Alain Pauly

Phylogeny of Halictidae with an emphasis on endemic African Halictinae

We review the literature on phylogeny, fossil record, biogeography, and social evolution in Halictidae. We then present a phylogenetic analysis of tribal, generic, and subgeneric relationships within the subfamily Halictinae using a combined data set of three nuclear genes: long-wavelength (LW) opsin, wingless, and EF-1α. The data set includes 89 species in 34 genera representing all four halictid subfamilies, and all tribes of the subfamily Halictinae. Our study provides several new insights into the phylogeny of the African Halictinae. First, our results support a close relationship between Mexalictus (a small genus of bees occurring at high elevations in the mountains of western North and Central America) and the African/Asian genus Patellapis. Second, our results support placement of the parasitic genus Parathrincostoma well within its host genus Thrinchostoma, suggesting that Parathrincostoma should be treated as a subgenus of Thrinchostoma. Finally, our data set provides strong support for the monophyly of Patellapis (sensu Michener, 2000) and establishes monophyletic groups within the African subgenera that could be the basis for future taxonomic studies.ZusammenfassungIn der Bienenfamilie Halictidae sind über 3500 Arten aus allen Kontinenten beschrieben. Diese Familie vereint die verschiedensten Bienentaxa, darunter solitäre, soziale, kleptoparasitische und sozialparasitische Arten. Viele Arten sind bezüglich ihrer Trachtpflanzen Generalisten, andere Spezialisten. Unsere Untersuchungen konzentrieren sich auf die Unterfamilie Halictinae, zu der die allermeisten Arten gehören. Wir analysierten die evolutionsbiologischen Beziehungen bei den verschiedenen Unterfamilien, Tribus und Gattungen der Halictiden unter besonderer Berücksichtigung der endemischen afrikanischen Halictinae. Verglichen mit Vertretern der Halictiden aus anderen Teilen der Welt hat man den afrikanischen Gattungen und Untergattungen bisher wenig Beachtung geschenkt. Wir wissen bisher sehr wenig über ihr Sozial- und Nistverhalten und wie diese Bienen in die Phylogenie der Halictiden einzuordnen sind. Wir analysierten die DNA-Sequenzen von drei Genen (long-wavelength opsin, wingless und elongation factor-1α) und einen Datensatz von insgesamt 2536 Nukleotide unter Verwendung von Alignment-Analysen und Parsimony-Methoden. Anhand unserer Ergebnisse identifizierten wir zwei distinkte endemische afrikanische Abstammungslinien. Erstens den Tribus Thrinchostomini (einschließlich der Gattungen Thrinchostoma und Parathrincostoma), der früh im Stammbaum abzweigt. Zweitens eine gut abgesicherte monophyletische Gruppe (Patellapis sensu lato), die offensichtlich nahe verwandt mit dem Genus Mexalictus ist, der ausschließlich in hochgelegenen Gebieten vom westlichen Nordamerika bis nach Zentralamerika vorkommt. Dieses ungewöhnliche biogeographische Muster könnte darauf hinweisen, dass eine ursprünglich über die gesamte nördliche Hemisphäre verbreitete Gruppe ausgestorben ist. Wir überprüfen die Literatur zu den Halictiden bezüglich ihrer Beziehung zu Trachtpflanzen, ihrer Nistbiologie und dem Vorkommen von Kleptoparasitismus. Unsere Untersuchungen erlauben erstmals eine Beurteilung der Verwandtschaftsverhältnisse innerhalb der Halictinae anhand umfangreicher Proben aller afrikanischen Gattungen und Untergattungen.

Contribution à la connaissance des Hymenoptera Apoidea de Nouvelle-Calédonie et de leurs relations avec la flore butinée

Résumé Seulement 21 espèces d’Apoidea sont répertoriées de Nouvelle-Calédonie. La pauvreté de cette faune est paradoxale si on la compare à la richesse et l’endémicité de la flore de l’île, mais s’expliquerait par le fait qu’elle a été isolée avant ou peu après l’apparition des Apoidea vers – 130 millions d’années. Les auteurs comparent aussi les données de plusieurs îles des océans Pacifique et Indien. Deux nouvelles espèces de Halictidae sont décrites: Homalictus cocos et Lasioglossum (Chilalictus) delobeli. Une nouvelle synonymie est établie: Homalictus risbeci (Cockerell, 1929) = Homalictus crotalariae (Cockerell, 1929). Deux taxons, Chalicodoma umbripenne et Megachile laticeps, sont signalés pour la première fois en Nouvelle-Calédonie. Les relations entre les Apoidea et les 22 espèces végétales, sur lesquelles ces insectes ont été capturés, sont présentées et commentées.

