Miocene honey bees from the Randeck Maar of southwestern Germany (Hymenoptera, Apidae)

The Miocene Randeck Maar (southwestern Germany) is one of the only sites with abundant material of fossil honey bees. The fauna has been the focus of much scrutiny by early authors who recognized multiple species or subspecies within the fauna. The history of work on the Randeck Maar is briefly reviewed and these fossils placed into context with other Tertiary and living species of the genus Apis Linnaeus (Apinae: Apini). Previously unrecorded specimens from Randeck Maar were compared with earlier series in an attempt to evaluate the observed variation. A morphometric analysis of forewing venation angles across representative Recent and Tertiary species of Apis as well as various non-Apini controls was undertaken to evaluate the distribution of variation in fossil honey bees. The resulting dendrogram shows considerable variation concerning the wing venation of Miocene Apini, but intergradation of other morphological characters reveals no clear pattern of separate species. This suggests that a single, highly variable species was present in Europe during the Miocene. The pattern also supports the notion that the multiple species and subspecies proposed by earlier authors for the Randeck Maar honey bee fauna are not valid, and all are accordingly recognized as Apis armbrusteri Zeuner.     

The phylogenetic position of the bumble bee inquiline Bombus inexspectatus and implications for the evolution of social parasitism

The high alpine European bumble bee Bombus (Thoracobombus) inexspectatus is one of three taxa in which obligate social parasitism has evolved in bumble bees. Until now, the phylogenetic placement of this species has not been analyzed quantitatively because it is rare in nature. Here a specimen of B. inexspectatus is sequenced for five genes to assess its phylogenetic position relative to other bumble bee species. Phylogenetic estimation of B. inexspectatus places this species within the subgenus Thoracobombus, as expected based on morphology. This provides strong support for the acquisition of social parasitism in B. inexspectatus separate from that found in the bumble bees of the subgenus Psithyrus and B. (Alpinobombus) hyperboreus. Furthermore, B. inexspectatus is not the sister taxon of its host but is a near relative, suggesting a loose but not strict adherence to Emery’s rule. B. inexspectatus is a sister taxon to a clade including B. veteranus, a frequent nest usurper that has been suggested to be a facultative social parasite. The phylogenetic placement of all bumble bee social parasites relative to their hosts is discussed.     

A NEW FOSSIL HOVERFLY (INSECTA, DIPTERA: SYRPHIDAE) FROM THE RANDECK MAAR (EARLY MIOCENE, SOUTH-WEST GERMANY)

Abstract:  Helophilus nothobombus sp. nov. (Diptera: Syrphidae) is described from Early Miocene lake sediments of the Randeck Maar (Swabian Alb, south-west Germany) and compared to other fossil species of the genus. Helophilus primarius Germar, 1837 and Helophilus villeneuvi Théobald, 1937 do not show diagnostic features that allow assignment to the genus Helophilus or even to the tribe Eristalini. Helophilus miocaenicus (Stackelberg, 1925) and H. nothobombus thus are the only plausible fossil species of the genus. The rat-tailed maggots of Helophilus most probably were part of the biocoenosis of the shores of the Randeck Maar lake. The frequency of larvae of Stratiomyidae shows that there were also suitable habitats for species of Helophilus and related genera.     

Gastropod palaeohabitats of Miocene Lake Randeck Maar and its hinterland defined by an actualistic genus-level approach

The Randeck Maar lake sediments from SW Germany were deposited during a climatic phase known as the Mid-Miocene Climatic Optimum, providing a unique window into this last favourable period for a more thermophilous fauna in Central Europe. Previous palaeoecological reconstructions of Randeck Maar's environment focused on the flora and vertebrate fauna. Here, we present a palaeoecological analysis using gastropods as proxies by means of an actualistic genus-level approach. This approach is grounded in comparisons of habitat preferences between extant and fossil congeners and thus obviously requires a sound taxonomic framework. Despite being commonly applied in the literature, this approach has hardly been defined so far. Therefore, we thoroughly delineate it, exploring its applicability, potential and shortcomings. The molluscan fauna of Randeck Maar counts with 32 species of continental snails, mostly terrestrial pulmonates, related to the following palaeohabitats: littoral, supralittoral, rocky walls, shrubland, forests and open habitats. The gastropods indicate a warm-temperate climate with a dry season, in line with previous works on Randeck Maar's flora and vertebrates.     

