Twenty-five new species of mining bees (Hymenoptera: Andrenidae: Andrena) from Israel and the Levant.

Andrena is one of the most diverse bee genera, comprising about 1,600 described species of ground-nesting solitary bees. Many Andrena species are plant specialists, and several taxa have been indicated to be important pollinators of wild and/or crop plants. The Eastern Mediterranean Basin and Israel in particular are one of the main world diversity hotspots of Andrena. Based on extensive examination of museum specimens combined with DNA barcoding, we hereby describe twenty-five Levantine species of Andrena new to science: Andrena anathema Pisanty sp. nov., A. ardentia Pisanty sp. nov., A. asluji Pisanty sp. nov., A. curviocciput Pisanty & Wood sp. nov., A. dividicincta Pisanty sp. nov., A. dorchini Pisanty sp. nov., A. euphorbiae Pisanty sp. nov., A. gageae Wood & Pisanty sp. nov., A. herodesi Pisanty & Wood sp. nov., A. hulae Pisanty sp. nov., A. igraeca Pisanty & Wood sp. nov., A. inusitata Pisanty sp. nov., A. janthinoides Pisanty sp. nov., A. longistilus Pisanty & Wood sp. nov., A. lunaris Pisanty & Wood sp. nov., A. macula Pisanty & Wood sp. nov., A. obtusa Pisanty sp. nov., A. ornithogali Pisanty & Wood sp. nov., A. petrae Wood sp. nov., A. protuber Pisanty sp. nov., A. sulfurea Wood sp. nov., A. turmalina Pisanty & Wood sp. nov., A. veronicae Pisanty & Wood sp. nov., A. veterana Pisanty sp. nov., and A. xera Pisanty sp. nov. We synonymise Andrena edentula Wood with A. tadauchii Gusenleitner syn. nov., and recognise four infraspecific names as valid species: Andrena mediterranea Pisanty & Scheuchl stat. nov., A. mizorhina Warncke stat. nov., A. noacki Alfken sp. resurr. and A. ochraceohirta Alfken sp. resurr. We additionally describe the hitherto unknown sexes of four species, provide new records for fifteen species previously unknown from Israel, and list fourteen taxa whose previously reported presence in Israel is considered erroneous or questionable.

Molecular phylogeny, historical biogeography and revised classification of andrenine bees (Hymenoptera: Andrenidae).

The mining bee subfamily Andreninae (Hymenoptera: Andrenidae) is a widely distributed and diverse group of ground-nesting solitary bees, including numerous species known to be important pollinators. Most of the species diversity of Andreninae is concentrated in the mainly Holarctic genus Andrena, comprising ca. 1550 described species. The subfamily and especially the genus have remained relatively neglected by recent molecular phylogenetic studies, with current classifications relying largely on morphological characters. We sampled ultraconserved element (UCE) sequences from 235 taxa, including all andrenine genera and 98 out of 104 currently recognized Andrena subgenera. Using 419,858 aligned nucleotide sites from 1009 UCE loci, we present a comprehensive molecular phylogenetic analysis of the subfamily. Our analysis supports the recognition of seven distinct genera in the Andreninae: Alocandrena, Ancylandrena, Andrena, Cubiandrena, Euherbstia, Megandrena, and Orphana. Within the genus Andrena, present-day subgeneric concepts revealed high degrees of paraphyly and polyphyly, due to strong homoplasy of morphological characters, necessitating a thorough, extensive revision of the higher classification of the genus. Based on our findings, we place the subgenus Calcarandrena in synonymy with Andrena (Lepidandrena); Hyperandrena, Nemandrena, Scoliandrena, Tylandrena and Zonandrena with A. (Melandrena); Distandrena, Fumandrena and Proxiandrena with A. (Micrandrena); Carandrena with A. (Notandrena); Agandrena with A. (Plastandrena); Geandrena and Xanthandrena with A. (Ptilandrena); Xiphandrena with A. (Scrapteropsis); and Platygalandrena and Poliandrena with A. (Ulandrena) (new synonymies). We additionally reestablish the groups known as Opandrena and Truncandrena as valid subgenera of Andrena. Our results also show that the MRCA of Andrena + Cubiandrena dispersed from the New World to the Palaearctic probably during the Eocene-early Oligocene, followed by 10-14 Neogene dispersal events from the Palaearctic to the Nearctic and 1-6 Neogene dispersals back into the Palaearctic, all within the genus Andrena.

Wild insect diversity increases inter-annual stability in global crop pollinator communities.