Monograph of the bees of the subfamily Nomioidinae (Hymenoptera: Halictidae) of Africa (excluding Madagascar)

Abstract The paper presents the results of a study of 11863 specimens of Nomioidinae caught in Africa (excepting Madagascar) from 43 institutions and private collections. In total, 33 species belonging to three genera of Nomioidinae are found in the African fauna. All of the species are redescribed, figured and keyed. Their variability is analysed. Distributional maps in Africa are provided for all species. The types of nearly all the nominal taxa described earlier are examined. Seven new species and one new subspecies are described: Cellariella inexpectata n. sp., C. schwarzi n. sp., Ceylalictus congoensis n. sp., Nomioides deceptor capverdensis n. ssp., N. griswoldi n. sp., N. kenyensis n. sp., N. micheneri n. sp., and N. paulyi Pesenko, n. sp. New status is established for the following nominal taxa: Nomioides maculiventris var. fulviventris Blüthgen 1925 and N. somalicus ssp. kalaharicus Cockerell 1936 are considered separate species in the genus Cellariella; Nomioides karachensis var. desertorum Blüthgen 1925, a separate species in the genus Ceylalictus; Nomioides canariensis Blüthgen 1937, an insular subspecies of N. deceptor Saunders 1908; Nomioides maurus Blüthgen 1925, an African subspecies of N. minutissimus (Rossi 1790). The names Nomioides elbanus Blüthgen 1934 and N. squamiger Saunders 1908 are resurrected. The following new synonymies are established: Nomioides somalica ssp. kalaharica Cockerell 1936 = Cellariella brooksi Pesenko 1993; Nomioides somalicus Magretti 1899 = N. somalicus var. completus Blüthgen 1934 = N. atomellus Cockerell 1936; Nomioides deceptor Saunders 1908 = N. minutissimus var. deserticola Blüthgen 1925; Ceratina maculiventris Cameron 1905 = Nomioides maculiventris var. convergens Blüthgen 1934 = N. callonotus Cockerell 1936 = N. maculiventris var. cyaneonotus Cockerell 1937; Nomioides minutissimus spp. maurus Blüthgen 1925 = N. senecionis Cockerell 1931 = N. maurus var. tingitanus Blüthgen 1933. Lectotypes are designated for the following nominal taxa: Ceratina maculiventris Cameron 1905; Nomioides callonotus Cockerell 1936; N. maculiventris var. cyaneonotus Cockerell 1937, N. somalicus ssp. kalaharicus Cockerell 1936. The morphological evolution, phylogeny and geographical history of the subfamily Nomioidinae are presented and discussed. The paper includes 228 line drawings on 39 figures in the text, 124 colour, 8 black and white, 48 SEM photos and 33 maps on 20 plates, 7 tables, and provided with a list of 140 papers cited.

