Low dispersing species : population genetic processes in space and time : the genus "Trochulus" (Gastropoda: Hygromiidae) as a case study

RÉSUMÉ Une espèce est rarement composée d'une population unique. Parce que les individus ont des capacités de dispersion limitées et que les paysages sont des mosaïques d'habitats, la plupart des espèces sont plutôt composées de sous-populations connectées par la migration. Cette variation spatiale influence directement la distribution de la variabilité génétique dans et entre les populations. Durant ce travail, nous avons abordé certains des processus populationnels qui ont joué un rôle supposé dans l'apparition de nouvelles espèces au sein du genre Trochulus. Plus précisément, nous avons tenté d'évaluer les impacts respectifs de l'isolement passé (facteurs historiques) et présent (facteurs locaux). Nous avons d'abord pu montrer que les faibles capacités de dispersion des escargots terrestres ont directement influencé leur histoire évolutive à toutes les échelles spatiales et temporelles. En réduisant l'effet homogénéisant de la migration, une faible dispersion maintient dans les populations les traces génétiques d'évènements passés. A l'échelle de la distribution globale de Trochulus villosus, ces traces ont permis de reconstruire une histoire faite d'isolements et d'expansions de populations. En combinant des données génétiques avec une modélisation de la niche climatique passée, il a été possible de proposer un scénario significativement meilleur que toutes les hypothèses alternatives que nous avons testées. A l'échelle locale par contre, l'héritage historique est difficile à distinguer de la dynamique actuelle. Ce fut le cas des lignées mitochondriales du complexe sericeus-hispidus : les deux principales lignées étaient phylogénétiquement éloignées, avaient eu des démographies passées différentes et corrélaient avec des différences morphologiques. D'un autre côté, le flux de gène nucléaire était fort, contredisant l'idée de deux espèces cryptiques isolées reproductivement. Pour pouvoir conclure à la présence ou non de deux espèces, il nous a manqué des informations locales sur la dynamique des populations et les conditions écologiques que l'on trouve dans la région d'étude. Enfin, nous avons pu souligner que la connectivité entre populations d'escargots est soumise à la qualité des habitats et à leur organisation spatiale. Les escargots sont dépendants d'un habitat et s'y adaptent, comme l'indiquent la présence de «poils » uniquement sur la coquille d'espèces vivant dans des habitats humides ou la corrélation entre morphologie et habitat au sein du complexe sericeus-hispidus. Logiquement donc, les escargots migrent préférentiellement au travers d'habitats favorables comme l'a montré la réduction de flux de gènes au travers des prairies chez T. villosus (une espèce forestière). De ces données, nous pouvons supposer que les populations d'escargots en particulier, et des espèces à faible dispersion en général, ont de fortes chances d'être affectées par les changements climatiques, avec de probables implications pour leurs histoires évolutives. SUMMARY : Species rarely consists in a single population. Because individuals have limited dispersal abilities, because landscapes are habitat patchworks, most species are made of several subpopulations connected by migration. This spatial variation has consequences on the distribution of genetic diversity within and between populations, creating a structure among the populations. During the present work, we investigated some of the population processes assumed to have played an important role on the speciation within the genus Trochulus. More specifically, we questioned the respective impacts of past (historical factors) or present (local factors) population isolations. We first could show that the poor dispersal abilities of land snails have had profound impacts on their evolutionary histories at all spatial and temporal scales. Low dispersal maintains a strong signature of past events in the populations by minimising the homogenising effects of geneflow. At the scale of Trochulus villosus global distribution, they allowed to retrieve the detailed history of this species population isolations and expansions. Combining a large genetic dataset with paleo-climatic niche modelling ended up with a historical scenario significantly better than all traditional alternatives we tested. At local scale on the contrary, past events become difficult to tease apart from ongoing processes. This was the case for the divergent mitochondria) lineages within the sericeus-hispidus complex: the two principal lineages appeared to be phylogenetically distant, to have experienced different demographic histories and to correlate with morphological differences. On the other hand, nuclear (present day) geneflow was high, contradicting the idea of two reproductively isolated cryptic species. Information on the local population dynamics and environmental conditions are lacking to be able to decide whether past isolation has indeed resulted here in new species. Finally, we emphasised the importance of the habitat types present in a landscape as well as their spatial organisation for the population connectivity of land snails. These species are tightly dependent on a habitat and adapt to it as shown by thé occurrence of hair-like structures only in species living in humid environments or by the correlation between shell morphology and habitat in the sericeus-hispidus complex. As a result, land snails preferentially migrate through favourable habitats: Trochulus villosus, a forest species, had its geneflow significantly reduced across meadows. From these data, we can hypothesise that the populations of land snails in particular and of low dispersing species in general are likely to be strongly affected by the ongoing climate changes, with potential major consequences on their evolutionary histories.