Mite‐y bees: bumble bees (Bombus spp., Hymenoptera: Apidae) host a relatively homogeneous mite (Acari) community,shaped by bee species identity but not by geographic proximity

1. Parasites can affect the communities of their hosts; and hosts, in turn, shape communities of parasites and other symbionts. This makes host–symbiont relationships a key but often overlooked aspect of community ecology. 2. Mites associated with bees have a range of lifestyles; however, little is known about mites associated with wild bees or about factors influencing the make‐up of bee‐associated mite communities. This study investigated how mite communities associated with bumble bees (Bombus spp.) are shaped by the Bombus community and geographic proximity. 3. Bees were collected from 15 sites in Ontario, Canada, and examined for mites. Mite abundance and species richness increased with local bee abundance. Several bee species also differed in mite abundance, species richness, prevalence, and diversity. Locally uncommon species tended to have more mites than other bees. Queen bees had the most mites, and males had more mites than workers. 4. Spatial proximity was not a predictor of mite community composition, despite a strong effect of proximity on bee community similarity. 5. On the 11 Bombus spp. examined, 33 mite species were found. Whereas nearly half of these mite species are obligate associates of bumble bees, none was restricted to particular Bombus species. 6. The best predictor of mite community composition was bee identity. Although many parasite communities show strong geographic patterns, the communities of primarily commensalistic bee‐mites in this study did not. These findings have implications for bumble bee conservation, given that pollen‐feeding commensals might become harmful at high densities or act as disease vectors.     

Tailoring your bee sampling protocol: Comparing three methods reveals the best approaches to capturing bees

1. Bees are prolific, vital pollinators in agricultural and natural settings, but some taxa are declining. Surveying bees is crucial to understand the needs of these taxa; however, we lack a fine-grained understanding of assemblages associated with different sampling methods that would enable us to analyse data range-wide.
2. Here we examine the difference in abundance and richness of bees (bee bowls and vane traps only) and bumble bees (genus Bombus; bee bowls, blue vane traps and target netting) sampled with these methods from mixed-grass prairie to alpine habitats in Wyoming, USA.
3. We collected four times more bees and twice as many genera of bees in vane traps than bee bowls. Vane traps captured more individuals of abundant genera than bee bowls.
4. Bombus species abundance did not vary between vane traps and target netting; however, richness was higher in vane traps. Bee bowls captured few Bombus species.
5. Overall, we recommend using vane traps to sample most bees, and a combination of vane traps and target netting to collect bumble bees. We evaluated how three sampling techniques perform when surveying for wild bees, which will aid in identifying declining species as well as monitoring species of conservation concern.

     

Local bumble bee decline linked to recovery of honey bees,drought effects on floral resources

Time series of abundances are critical for understanding how abiotic factors and species interactions affect population dynamics, but are rarely linked with experiments and also scarce for bee pollinators. This gap is important given concerns about declines in some bee species. I monitored honey bee (Apis mellifera) and bumble bee (Bombus spp.) foragers in coastal California from 1999, when feral A. mellifera populations were low due to Varroa destructor, until 2014. Apis mellifera increased substantially, except between 2006 and 2011, coinciding with declines in managed populations. Increases in A. mellifera strongly correlated with declines in Bombus and reduced diet overlap between them, suggesting resource competition consistent with past experimental results. Lower Bombus numbers also correlated with diminished floral resources. Declines in floral abundances were associated with drought and reduced spring rainfall. These results illustrate how competition with an introduced species may interact with climate to drive local decline of native pollinators.     

CAESALPINIA SUBGENUS MEZONEURON (LEGUMINOSAE,CAESALPINIOIDEAE) FROM THE TERTIARY OF NORTH AMERICA

Legume fruits from the Eocene of Tennessee and Wyoming and the Miocene of Idaho are described and assigned to Caesalpinia subgenus Mezoneuron (Caesalpinioideae), an extant Paleotropical taxon that does not occur in North or South America today. Morphological and anatomical details of the fruits are used in evaluating their systematic relationships. The features of the fossil fruits are accommodated only within this extant subgenus. These fossils represent the only reliable known occurrence of C. subgenus Mezoneuron in the paleobotanical record. These fossils suggest that subgenus Mezoneuron was distinct from subgenus Caesalpinia by the Middle Eocene. Further, they document the widespread occurrence of this currently Paleotropical group for at least 30 million years in North America.     