Flower-Visiting Records of the Native Bees of New Caledonia1

Abstract The flower-visiting records for the 43 species of bees considered to be native to New Caledonia show that females of 21 species visited 116 native species of plants in 69 genera and 41 families, and the bees were documented to carry pollen from 64 species and possibly four more. The plant families with the greatest number of species documented for visits by female bees were, in descending order: Myrtaceae (21), Dilleniaceae (10), Cunoniaceae (nine), Araliaceae (seven), Fabaceae (seven, encompassing the Caesalpinioideae, Mimosoideae, and Papilionoideae), Goodeniaceae (five), Proteaceae (five), Apocynaceae (four), Sapindaceae (four), and remaining families with one to three species. Females of six and possibly one more species carried pollen from each of Dilleniaceae and Myrtaceae, six carried pollen from Araliaceae, five from Goodeniaceae, four and possibly one more from Cunoniaceae, four from each of Fabaceae and Sapindaceae, and none to three from the remaining 34 families observed. For introduced plants, female bees of 12 species visited 54 species in 43 genera among 19 plant families and were documented to carry pollen from 31 and possibly one more species. For introduced plants, families with the highest number of species visited by female bees, in descending order, were: Asteraceae (12); Fabaceae (eight); Verbenaceae (seven); and Euphorbiaceae, Myrtaceae, and Solanaceae each with three. The remaining 13 families had either one or two species visited by bees. Females of seven species of bees carried pollen from Fabaceae, six from Asteraceae, three each from Myrtaceae and Solanaceae, and none to “2 + 1?” (two or possibly three) from remaining families. Only half a dozen species of native bees can be considered to be common, in that they can be expected to be observed reasonably regularly on a range of flowers. The ubiquitous introduced honey bee Apis mellifera L. and its constant foraging for nectar and pollen on a very wide range of flowers may outcompete many species of native bees, potentially reducing their numbers, and consequently obscuring their relationships with the flora.

Une abeille afrotropicale spécialisée dans la récolte du pollen de Graminées (Poaceae): Lipotriches notabilis (Schletterer 1891) (Hymenoptera Apoidea Halictidae)

Résumé Dans les zones de savanes de l’Afrique, un genre d’abeille, Lipotriches Gerstaecker 1858, s’est spécialisé dans la collecte du pollen de graminées. Un site de nidification et l’aire de butinage de Lipotriches notabilis ont été suivis pendant trois années dans la région de Ngaoundéré au Cameroun. Le régime alimentaire pollinique est composé presque exclusivement du pollen de graminées, notamment Brachiaria ruziziensis abondant dans cette région. Le maïs est aussi visité et l’abeille contribue indirectement à la pollinisation par la mise en suspension dans l’air du pollen. Comme les graminées n’offrent pas de ressource sucrée, les femelles comme les mâles de L. notabilis butinent de temps en temps les Asteraceae pour la collecte de nectar. La consommation du pollen de graminées par les femelles a été aussi observée. Lespèce niche en bourgades dont la taille varie d’une dizaine à une centaine de nids. Le nid creusé dans un sol horizontal est du type progressif. Il comprend un tumulus, une cheminée verticale, un conduit principal vertical pouvant atteindre 65 cm de profondeur et en moyenne 3 conduits latéraux obliques de 4 à 16 cm aboutissant dans une ou plusieurs cellules successives. Les cellules des conduits latéraux sont approvisionnées simultanément et fermées de manière régressive. On compte un maximum de 10 cellules par nid. Généralement un nid est habité par une seule femelle, mais certains nids sont habités par deux ou trois. Lactivité de butinage de cette espèce est limitée dans la matinée. Dès 7 heures, elle s’envole pour le site de butinage. Après 11 heures, il n’y a généralement plus de nids ouverts. Les mâles n’ont pas été aperçus au niveau du site de nidification et l’accouplement a lieu sur le site de butinage. La période d’activité commence avec la saison des pluies en avril et se termine au début de la saison sèche en décembre, avec la fanaison des graminées.

Mechanism of self-pollination in Hybanthus enneaspermus (L.) F. Muell. and notes on the floral biology of some Rinorea species (Violaceae) in Ivory Coast

Abstract Few studies have examined insects visiting flowers of tropical Violaceae. This study presents results from field work in Ivory Coast and from the examination of cultivated plants of three species in the two largest tropical genera, Rinorea and Hybanthus. Field observations showed that Hybanthus enneaspermus flowers opened only one morning and were visited by a single bee species. In contrast, the two species of Rinorea observed, which have flowers that do not close, were visited by many and various insects. Cultivation showed that self-pollination, with viable seeds production, appears likely in H. enneaspermus, perhaps as a means of compensating for the possible scarcity of pollinators. Floral structures would support allogamy first, and then lead to autogamy.