Thermal energetics of the New-Guinean moss-forest rat (Rattus niobe) in comparison with other tropical murid rodents.

The thermal energetics of rodents from cool, wet tropical highlands are poorly known. Metabolic rate, body temperature and thermal conductance were measured in the moss-forest rat, Rattus niobe (Rodentia), a small murid endemic to the highlands of New Guinea. These data were evaluated in the context of the variation observed in the genus Rattus and among tropical murids. In 7 adult R. niobe, basal metabolic rate (BMR) averaged 53.6±6.6mLO2h(-1), or 103% of the value predicted for a body mass of 42.3±5.8g. Compared to other species of Rattus, R. niobe combines a low body temperature (35.5±0.6°C) and a moderately low minimal wet thermal conductance cmin (5.88±0.7mLO2h(-1)°C(-1), 95% of predicted) with a small size, all of which lead to reduced energy expenditure in a constantly cool environment. The correlations of mean annual rainfall and temperature, altitude and body mass with BMR, body temperature and cmin were analyzed comparatively among tropical Muridae. Neither BMR, nor cmin or body temperature correlated with ambient temperature or altitude. Some of the factors which promote high BMR in higher latitude habitats, such as seasonal exposure to very low temperature and short reproductive season, are lacking in wet montane tropical forests. BMR increased with rainfall, confirming a pattern observed among other assemblages of mammals. This correlation was due to the low BMR of several desert adapted murids, while R. niobe and other species from wet habitats had a moderate BMR.

Revision of the myrmecophilous diapriid genus Bruchopria Kieffer (Hymenoptera, Proctotrupoidea, Diapriidae)

A redescription of the genus Bruchopria and its two species, B. hexatoma Kieffer (female and male) and B. pentatoma Kieffer (male only) is presented. New record for B. hexatoma from Argentina is provided. Some aspects of diapriid biology with emphasis on their myrmecophilic evidence and association with ants are discussed. Scanning photographs of morphological structures are included.

Review of the genus Chalcolepidius Eschscholtz, 1829 (Coleoptera, Elateridae, Agrypninae)

