaptation for cave life are loss of reproductive seasonality and production of fewer and larger eggs. Significant change in pre-imaginal life-cycle stages was not found. This is in contrast to cavernicolous bathysciine beetles of Europe which show remarkable adaptive trends in pre-imaginal stages, but larval adaptations are not strongly coevolved with adult cave-adaptive characters. This suggests that evolution of cave adaptation in adult endopterygote insects occurs before and independently from that in larvae.     

Anophthalmia and elongation of body appendages in cave scale worms (Annelida: Aphroditiformia)

This study addresses whether cave dwelling annelids exhibited similar reductive and constructive traits equally as strong as those of arthropods and vertebrates inhabiting caves. Known as troglomorphism, these adaptations bring about striking morphologies across invertebrates and vertebrates from both aquatic and terrestrial cave habitats, and include varying degrees of eye and pigmentation loss, as well as hypertrophy of body appendages and sensorial structures. Employing phylogenetic comparative methods and ancestral character reconstructions on a worldwide data set of a group of annelids, the scale worms (Aphroditiformia), we investigate the behavioural and morphological traits of species living in marine caves in comparison with those species living outside caves. Our work demonstrated that cave scale worms respond similar to arthropods in cave environments, showing a significant elongation of sensory parapodial cirri, while lacking eyes and pigmentation. However, whereas elongation of sensory appendages likely occurred in correlation to cave colonization, eyes were plausibly lost in correlation with specialization and colonization of deep‐sea habitats.     

Tectonic vicariance versus Messinian dispersal in western Mediterranean ground beetles

The complex geological history of the western Mediterranean region conceals the interpretation of the evolutionary history of its current fauna, as similar distribution patterns may have very different temporal and geographic origins. Particularly intriguing are some subterranean species in islands, which origin is usually difficult to interpret as their strongly modified morphologies obscure their relationships. We studied subterranean taxa and their likely relatives of two groups of ground beetles in the western Mediterranean: the Duvalius lineage (“isotopic” Trechini) and Molopina (Pterostichini). We included specimens from the islands of Mallorca, Sardinia and Sicily, plus mainland Europe and North Africa. Phylogenetic relationships were reconstructed with a combination of mitochondrial and nuclear data, and divergence dates were estimated with Bayesian methods using the same a priori molecular evolutionary rates for the same gene fragments in the two groups. In the Duvalius lineage, the subgenus Trechopsis, including all the highly modified cave or nivicolous species, was found to be polyphyletic: the species from Mallorca was found to be of Pleistocene origin and sister to the less modified species of subgenus Duvalius from the same island, whereas the Algerian species of Trechopsis were, on the contrary, related to the Sicilian Duvalius, indicating a northern colonisation route during the late Pliocene. Molopina was divided into three main lineages: the genera Abax, Percus, and the Molops groups of genera. The basal diversification of the latter was dated within a temporal window (35–25 Ma) fully congruent with the tectonic opening of the western Mediterranean basin and included six main lineages with uncertain relationships among them: the epigean genera (a) Molops and (b) Tanythrix; and the subterranean (c) Typhlochoromus (Eastern Alps), (d) Speomolops (Sardinia), (e) Henrotius (Mallorca) and (f) a strongly supported clade including the Pyrenean genera Zariquieya, Oscadytes and Molopidius. Despite the similar distribution of some of their subterranean taxa, the two studied groups show a strongly contrasting origin and mode of diversification. While the Duvalius lineage had a recent origin, with complex colonisation patterns and widespread morphological convergence among the subterranean species, the subterranean Molopina had an ancient vicariant origin resulting from the tectonic opening of the western Mediterranean basin.     

First report of the genus Kisaura Ross (Trichoptera, Philopotamidae) from India with the description of six new species

The genus Kisaura Ross, 1956 (Trichoptera, Philopotamidae, Philopotaminae) is reported from India the first time. Six new species are described and illustrated: Kisaura elongatasp. n., Kisaura eloctsp. n., Kisaura clavatasp. n., Kisaura gangtokensissp. n., Kisaura truncatasp. n., all from Gangtok (Sikkim) and Kisaura himachalicasp. n. from Barot (Himachal Pradesh). Male genitalia of this genus are distinguishable from those of other genera of the family by the pair of long lateral processes of tergum X, the well-developed mesoventral plates between two segments of the inferior appendages, and by the brush-like row of dark setae on the inner surfaces of the terminal segments (harpago) of the inferior appendages.     

Species diversity and ecological range in relation to ploidy level in the flora of the Pyrenees

Polyploidy is a major process in plant evolution. Surprisingly, no study has examined its role in species diversification and ecological distribution in relation to other life history traits. In this study, we examine to what extent polyploidy and the other traditionally examined biological traits (pollination mode, dispersal mode and growth form) account for ecological and taxonomic diversity in the flora of the Pyrenees. Fifty genera (in 22 angiosperm families) were classified according to ploidy level, growth form, pollination mode and dispersal mode, and 451 species and/or subspecies in these 50 genera were classified according to ploidy level and growth form. We examined the contribution of ploidy level, pollination and dispersal modes and growth form to (i) the ecological range of species and genera, i.e., the number of natural habitats (defined by a combination of ecological characteristics) where they occur, and (ii) the taxonomic diversity of the 50 genera. Ploidy level and dispersal mode had significant effects on the taxonomic diversity of the 50 genera. Taxonomic diversity, but not polyploidy per se, was significantly correlated with ecological range of genera. For individual species, diploids had a larger ecological range than polyploids, and herbaceous growth forms had wider ecological distributions than other growth forms. Our results indicate that polyploidisation may be a source of ecological diversification of genera, not by increasing the ecological range of particular polyploid species compared to diploids, but rather by creating taxonomic diversity that leads in some genera to a diversification of the habitats occupied by different ploidy levels. This observation is consistent with previous observations of ecological divergence of chromosomal races in some species in the Alps and in the Pyrenees. As found in other studies, species diversification in the studied flora appears to be greatly influenced by the occurrence of multiple dispersal modes, while ecological range of species or subspecies is significantly increased by the presence of herbaceous species.     

Phylogeny of haplo–diploid,fungus‐growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data

Cognato, A. I., Hulcr, J., Dole, S. A. & Jordal, B. H. (2010). Phylogeny of haplo‐diploid, fungus‐growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data. —Zoologica Scripta, 40, 174–186. The ambrosia beetle tribe Xyleborini currently contains 30 genera and approximately 1200 species which are distributed throughout worldwide forests with most diversity located in the tropics. They also represent the most invasive scolytines in North America. Despite economic concerns and biological curiosity with this group, a comprehensive understanding of generic boundaries and the evolutionary relationship among species is lacking. In this study, we include 155 xyleborine species representing 23 genera in parsimony and Bayesian analyses using 3925 nucleotides from mitochondrial (COI) and nuclear genomes (28S, ArgK, CAD, EF‐1α) and 39 morphological characters. The phylogenies resulting from the parsimony analyses, which treated gap positions either as missing or fifth character states, and the Bayesian analysis were generally similar. Clades with high support or posterior probabilities were found in all trees, while those with low support were not recovered by all analyses. Fourteen of the 23 genera were monophyletic although not all relationships among the genera were resolved. We show monophyly of several species groups associated with particular morphological and biological characters and suggest recognition of these groups as genera. Most interesting was the monophyly of South and Central American species representing several genera. This finding suggests recent and fast radiation of xyleborines in the New World accompanied by morphological and biological diversification.