The frequency of multiple paternity in bumble bee (<Emphasis Type="Italic">Bombus</Emphasis>) colonies based on microsatellite DNA at the B10 locus

Summary This study used the microsatellite locus B10 to determine the frequency of colonies with multiple patrilines in a previously unexamined group, the North American bumble bees (Bombus). The effective mating frequency (m_e) was greater than 1 in six of 28 colonies. Five of 11 species tested showed at least one incidence of polyandry: four species from the subgenus Pyrobombus (B. bimaculatus, B. impatiens, B. mixtus, B. ternarius) and one species from the parasitic subgenus Psithyrus (B. citrinus). The B10 locus showed high cross-species amplification success for North American Bombus.     

Reduced abundance and earlier collection of bumble bee workers under intensive cultivation of a mass‐flowering prairie crop

One of the most commonly seeded crops in Canada is canola, a cultivar of oilseed rape (Brassica napus). As a mass‐flowering crop grown intensively throughout the Canadian Prairies, canola has the potential to influence pollinator success across tens of thousands of square kilometers of cropland. Bumble bees (Bombus sp.) are efficient pollinators of many types of native and crop plants. We measured the influence of this mass‐flowering crop on the abundance and phenology of bumble bees, and on another species of social bee (a sweat bee; Halictus rubicundus), by continuously deploying traps at different levels of canola cultivation intensity, spanning the start and end of canola bloom. Queen bumble bees were more abundant in areas with more canola cover, indicating that this crop is attractive to queens. However, bumble bee workers were significantly fewer in these locations later in the season, suggesting reduced colony success. The median collection dates of workers of three bumble bee species were earlier near canola fields, suggesting a dynamic response of colonies to the increased floral resources. Different species experienced this shift to different extents. The sweat bee was not affected by canola cultivation intensity. Our findings suggest that mass‐flowering crops such as canola are attractive to bumble bee queens and therefore may lead to higher rates of colony establishment, but also that colonies established near this crop may be less successful. We propose that the effect on bumble bees can be mitigated by spacing the crop more evenly with respect to alternate floral resources.     

Assessing the Threatened Status of Bumble Bee Species (Hymenoptera: Apidae) in Hungary,Central Europe

Decline in the populations of bumble bees and other pollinators stress the need for more knowledge about their conservation status. Only one of the 25 bumble bee species present in Hungary is included in the Hungarian Red List. We estimated the endangerment of the Hungarian bumble bee (Bombus Latr.) species using the available occurrence data from the last 50 years of the 20th century. Four of the 25 species were data deficient or extinct from Hungary. About 60% of species were considered rare or moderately rare. Changes in distribution and occurrence frequency indicated that 10 of the 21 native species showed a declining trend, while only three species increased in frequency of occurrence. According to the IUCN Red List categories, seven species (33% of the native fauna) should be labelled as critically endangered (CR) and 3 (14%) as endangered (EN). Our results stress an urgent need of protection plans for bumble bees in Hungary, and further underlines the validity of concern over bumble bees all over Europe.     

Ancient DNA from an extinct Mediterranean micromammal—Hypnomys morpheus (Rodentia: Gliridae)—Provides insight into the biogeographic history of insular dormice

The dormice (Gliridae) are a family of rodents represented by relatively few extant species, though the family was much more species-rich during the Early Miocene. Intergeneric phylogenetic relationships among glirids in some cases remain unresolved, despite extensive molecular and morphological analyses. Uncertainty is greatest with respect to the relationships among fossil taxa and how extinct lineages are related to modern species. The fossil genus Hypnomys from the Balearic Islands (western Mediterranean Sea) includes the Late Pleistocene–Holocene species Hypnomys morpheus, which has variously been considered a close relative or subgenus of the extant Eliomys. In the present study, we sequenced ancient mitochondrial DNA from H. morpheus, which suggests a sister relationship with the extant members of Eliomys. In addition, the pairwise sequence variation between Hypnomys and Eliomys is higher than that observed between congeneric glirid species (e.g., many Graphiurus spp.), which allows us to reject the hypothesis that Hypnomys is a subgenus of Eliomys. Our molecular dating analyses suggest that Hypnomys and Eliomys diverged 13.67 million years ago (95% highest posterior density [HPD] = 7.39–20.07). The relatively early split between these genera together with the molar morphology of early representatives of Hypnomys points to a Middle-Late Miocene origin from a continental glirid with a complex molar pattern, such as Vasseuromys or a closely related genus.     