The genus Chalcolepidius is revised. Type specimens of 65 nominal species, except C. costatus Pjatakowa, 1941, C. fleutiauxi Pjatakowa, 1941 and C. viriditarsus Schwarz, 1906, are examined. Eighty five species are studied, of which 34 are synonymyzed and 12 new species described; three species, C. alicii Pjatakowa, 1941, C. haroldi Candèze, 1878 and C. unicus Fleutiaux, 1910, formely included in this genus, are not congeneric and are removed; C. validus Candèze, 1857 is revalidated. The genus is now formed by 63 species. Redescriptions, illustrations and a key for the examined species, and a cladistic analysis for groups of species are also included. New synonyms established: C. apacheanus Casey, 1891 = C. simulans Casey, 1907 syn. nov. = C. acuminatus Casey, 1907 syn. nov. = C. nobilis Casey, 1907 syn. nov.; C. approximatus Erichson, 1841 = C. aztecus Casey, 1907 syn. nov. = C. niger Pjatakowa, 1941 syn. nov.; C. attenuatus Erichson, 1841 = C. cuneatus Champion, 1894 syn. nov. = C. tenuis Champion, 1894 syn. nov.; C. aurulentus Candèze, 1874 = C. candezei Dohrn, 1881 syn. nov. = C. grossheimi Pjatakowa, 1941 syn. nov.; C. bomplandii Guérin, 1844 = C. humboldti Candèze, 1881 syn. nov.; C. chalcantheus Candèze, 1857 = C. violaceous Pjatakowa, 1941 syn. nov.; C. cyaneus Candèze, 1881 = C. scitus Candèze, 1889 syn. nov. = C. abbreviatovittatus Pjatakowa, 1941 syn. nov.; C. desmarestii Chevrolat, 1835 = C. brevicollis Casey, 1907 syn. nov.; C. gossipiatus Guérin, 1844 = C. erichsonii Guérin-Méneville, 1844 syn. nov. = C. lemoinii Candèze, 1857 syn. nov.; C. inops Candèze, 1886 = C. murinus Champion, 1894 syn. nov.; C. jansoni Candèze, 1874 = C. mucronatus Candèze, 1889 syn. nov.; C. lacordairii Candèze, 1857 = C. exquisitus Candèze, 1886 syn. nov. = C. monachus Candèze, 1893 syn. nov.; C. lenzi Candèze, 1886 = C. behrensi Candèze, 1886 syn. nov.; C. oxydatus Candèze, 1857 = C. jekeli Candèze, 1874 syn. nov.; C. porcatus (Linnaeus, 1767) = C. peruanus Candèze, 1886 syn. nov. = C. flavostriatus Pjatakowa, 1941 syn. nov. = C. herbstii multistriatus Golbach, 1977 syn. nov.; C. rugatus Candèze, 1857 = C. amictus Casey, 1907 syn. nov.; C. smaragdinus LeConte, 1854 = C. ostentus Casey, 1907 syn. nov. = C. rectus Casey, 1907 syn. nov.; C. sulcatus (Fabricius, 1777) = C. herbstii Erichson, 1841 syn. nov; C. virens (Fabricius, 1787) = C. perrisi Candèze, 1857 syn. nov.; C. virginalis Candèze, 1857 = C. championi Casey, 1907 syn. nov.; C. viridipilis (Say, 1825) = C. debilis Casey, 1907 syn. nov.; C. webbi LeConte, 1854 = C. sonoricus Casey, 1907 syn. nov.; C. zonatus Eschscholtz, 1829 = C. longicollis Candèze, 1857 syn. nov. New species described: C. albisetosus sp. nov. (Ecuador), C. albiventris sp. nov. (Mexico: Veracruz), C. copulatuvittatus sp. nov. (Venezuela), C. extenuatuvittatus sp. nov. (Venezuela), C. fasciatus sp. nov. (Mexico: Durango), C. ferratuvittatus sp. nov. (Ecuador), C. proximus sp. nov. (Mexico: Sinaloa), C. serricornis sp. nov. (Mexico: Veracruz), C. spinipennis sp. nov. (Mexico: Veracruz), C. supremus sp. nov. (Venezuela), C. truncuvittatus sp. nov. (Mexico: Tamaulipas) and C. virgatipennis sp. nov. (Mexico: Durango). Redescribed species: C. angustatus Candèze, 1857, C. apacheanus Casey, 1891, C. approximatus Erichson, 1841, C. attenuatus Erichson, 1841, C. aurulentus Candèze, 1874, C. bomplandii Guérin-Méneville, 1844, C. boucardi Candèze, 1874, C. chalcantheus Candèze, 1857, C. corpulentus Candèze, 1874, C. cyaneus Candèze, 1881, C. desmarestii Chevrolat, 1835, C. dugesi Candèze, 1886, C. erythroloma Candèze, 1857, C. eschscholtzi Chevrolat, 1833, C. exulatus Candèze, 1874, C. fabricii Erichson, 1841, C. forreri Candèze, 1886, C. fryi Candèze, 1874, C. gossipiatus Guérin-Méneville, 1844, C. inops Candèze, 1886, C. jansoni Candèze, 1874, C. lacordairii Candèze, 1857, C. lafargi Chevrolat, 1835, C. lenzi Candèze, 1886, C. limbatus (Fabricius, 1777), C. mexicanus Castelnau, 1836, C. mniszechi Candèze, 1881, C. mocquerysii Candèze, 1857, C. morio Candèze, 1857, C. obscurus Castelnau, 1836, C. oxydatus Candèze, 1857, C. porcatus (Linnaeus, 1767), C. pruinosus Erichson, 1841, C. rodriguezi Candèze, 1886, C. rostainei Candèze, 1889, C. rubripennis LeConte, 1861, C. rugatus Candèze, 1857, C. silbermanni Chevrolat, 1835, C. smaragdinus LeConte, 1854, C. sulcatus (Fabricius, 1777), C. tartarus Fall, 1898, C. validus Candèze, 1857, reval., C. villei Candèze, 1878, C. virens (Fabricius, 1787), C. virginalis Candèze, 1857, C. viridipilis (Say, 1825), C. webbi LeConte, 1854, C. zonatus Eschscholtz, 1829.