Addition to the checklist of IUCN European wild bees (Hymenoptera: Apoidea)

Summary The present study is an update to the first Red List of European Bees published in 2014. The additional records are based on (i) comprehensive review of literature; (ii) new data provided by bee specialists in response to the publication of the first Red List; (iii) new developments in taxonomy of European bees including description of new cryptic species; (iv) new specimens from recent field collections. While the first Red List included a list of 1965 wild bee species with 75 genera, we found 86 additional species, while two more genera have been erected (Seladonia and Vestitohalictus), giving an updated total of 2051 species and 77 genera. The authors discuss the artificial framework of the study considered by International Union for Conservation of Nature (IUCN) to produce the first Red List and they propose the more meaningful West Palaearctic biogeographical region. For this whole region, a first estimation gives 3408 wild bee species in 105 genera. The next taxa have been erected from subspecies to species status: Andrena (Euandrena) limosa Warncke, 1969, stat. n., Andrena (Plastandrena) oligotricha Mavromoustakis, 1952, stat. n.

The bees (Hymenoptera: Apoidea) of the Maltese Islands

This study presents the first checklist of the bees of the Maltese Islands and includes notes on the distribution of each species. A total of 95 species belonging to five bee families are recorded: Andrenidae (17 species), Apidae (34 species), Colletidae (6 species), Halictidae (15 species) and Megachilidae (23 species). Lasioglossum callizonium (Pérez, 1896) is recorded for the first time from the Maltese Islands. Records of three previously reported species are listed as dubious. The bee fauna of the Maltese Archipelago is dominated by widespread West-Palaearctic species, and most of the species recorded are also found in the Western Mediterranean Basin. Bees that have been recorded from Malta are also known from Southern Europe. The study provides a biogeographical analysis of the Maltese bee fauna, and discusses the conservation of this group and their important role in the delivery of ecosystem services in the Maltese Islands.

Impact of human disturbance on bee pollinator communities in savanna and agricultural sites in Burkina Faso, West Africa

Abstract All over the world, pollinators are threatened by land‐use change involving degradation of seminatural habitats or conversion into agricultural land. Such disturbance often leads to lowered pollinator abundance and/or diversity, which might reduce crop yield in adjacent agricultural areas. For West Africa, changes in bee communities across disturbance gradients from savanna to agricultural land are mainly unknown. In this study, we monitored for the impact of human disturbance on bee communities in savanna and crop fields. We chose three savanna areas of varying disturbance intensity (low, medium, and high) in the South Sudanian zone of Burkina Faso, based on land‐use/land cover data via Landsat images, and selected nearby cotton and sesame fields. During 21 months covering two rainy and two dry seasons in 2014 and 2015, we captured bees using pan traps. Spatial and temporal patterns of bee species abundance, richness, evenness and community structure were assessed. In total, 35,469 bee specimens were caught on 12 savanna sites and 22 fields, comprising 97 species of 32 genera. Bee abundance was highest at intermediate disturbance in the rainy season. Species richness and evenness did not differ significantly. Bee communities at medium and highly disturbed savanna sites comprised only subsets of those at low disturbed sites. An across‐habitat spillover of bees (mostly abundant social bee species) from savanna into crop fields was observed during the rainy season when crops are mass‐flowering, whereas most savanna plants are not in bloom. Despite disturbance intensification, our findings suggest that wild bee communities can persist in anthropogenic landscapes and that some species even benefitted disproportionally. West African areas of crop production such as for cotton and sesame may serve as important food resources for bee species in times when resources in the savanna are scarce and receive at the same time considerable pollination service.

Using next-generation sequencing to improve DNA barcoding: lessons from a small-scale study of wild bee species (Hymenoptera, Halictidae)

The parallel sequencing of targeted amplicons is a scalable application of next-generation sequencing (NGS) that can advantageously replace Sanger sequencing in certain DNA barcoding studies. It can be used to sequence different PCR products simultaneously, including co-amplified products. Here, we explore this approach by simultaneously sequencing five markers (including the DNA barcode and a diagnostic marker of Wolbachia) in 12 species of Halictidae that were previously DNA barcoded using Sanger sequencing. Consensus sequences were obtained from fresh bees with success rates of 74–100% depending on the DNA fragment. They improved the phylogeny of the group, detected Wolbachia infections (in 8/21 specimens) and characterised haplotype variants. Sequencing cost per marker and per specimen (11.43 €) was estimated to decrease (< 5.00 €) in studies aiming for a higher throughput. We provide guidelines for selecting NGS or Sanger sequencing depending on the goals of future studies.