Survey of bumble bee (Bombus) pathogens and parasites in Illinois and selected areas of northern California and southern Oregon

Pathogens have been implicated as potential factors in the recent decline of some North American bumble bee (Bombus) species, but little information has been reported about the natural enemy complex of bumble bees in the United States. We targeted bumble bee populations in a state-wide survey in Illinois and several sites in California and Oregon where declines have been reported to determine presence and prevalence of natural enemies. Based on our observations, most parasites and pathogens appear to be widespread generalists among bumble bee species, but susceptibility to some natural enemies appeared to vary.     

Bumble bee colony dynamics: quantifying the importance of land use and floral resources for colony growth and queen production

Bumble bee (Bombus) species are ecologically and economically important pollinators, and many species are in decline. In this article, we develop a mechanistic model to analyse growth trajectories of Bombus vosnesenskii colonies in relation to floral resources and land use. Queen production increased with floral resources and was higher in semi‐natural areas than on conventional farms. However, the most important parameter for queen production was the colony growth rate per flower, as opposed to the average number of available flowers. This result indicates the importance of understanding mechanisms of colony growth, in order to predict queen production and enhance bumble bee population viability. Our work highlights the importance of interpreting bumble bee conservation efforts in the context of overall population dynamics and provides a framework for doing so.     

Pollen morphology and fossil record of the feathery mistletoe family Misodendraceae

Misodendraceae is a small family of mistletoes in the order Santalales. Its distribution is restricted to the southern South American temperate forests. The family comprises the sole genus Misodendrum with eight species of hemiparasitic shrubs, mainly parasitising the southern beech Nothofagus. This contribution presents palynological evidence from seven species, using light microscopy and scanning electron microscopy. Pollen grains are consistently small, periporate and echinate, although differences in the length of echini and number and size of pores were noted. Pollen features can be used to distinguish groups of species and, in some cases, individual species. Cluster analysis of pollen characters differentiates two main groups: one includes M. brachystachyum, M. oblongifolium and M. quadriflorum; and the other includes M. gayanum, M. linearifolium, M. punctulatum and M. angulatum. Palynological results are compared with previous systematic studies of the family. The South American fossil pollen record is summarised and characters of the fossil pollen are analysed using UPGMA to test the relationships between extant and fossil species. Miocene pollen resulted similar to species of subgenus Angelopogon while Eocene pollen is disimilar to extant species of Misodendraceae.     

Longer tongues and swifter handling: why do more bumble bees (Bombus spp.) than honey bees (Apis mellifera) forage on lavender (Lavandula spp.)?

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Bumble bee species exhibit divergent responses to urbanisation in a Southern California landscape

1. Bees are ecologically important pollinators that are threatened by disease, habitat degradation, and habitat loss. Bee species that share ecological traits (e.g. body size, diet breadth, and sociality) may often respond similarly to environmental disturbance; however, few studies have examined the extent to which closely related, ecologically similar species exhibit divergent responses to the same forms of disturbance. 2. In the present study, the responses of bumble bees (Apidae: genus Bombus) to urbanization are examined by combining field surveys with the quantification of local‐ and landscape‐level variables using Geographic Information Systems (GIS). During the spring and summer of 2012 and 2013, 57, bumble bees were surveyed in 1‐ha plots in San Diego County, California that spanned a gradient of urbanisation. Species‐level, occurrence (presence/absence) data and logistic regressions were then used to identify determinants of Bombus occurrence across this landscape. 3. Approximately 1000 Bombus individuals belonging to three species were observed: B. californicus Smith, B. melanopygus Cresson, and B. vosnesenskii Radoszkowski. The occurrence of all Bombus species was negatively related to impermeable surface cover within study plots, but species responded uniquely to other plot‐scale variables and exhibited distinctive patterns of scale‐dependency with respect to impermeable surface cover surrounding plots. 4. Urbanisation negatively affected all three focal bumble bee species in this study. Species‐specific responses to plot‐scale and landscape‐scale variables presumably reflect interspecific trait differences (e.g. body size, tongue length, and foraging behaviour). Unique responses to urbanisation caution against pooling species into functional groups based merely on taxonomic relationships or perceived ecological similarities.     