Tornabenea ribeirensis (Apiaceaeb a new species from São Nicolau, Cape Verde Islands (West Africa)

Many species of Apiaceae are found in the Macaronesien Region. Several have been introduccd by human activities, but a number of taxa is endemic to the different archipelagos or even rrstrictcd to a single island. The following enumeration is based mainly on HANSEN & SUNDING ( 1993). In the Arores 28 different taxa of Apiaceae occur; among them four endemic species [AUIMI; hrrrrii WATSON, A. trifoliatum (WATSON) TKEL., Clrtrc~,y~l~~ll~r~~~ cl:oricrm TREL.. SOErich trwrictr GUTIINICK ex SEUB.]. In Madeira the Apiaceae are very diverse and consist ol’ 29 species and subspecies. From the archipelago two monotypic genera, rC/c/trtio. velitru~rr t/ccipicvr.s (SCHRAD. & J. C. WENDL.) Ho~+hl. md kJorli:ia edu[is LOWE and ~hrcc cndcmic species [Oemmrlre diwricore (R. BR.) MABB.. I/nperrr/orio lotvei COSS. and Burrirr~r hre~$~lirrnr LOWE] are described. The Canary Islands have the highest numbcr of plant-species and a high level of endemism. 5-l taxa of Apiaceae are recorded including three endemic genera (Rtrrheopsis A. HANSEX & KUNKEL, Todm-oa PARL. and Tiqyrmm PARL.) and further I5 endemic taxa. The Apiaceae are represented in the Cape Verde Islands by I2 species. Most of the taxa have been introduced by human activities (LOBIN & ZIZKA 1957) like Amvhm grm’eo- 1efr.s L., Apirm grmvolerrs L, Foerricrrhrr urlgore MILL.. Corimrtlru~t~ srrtirvrrrr L. or Petrosilerrm crisprrm (FRILL.) A.W.HILL. These species are cultivated and some of them later became \\esdy. Other species like Ciclosper- UWL /e/~fo/~/l~ll~rrtr (PER%) SPRAGUE (= Apimr leproplr~llrr~rr) are weeds of cultivated grounds or wasted lands. All these species are today widespread in temperate. subtropical or tropical regions all over the world. The only native species are to be found in the endemic genus To~wI~~I~~~ PARL.

The genus Agelaia Lepeletier (Hymenoptera, Vespidae, Polistinae) in Rio Grande do Sul, Brazil

The genus Agelaia Lepeletier 1836 belongs to the swarming genera of the Polistinae, with species distributed from Mexico to northern Argentina. Fifteen of the 31 described species are found in Brazil. Four species occur in the State of Rio Grande do Sul, two of them recorded herein for the first time. Redescriptions and a key to these species are provided.

New records of Chironomus Meigen (Diptera, Chironomidae) in the State of São Paulo, Brazil

In this paper seven species of the genus Chironomus Meigen, 1803 are recorded for the first time in the State of São Paulo (Brazil). C. stigmaterus Say, 1823 and C. columbiensis Wülker et al., 1989 are new records for Brazil, while C. calligraphus Goeldi, 1905, C. strenzkei Fittkau, 1968, C. gigas Reiss, 1974, C. latistylus Reiss, 1974 and C. paragigas Reiss, 1974 had already been registered in aquatic systems in North and Northeast regions of Brazil.

A new species of Cervellus Szépligeti (Hymenoptera, Braconidae, Braconinae) with biological notes

A new Cervellus species is described from Brazil. The male to this genus is described for the first time as well its cocoon is illustrated. The specimens parasitize the papaya borer weevil Pseudopiazurus obesus (Boheman, 1838) (Coleoptera, Curculionidae) associated with Carica papaya Linnaeus, 1753 (Caricaceae) at Rio Grande do Norte and Bahia Brazilian States.

New and newly illustrated Gnomidolon Thomson (Coleoptera, Cerambycidae, Hexoplonini) from Bolivia

Three new species of Gnomidolon Thomson, 1864 (Coleoptera, Cerambycidae, Hexoplonini) from Bolivia are described and illustrated, and G. proseni Martins, 1962 is redefined. Photographs of G. proseni and four other Bolivian species of this genus are provided: G. bonsae Martins, 1967, G. nanum Martins, 1962, G. musivum (Erichson, 1847), and both pale and dark forms of G. wappesi Martins, 2006.