While an increasing number of studies indicate that the range, diversity and abundance of many wild pollinators has declined, the global area of pollinator-dependent crops has significantly increased over the last few decades. Crop pollination studies to date have mainly focused on either identifying different guilds pollinating various crops, or on factors driving spatial changes and turnover observed in these communities. The mechanisms driving temporal stability for ecosystem functioning and services, however, remain poorly understood. Our study quantifies temporal variability observed in crop pollinators in 21 different crops across multiple years at a global scale. Using data from 43 studies from six continents, we show that (i) higher pollinator diversity confers greater inter-annual stability in pollinator communities, (ii) temporal variation observed in pollinator abundance is primarily driven by the three-most dominant species, and (iii) crops in tropical regions demonstrate higher inter-annual variability in pollinator species richness than crops in temperate regions. We highlight the importance of recognizing wild pollinator diversity in agricultural landscapes to stabilize pollinator persistence across years to protect both biodiversity and crop pollination services. Short-term agricultural management practices aimed at dominant species for stabilizing pollination services need to be considered alongside longer term conservation goals focussed on maintaining and facilitating biodiversity to confer ecological stability.

Taxonomic review of the subgenus Andrena (Poecilandrena) (Hymenoptera: Andrenidae) in Israel and the Levant.

Andrena (Poecilandrena) Hedicke is a subgenus of small solitary bees, with the greatest diversity in the Eastern Mediterranean Region-an important but understudied biodiversity hotspot for bees. We studied Andrena (Poecilandrena) collected mostly in Israel and the West Bank, and make several additions to the regional fauna. We provide the first comprehensive review of Andrena (Poecilandrena) species currently known from Israel, the West Bank, Jordan, Lebanon and Syria, including diagnostic keys to females and males, descriptions of new species and unknown sexes, and detailed information for each taxon regarding distribution, phenology and flower visitation. Our review includes fourteen species of Andrena (Poecilandrena) from the Levant, including five species new to science: A. freidbergi Pisanty Scheuchl n. sp., A. galilaea Pisanty Scheuchl n. sp., A. hierosolymitana Pisanty Scheuchl n. sp., A. sedumella Scheuchl Pisanty n. sp., and A. stenofovea Scheuchl Pisanty n. sp. We also report four species as new to the region, and provide the first description of the male of A. rusticola Warncke. We exclude A. arabica Scheuchl Gusenleitner from the subgenus, and synonymize it with A. helouanensis Friese. We anticipate that more collecting work in this region will yield additional undescribed taxa as well as new records, especially of taxa already known from Turkey. In the appendix to this work we provide information on 35 new records of other Andrena subgenera from Israel and the West Bank that have accumulated in recent years.

Eight new species of Andrena Fabricius (Hymenoptera: Apoidea: Andrenidae) from Israel-a Mediterranean hotspot for wild bees.

More than 150 solitary bee species of the genus Andrena Fabricius are known from Israel and the West Bank, where they are distributed along a broad climatic gradient and diverse habitats and vegetation types. Extensive collecting throughout Israel in recent years has yielded eight new species and one new subspecies, adding to the rich bee fauna of the region:   A. crocusella Pisanty & Scheuchl n. sp., A. danini Pisanty & Scheuchl n. sp., A. hermonella Scheuchl & Pisanty n. sp., A. israelica Scheuchl & Pisanty n. sp., A. judaea Scheuchl & Pisanty n. sp., A. menahemella Scheuchl & Pisanty n. sp., A. palaestina Pisanty & Scheuchl n. sp., A. perahia Pisanty & Scheuchl n. sp., and A. sphecodimorpha mediterranea Pisanty & Scheuchl n. ssp. The previously unknown female of A. fimbriatoides Scheuchl 2004 and male of A. wolfi Gusenleitner & Scheuchl 2000 are also described here for the first time. The discovery of males of A. wolfi lead us to reinstate A. iohannescaroli Nobile 2000 as a valid taxon. Detailed morphological description and differential diagnosis against closest relatives are provided for all species, as well as information on the distribution, phenology and flower visitation, when available. A neotype is designated for A. sphecodimorpha Hedicke, the holotype of which is considered to be lost. Additional collecting efforts in diverse habitats and seasons, incorporating diverse collecting techniques, are required in order to deepen our knowledge of the rich bee fauna in threatened habitats in the Mediterranean Basin, which constitutes one of the world's major hotspots for wild bees.

Non-bee insects are important contributors to global crop pollination.