Endocranial morphology of Pygoscelis calderensis (Aves,Spheniscidae) from the Neogene of Chile and remarks on brain morphology in modern Pygoscelis

The endocranial anatomy of Pygoscelis calderensis, a fossil species from the Bahía Inglesa Formation (Middle Miocene–Pliocene) of Chile, South America, was described through CT scans. Reconstructions of the fossil P. calderensis and endocasts for the living Pygoscelis adeliae, and Pygoscelis papua are provided here for the first time. Comparisons with the extant congeneric species P. adeliae, Pygoscelisantarctica and P. papua indicate that the morphological pattern of the brain and inner ear of the extant pygoscelids has been present already in the Middle Miocene. The neurological morphology suggests that the paleobiology of the extinct form would have been similar to the extant species. It was probably true for diet, feeding behaviour and diving kinematics.     

Bridging the extant and fossil record of planktonic foraminifera: implications for the Globigerina lineage

We conducted a morphometric study and wall texture analysis on extant and fossil specimens of the planktonic foraminifera Globigerina falconensis plexus. Our global data reveal morphological inconsistencies between fossil and extant populations. Our results are significant as G. falconensis is widely used in palaeoceanographic studies in conjunction with its sister taxon G. bulloides. Morphologically these two species are similar, with the main difference being the distinctive apertural lip present in G. falconensis. We selected cores covering the entire stratigraphic range of G. falconensis, from the early Miocene to current day, spanning sites from high latitudes in the North Atlantic Ocean and the southern Indian Ocean to sites in equatorial regions. The morphology found in the modern ocean is not consistent with the Miocene holotype of Globigerina falconensis Blow described from lower Miocene sediments in Venezuela. A more lobate morphology evolved in the late Miocene, thus, a new name is required for this morphotype, coexisting in the modern oceans with G. falconensis s.s. We thus describe the new morphospecies, G. neofalconensis for the more lobate forms which evolved in the late Miocene and inhabit the modern oceans. Additionally, we report a pseudocancellate wall texture present in the G. falconensis plexus. We use the molecular sequences from the PR² database to explore the generic attribution of the G. falconensis lineage, confirming its close relationship with G. bulloides and its retention in the genus Globigerina.     

Cryptic species diversity in a widespread bumble bee complex revealed using mitochondrial DNA RFLPs

Cryptic species diversity is thought to be common within the class Insecta, posing problems for basic ecological and population genetic studies and conservation management. Within the temperate bumble bee (Bombus spp.) fauna, members of the subgenus Bombus sensu stricto are amongst the most abundant and widespread. However, their species diversity is controversial due to the extreme difficulty or inability morphologically to identify the majority of individuals to species. Our character-based phylogenetic analyses of partial CO1 (700 bp) from 39 individuals spread across their sympatric European ranges provided unequivocal support for five taxa (3–22 diagnostic DNA base pair sites per species). Inclusion of 20 Irish specimens to the dataset revealed ≥2.3% sequence divergence between taxa and ≤1.3% within taxa. We developed a PCR-RFLP based method for unequivocally distinguishing amongst the four cryptic European taxa of this subgenus, B. cryptarum, B. lucorum, B. magnus and B. terrestris, and used it to analyse 391 females of the former three species collected across Ireland, all of which could be unambiguously assigned to species. Bombus lucorum was the most widely distributed and abundant of the cryptarum–lucorum–magnus species complex, comprising 56% of individuals, though it was significantly less abundant at higher altitudes (>200 m) whilst B. cryptarum was relatively more abundant at higher altitudes. Bombus magnus was rarely encountered at urban sites. Both B. lucorum and B. terrestris are nowadays reared commercially for pollination and transported globally. Our RFLP approach to identify native fauna can underpin ecological studies of these important cryptic species as well as the impact of commercial bumble bees on them.