New enigmatic Andean bee species of Protandrena (Hymenoptera, Andrenidae, Panurginae)

Panurgine bees are diverse and abundant in temperate areas of the Americas but poorly represented to nearly absent in the tropics. We describe and illustrate five distinctive new species of the genus Protandrena that occur at high altitudes (2000-3400 m) in the Andes, from Venezuela to Ecuador. The species are also described to make the names available in forthcoming papers on their biology. These Andean species resemble some members of the subgenus Heterosarus but differ from it, as well as from any other subgenera of Protandrena, primarily in characters of the male genitalia and hidden sterna. The South American Protandrena s. l. are morphologically highly diverse and a complete study of the group is needed before supraspecific names are proposed for unusual species. Thus, to avoid further nomenclatural changes, we decided not to place these species in a new subgenus or any of the available subgenera. We also provide notes on the biology for some of the species.

The presence of Chane Nieto and Guajirolus Flowers (Ephemeroptera, Baetidae) in Brazil with the description of a new species

The related genera Chane and Guajirolus are reported for the first time from Brazil. Guajirolus rondoni, sp. n. is described based on nymphs from Rondônia State and can be differentiated from the other species of the genus by the following combination of characteristics: 1) mandibles with minute spicules on outer margin; 2) maxillary palp three-segmented; 3) hypopharynx lingua with three rounded processes; 4) projection of second segment of labial palp with apex narrower than base; 5) abdominal color pattern; and 6) paraprocts with 13 to 15 organized spines. An updated key to the species of both genera is presented.

Phylogeny of wasps of the genus Mischocyttarus de Saussure (Hymenoptera, Vespidae, Polistinae)

A phylogenetic analysis is presented of subgenera and species-groups of Mischocyttarus de Saussure, the largest genus of social wasps. The analysis is based on 62 morphological and nest architecture characters, coded for 71 terminals representing much of the taxonomic diversity within the genus, plus three outgroup terminals representing other polistine tribes. The main conclusions about phylogenetic relationships within the genus are based on parsimony analysis under implied weights. Monophyly of Mischocyttarus is confirmed as well as that of most of the previously recognized subgenera: Mischocyttarus s. str., Clypeopolybia, Monogynoecus, Scytokeraia, Phi, Kappa, Megacanthopus and Omega sensu Richards (1978). Haplometrobius as conceived by Richards (1978) is not a monophyletic taxon, but some of its species-groups are monophyletic. The groups of M.artifex and M.cerberus are raised to subgenus level, and a new concept of Haplometrobius restricts it to the group of M.iheringi (the type species of this subgenus) in the sense of this work. The concept of subgenus Omega is widened to include the species-groups of M.surinamensis and M.prominulus. Besides the new subgeneric classification presented, limits and diagnoses of all species-groups of the subgenera Phi and Haplometrobius sensu Richards (1978) are discussed, and a new key for all subgenera and species-groups of Mischocyttarus is also presented.

A new species of Dicrepidius Eschscholtz from Brazil and new records for D. ramicornis (Palisot de Beauvois) (Coleoptera, Elateridae, Elaterinae)

Dicrepidius brasilianus sp. nov., from Pará and Mato Grosso is described and illustrated. This is the second species of this genus recorded from Brazil. D. ramicornis (Palisot de Beauvois, 1805) is widely distributed from south of United States to south of Brazil. From Brazil, it was recorded from Minas Gerais, Rio de Janeiro, São Paulo and Santa Catarina states, but now other records are included. A comparison between the two Brazilian species and a discussion, including intraspecific variations, are presented.

A new cluster-brood building species of Plebeia (Hymenoptera, Apidae) from eastern Brazil

A new species of Plebeia, the second largest genus of stingless bees in the Neotropical region, is described from eastern Brazil. Plebeia grapiuna sp. nov., known only from the lowland forests of southern Bahia, is most similar to P. lucii Moure, a species recently described from Minas Gerais. The lack of yellow marks and the smoother integument of the frons and mesoscutum in P. grapiuna sp. nov. distinguish them. Main features of the nesting habits of the new species are described and illustrated.

A new species of Seira Lubbock (Collembola, Entomobryidae), with a key to the species of Paraíba, Brazil

A new species of Entomobryidae of the genus Seira Lubbock, S. paraibensis sp. nov. is described and illustrated. The type locality of the species is the municipality of Areia, state of Paraíba, Brazil. The species was found inhabiting the Atlantic forest litter and top soil environment. S. paraibensis sp. nov. resembles S. pseudoannulata in many morphological features. This is the sixth species of the genus found and described in Paraiba so far. A key with the 11 species of Seira registered so far in Paraíba is provided. Also a list with all species of the genus from Brazil is presented.