Wild and managed bees are well documented as effective pollinators of global crops of economic importance. However, the contributions by pollinators other than bees have been little explored despite their potential to contribute to crop production and stability in the face of environmental change. Non-bee pollinators include flies, beetles, moths, butterflies, wasps, ants, birds, and bats, among others. Here we focus on non-bee insects and synthesize 39 field studies from five continents that directly measured the crop pollination services provided by non-bees, honey bees, and other bees to compare the relative contributions of these taxa. Non-bees performed 25-50% of the total number of flower visits. Although non-bees were less effective pollinators than bees per flower visit, they made more visits; thus these two factors compensated for each other, resulting in pollination services rendered by non-bees that were similar to those provided by bees. In the subset of studies that measured fruit set, fruit set increased with non-bee insect visits independently of bee visitation rates, indicating that non-bee insects provide a unique benefit that is not provided by bees. We also show that non-bee insects are not as reliant as bees on the presence of remnant natural or seminatural habitat in the surrounding landscape. These results strongly suggest that non-bee insect pollinators play a significant role in global crop production and respond differently than bees to landscape structure, probably making their crop pollination services more robust to changes in land use. Non-bee insects provide a valuable service and provide potential insurance against bee population declines.

Profiling crop pollinators: life history traits predict habitat use and crop visitation by Mediterranean wild bees.

Wild pollinators, bees in particular, may greatly contribute to crop pollination and provide a safety net against declines in commercial pollinators. However, the identity, life history traits, and environmental sensitivities of main crop pollinator species.have received limited attention. These are crucial for predicting pollination services of different communities and for developing management practices that enhance crop pollinators. We sampled wild bees in three crop systems (almond, confection sunflower, and seed watermelon) in a mosaic Israeli Mediterranean landscape. Bees were sampled in field/orchard edges and interiors, and in seminatural scrub surrounding the fields/orchards. We also analyzed land cover at 50-2500 m radii around fields/orchards. We used this data to distinguish crop from non-crop pollinators based on a set of life history traits (nesting, lecty, sociality, body size) linked to habitat preference and crop visitation. Bee abundance and species richness decreased from the surrounding seminatural habitat to the field/orchard interior, especially across the seminatural habitat-field edge ecotone. Thus, although rich bee communities were found near fields, only small fractions crossed the ecotone and visited crop flowers in substantial numbers. The bee assemblage in agricultural fields/orchards and on crop flowers was dominated by ground-nesting bees of the tribe Halictini, which tend to nest within fields. Bees' habitat preferences were determined mainly by nesting guild, whereas crop visitation was determined mainly by sociality. Lecty and body size also affected both measures. The percentage of surrounding seminatural habitat at 250-2500 m radii had a positive effect on wild bee diversity in field edges, for all bee guilds, while at 50-100 m radii, only aboveground nesters were positively affected. In sum, we found that crop and non-crop pollinators are distinguished by behavioral and morphological traits. Hence, analysis of life-history traits of bee communities can help assess the pollination services they are likely to provide (when taking into account single-visit pollination efficiency). The ecotone between agricultural fields and surrounding habitats is a major barrier that filters many bee species, particularly with regard to their nesting requirements. Thus, greater attention should be given to management practices that encourage pollinators to live and nest, and not only forage, within fields.

Delivery of crop pollination services is an insufficient argument for wild pollinator conservation.

There is compelling evidence that more diverse ecosystems deliver greater benefits to people, and these ecosystem services have become a key argument for biodiversity conservation. However, it is unclear how much biodiversity is needed to deliver ecosystem services in a cost-effective way. Here we show that, while the contribution of wild bees to crop production is significant, service delivery is restricted to a limited subset of all known bee species. Across crops, years and biogeographical regions, crop-visiting wild bee communities are dominated by a small number of common species, and threatened species are rarely observed on crops. Dominant crop pollinators persist under agricultural expansion and many are easily enhanced by simple conservation measures, suggesting that cost-effective management strategies to promote crop pollination should target a different set of species than management strategies to promote threatened bees. Conserving the biological diversity of bees therefore requires more than just ecosystem-service-based arguments.

Wild pollinators enhance fruit set of crops regardless of honey bee abundance.

The diversity and abundance of wild insect pollinators have declined in many agricultural landscapes. Whether such declines reduce crop yields, or are mitigated by managed pollinators such as honey bees, is unclear. We found universally positive associations of fruit set with flower visitation by wild insects in 41 crop systems worldwide. In contrast, fruit set increased significantly with flower visitation by honey bees in only 14% of the systems surveyed. Overall, wild insects pollinated crops more effectively; an increase in wild insect visitation enhanced fruit set by twice as much as an equivalent increase in honey bee visitation. Visitation by wild insects and honey bees promoted fruit set independently, so pollination by managed honey bees supplemented, rather than substituted for, pollination by wild insects. Our results suggest that new practices for integrated management of both honey bees and diverse wild insect assemblages will enhance global crop yields.