Evolution of host- and habitat association in the wood-feeding cockroach, Cryptocercus

Members of the cockroach genus Cryptocercus are subsocial insects that live in temperate forests and feed on decomposing logs. At present, seven species are recognized worldwide: four in the eastern USA, one in the western USA, and one each in Russia and China. Genetic variation within and among the Nearctic species has been characterized extensively in previous studies. However, whether there has been a corresponding divergence in the host and habitat association of Cryptocercus species is not known. Here, we report on differences in host and habitat association among six of the seven Cryptocercus species, estimated from field observations, elevation data, and land cover data. Our results indicated that the eastern and western USA species differ from one another in their distribution patterns, abundance, and habitat association. The eastern USA species are associated largely with deciduous forests, whereas the western USA and the Russian species are associated with evergreen forests. Thus, the eastern USA species, which are evolutionarily the most recent ones, have adapted to a different set of tree species relative to the basal species. There were also differences in the habitat association of the various species. Specifically, in the eastern USA, Cryptocercus darwini , evolutionarily the most recent species, occupied a habitat that is predominantly at low elevation [<400 m above sea level (ASL)] while all the other Nearctic species and the Russian species occupied a habitat that is at relatively higher elevations (>400 m ASL). Mapping of the above traits on a phylogenetic tree revealed that the evolutionary trend in Cryptocercus with regard to host and habitat association has been toward the utilization of low elevation habitats dominated by deciduous forests. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 163–172.     

Uncovering tropical diversity: six sympatric cryptic species of Blepharoneura (Diptera: Tephritidae) in flowers of Gurania spinulosa (Cucurbitaceae) in eastern Ecuador

Diversification of phytophagous insects is often associated with changes in the use of host taxa and host parts. We focus on a group of newly discovered Neotropical tephritids in the genus Blepharoneura , and report the discovery of an extraordinary number of sympatric, morphologically cryptic species, all feeding as larvae on calyces of flowers of a single functionally dioecious and highly sexually dimorphic host species ( Gurania spinulosa ) in eastern Ecuador. Molecular analyses of the mitochondrial cytochrome oxidase-I gene from flies reared from flowers of G. spinulosa reveal six distinct haplotype groups that differ by 7.2–10.1% bp (uncorrected pairwise distances; N  = 624 bp). Haplotype groups correspond to six distinct and well-supported clades. Members of five clades specialize on the calyces of flowers of a particular sex: three clades comprise male flower specialists; two clades comprise female flower specialists; the sixth clade comprises generalists reared from male and female flowers. The six clades occupy significantly different morphological spaces defined by wing pigmentation patterns; however, diagnostic morphological characters were not discovered. Behavioural observations suggest specific courtship behaviours may play a role in maintaining reproductive isolation among sympatric species. Journal compilation  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 779–797. No claim to original US government works.     

Species' diversity in the New Caledonian endemic genera Cephalidiosus and Nobarnus (Insecta: Heteroptera: Tingidae), an approach using phylogeny and species' distribution modelling

The patterns of local endemism in New Caledonia were analysed in two endemic genera of Tingidae (Insecta, Heteroptera), Cephalidiosus and Nobarnus , through a phylogenetic analysis and species' distribution modelling. The aim was to determine the possible causes of diversification and endemism in New Caledonia. Our results show that environmental conditions are probably important for the distribution of the genus Cephalidiosus , in conjunction with other factors such as resource (host plant) distribution, but suggest that the same environmental conditions have not influenced the speciation processes and diversification in the genus.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 177–184.     

Convergence and parallelism among cicadas of Argentina and the southwestern United States (Hemiptera: Cicadoidea)

Cicadas of the genera Derotettix in Argentina and Okanagodes in the south-western United States resemble each other in colour, live on salt-tolerant plants ( Atriplex spp. etc.), nearly match the colour of their respective host plants and produce songs above the range of avian hearing. The Argentine cicadas are smaller, but have nearly identical thermal limits for activity measured by the minimum temperature for flight (20–24 °C) and a body temperature at heat torpor (48–49 °C). The species shift activity from basking sites to shade at temperatures above 37 °C, although O .  gracilis rises to a significantly higher temperature (40.7 °C) than its congener (38.2 °C) or Derotettix (37.2 °C). The thermal tolerances are the highest reported for cicada species. A third group using halophytes in Argentina ( Babras sonorivox ) has similar temperature tolerances and is cryptically coloured. The genera are convergent with respect to morphology, coloration, body size, behaviour, habitat choice and host plant selection. The similarities of thermal tolerances and their influence on behaviour can be viewed as parallelism because the underlying mechanisms are the same in all species studied.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 281–288.     

A new species of Clivia (Amaryllidaceae) endemic to the Pondoland Centre of Endemism, South Africa

Clivia robusta B.G. Murray, Ran, de Lange, Hammett, Truter et Swanevelder sp. nov. (Amaryllidaceae) is a tubular, pendulous-flowered Clivia species, restricted to the Pondoland Centre of Endemism, South Africa. The unique morphology, distribution, karyotype and molecular fingerprint distinguish it from all other pendulous-flowered species in the genus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 369–374.     

Phylogenetic analysis of Blattabacterium,endosymbiotic bacteria from the wood roach,Cryptocercus (Blattodea: Cryptocercidae), including a description of three new species

Members of the cockroach genus Cryptocercus are wood-feeding, subsocial insects that live in temperate forests of the Nearctic and Palaearctic. At present, nine species are recognized: Cryptocercus relictus and Cryptocercus kyebangensis in eastern Asia and Russia, Cryptocercus primarius and Cryptocercus matilei in southwestern China, Cryptocercus clevelandi in the western USA, and Cryptocercus darwini, Cryptocercus garciai, Cryptocercus punctulatus, and Cryptocercus wrighti in the eastern USA. Like all extant cockroaches, Cryptocercus harbor endosymbiotic bacteria, Blattabacterium, in their fat bodies. The endosymbionts in all cockroaches have been considered a single species, Blattabacterium cuenoti, since their discovery about a century ago. However, a recent analysis of DNA sequences from representatives of four cockroach families has indicated that there is considerable DNA sequence divergence among B. cuenoti from different host species. As a part of our studies on the evolution of Cryptocercus, we examined DNA sequence divergence among B. cuenoti from six of the nine known Cryptocercus species. Specifically, we sequenced approximately 2,400 bp of the 16S rRNA and 23S rRNA genes of B. cuenoti from six species of Cryptocercus. We found that B. cuenoti in Cryptocercus has differentiated into multiple monophyletic lineages distinguishable by DNA sequence of rRNA genes and host association. Our sequence divergence estimates were consistent with those reported for other, congeneric bacterial species. We propose the recognition of three new species of Blattabacterium within Cryptocercus species as follows: Blattabacterium relictus sp. nov. in C. relictus, Blattabacterium clevelandi sp. nov. in C. clevelandi, and Blattabacterium punctulatus sp. nov. in C. darwini, C. garciai, C. punctulatus, and C. wrighti.     

Historical influences on genital morphology among sympatric species: gonopod evolution and reproductive isolation in the crayfish genus Orconectes (Cambaridae)

Freshwater crayfishes represent an incredibly diverse component of temperate aquatic ecosystems. The genus Orconectes (Cambaridae) comprises approximately 17% of total global crayfish diversity and is native to central and eastern North America. Using both Bayesian and parsimony-based phylogenetic hypotheses from complete cytochrome c oxidase subunit I (COI) mtDNA sequence data (1545 bp) along with information on male gonopod shape and species distribution patterns, we examined the evolution of male genital morphology and its potential influence on species co-occurrence. We found that sympatric species exhibited similar male genital morphology more often than expected based on the frequency of genital shapes within the genus. When evolutionary history was incorporated into this analysis, the result was no longer significant, suggesting that species co-occurrence and the associated genital shapes of sympatric congeners cannot be explained solely by interspecific ecological interactions. Our results also do not support the current subgeneric classifications within Orconectes or the monophyly of the genus, both of which have been based previously on male genital morphology.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 1–12.     

Two new species of Multiclavula (lichenized basidiomycetes) from savanna soils in Rwanda (East Africa)

Two new species of Multiclavula are described from Rwanda: M. akagerae , with a thallus consisting of turgescent glomerules, and M. rugaramae , with a thallus composed of flattened, rounded, marginate, and dispersed squamules. They grow on soil in frequently burned savannas or on lateritic soils in eastern Rwanda. The genus Lepidostroma should probably be reduced into synonymy with Multiclavula .  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 457–465.     

Lineage specific evolution of an alternative social strategy in Tetramorium ants (Hymenoptera: Formicidae)

The western Palearctic ant, Tetramorium moravicum , is the only species of the large genus known to display two alternative social strategies: macrogyny-monoygyny (characterized by one large queen per colony) and microgyny-polygyny (multiple small queens per colony). The microgyne colonies of this ant are confined to the westernmost part of its distribution area and occasionally co-occur with macrogyne colonies. We investigated phylogenetic and geographical patterns in relation to social strategy by analysing 1031 bp of the mitochondrial cytochrome c oxidase subunit I gene of T. moravicum from the entire known species range. Phylogenetic analyses revealed two clear-cut lineages coinciding with the eastern and western populations, which probably were separated after the late Quaternary glacial periods. Geographical distance analysis suggested allopatric fragmentation of the two lineages. We hypothesize that the two lineages colonized Central Europe postglacially from two directions. The current distribution gap of 500 km between the tip of the eastern lineage, in the Czech Republic, Austria and Hungary, and the western lineage, in Germany, was confirmed by hierarchical population analysis. Macrogyny-monogyny probably was the original condition in T. moravicum ; the alternative strategy of microgyny-polygyny may have evolved at the tip of the western lineage, as recently as the early Holocene.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 247–255.     

Uniformity of karyotypes in Ligularia (Asteraceae: Senecioneae), a highly diversified genus of the eastern Qinghai–Tibet Plateau highlands and adjacent areas

Ligularia , a highly diversified genus in the eastern Qinghai–Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x  = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2 n  = 58 and 87. Other species are only diploid, with 2 n  = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia , are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium , Ligulariopsis , Parasenecio , and Sinacalia . Aneuploid reduction of chromosome number from 2 n  = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 329–342.     

Species: towards new, well-grounded practices

Velasco recently criticized our formal definition of the species concept, arguing for its inappropriateness both in fundamental and practical aspects [Velasco JD (2008) Biological Journal of the Linnean Society , 93 , 865–869]. Here, we clarify some misunderstandings that are at the basis of Velasco's paper. First, we show why and how the definition of the species concept can be grounded in the theory of evolution and what that implies. Then, we explain why Velasco's formal criticisms are unjustified. Finally, we point out the practical and methodological consequences of a rigorous conceptual framework for species study, and we show that today's development of species delimitation methods fully agrees with our proposal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 217–222.     

Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodontidae) and its evolutionary and biogeographical implications

The largest genus of salamanders, Bolitoglossa (Plethodontidae), is widespread in tropical America, where it occurs in diverse habitats and elevations, from high elevation grasslands to lowland rain forest . It has the most extensive geographical range of any salamander genus. While most species occur in Middle America, it ranges throughout most of tropical South America as well. Phylogenetic analysis of 1196 bp of two mitochondrial genes (cytochrome b , 16S RNA) from 55 species offers strong support for the monophyly of the genus and sorts the species into a number of clades. Taking into account morphology, distribution, general ecology, and prior systematic and taxonomic studies, we recognize seven subgenera, four of them new: Bolitoglossa Duméril, Bibron et Duméril, 1854, Eladinea Miranda Ribeiro, 1937, Magnadigita Taylor, 1944, Mayamandra , Nanotriton , Oaxakia and Pachymandra . All South American and some lower Middle American species are included in a single well -supported clade, Eladinea . At the species level our analyses uncover the existence of large genetic diversity within morphologically homogeneous taxa. We propose the new combination: B. (Eladinea) paraensis (Unterstein, 1930) stat. nov. , for Brazilian salamanders previously included under B. altamazonica . We evaluate evidence for the multiple colonization of the tropical lowlands by morphologically derived species groups. South America was invaded by members of one clade, Eladinea , which we infer to have dispersed to South America prior to closure of the Panamanian Portal. Despite the relatively long history of salamanders in South America, that continent now accounts for a relatively small proportion of the lineages and species of neotropical salamanders.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 325–346.     

Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s . l ., Argynnina comprising Prokuekenthaliella , Issoria , Brenthis , and all the 'large fritillary' species joined in the genus Argynnis s . l . and a new subtribe Euptoietina comprising only the genus Euptoieta . The classical genus Issoria s . l . is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella . Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas , Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 627–673.     

Arbuscular mycorrhizal fungi: genetics of multigenomic, clonal networks and its ecological consequences

Arbuscular mycorrhizal fungi are thought to have remained asexual for 400 million years, although recent studies have suggested that considerable genetic and phenotypic variation could potentially exist in populations. A brief discussion of these multigenomic organisms is presented.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 59–60.     

Host plant toxicity affects developmental rates in a polyphagous fruit fly: experimental evidence

Various, non-exclusive mechanisms have been invoked to explain the observed association between host plant use and speciation in insect species. In the Afrotropical genus Ceratitis (Diptera: Tephritidae), morphological, molecular, and biochemical data suggest that evolutionary radiation of stenophagous clades originates from their ability to exploit toxic hosts. To test whether, and to what extent, the development and fitness of Ceratitis fasciventris , a polyphagous congener known to infest over 43 host species, is adversely affected by host plant toxicity, we compared the rates of development, survival, and reproduction of captive bred individuals in four media that differ in alkaloid concentration. Despite reduced pupal and adult sizes, C. fasciventris larvae developing under low alkaloid concentrations successfully developed to the adult stage, probably as a result of accelerated pupation and ensuing restricted exposure to the toxic environment. High alkaloid concentrations, however, impaired their developmental process and prevented subsequent reproduction. The adverse effects of host plant toxicity on larval development in polyphagous fruit flies indicate that high alkaloid concentrations pose a significant constraint on host use by polyphagous Ceratitis species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 728–737.     

Genetic cohesion of Eresus walckenaeri (Araneae, Eresidae) in the eastern Mediterranean

The eresid spider genus Eresus is morphologically and ecologically conservative. At least three species occur in Europe. However, deep genetic divergence among geographical samples within two species, E. cinnaberinus and E. sandaliatus , may suggest more cryptic species. In the present study we investigate the genetic cohesion of the third species, Eresus walckenaeri , throughout its eastern Mediterranean distribution range, relative to the E. cinnaberinus–E. sandaliatus species complex. Eresus walckenaeri specimens were monophyletic. Genetic discreteness of E. walckenaeri in a region of sympatry with its sister species in Greece provides evidence for species integrity of E. walckenaeri within the European Eresus species complex. Eresus walckenaeri exhibited high concordance between geographical location and mtDNA genealogy. Two major phylogeographical clades were found in the Greek–Turkish and Syrian–Israel parts of the investigated area, respectively (∼6.5% sequence divergence). Concordance between geography and genetic divergence was further observed between Aegean island samples and their corresponding Greek and Turkish mainland samples, suggesting regional subdivision with gradual but potentially high dispersal propensity. Monophyly and limited regional distribution indicate Mediterranean endemic origin.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 1–9.     

Host specialization by Cotesia wasps (Hymenoptera: Braconidae) parasitizing species-rich Melitaeini (Lepidoptera: Nymphalidae) communities in north-eastern Spain

In order to investigate parasitoids of the genus Cotesia (Hymenoptera: Braconidae), larvae of a speciose group of butterflies, the tribe Melitaeini (Lepidoptera: Nymphalidae), were collected from several sites in Catalonia, northern Spain, a region that harbours ten out of the 20 European species of Melitaeini. New information on the natural history of the butterflies is presented, and the structure of their communities and patterns of larval parasitism are described. On the basis of mtDNA sequence data (COI gene), microsatellite data (ten loci) and behavioural experiments, we recognize seven biologically distinct species of Cotesia parasitizing the Melitaeini communities within this relatively small geographical area. In particular, the notional species C. melitaearum and C. acuminata each represents a series of cryptic species with narrow host associations. The possibility of direct competition among the parasitoids and/or indirect interactions between butterflies mediated by Cotesia parasitoids is explored.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 45–65.     

Landmark superimposition for taxonomic identification

The identification of organisms is a time-consuming task even for highly specialized researchers. Many ways to accelerate the identification process have been devised – from pictorial keys to automatic, machine identification – with different degrees of success. This paper explores landmark configurations as an aid to taxonomic identification. The basic hypothesis rests on the analogy between human fingerprints and organism landmark configuration. Translated into biological terms, it asks whether individual landmark configurations can be used as diagnostic characteristics for species identification. Water mites of the genus Torrenticola were used as test organisms. The results show that identification can be made simpler through the use of landmark configurations.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 267–274.