Global stingless bee phylogeny supports ancient divergence,vicariance, and long distance dispersal

Stingless bees (Meliponini) are one of only two highly eusocial bees, the other being the well studied honey bee (Apini). Unlike Apini, with only 11 species in the single genus Apis, stingless bees are a large and diverse taxon comprising some 60 genera, many of which are poorly known. This is the first attempt to infer a phylogeny of the group that includes the world fauna and extensive molecular data. Understanding the evolutionary relationships of these bees would provide a basis for behavioural studies within an evolutionary framework, illuminating the origins of complex social behaviour, such as the employment of dance and sound to communicate the location of food or shelter. In addition to a global phylogeny, we also provide estimates of divergence times and ancestral biogeograhic distributions of the major groups. Bayesian and maximum likelihood analyses strongly support a principal division of Meliponini into Old and New World groups, with the Afrotropical+Indo‐Malay/Australian clades comprising the sister group to the large Neotropical clade. The meliponine crown clade is inferred to be of late Gondwanan origin (approximately 80 Mya), undergoing radiations in the Afrotropical and Indo‐Malayan/Australasian regions, approximately 50–60 Mya. In the New World, major diversifications occurred approximately 30–40 Mya. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 206–232.     

Climate,physiological tolerance and sex‐biased dispersal shape genetic structure of Neotropical orchid bees

Understanding the impact of past climatic events on the demographic history of extant species is critical for predicting species' responses to future climate change. Palaeoclimatic instability is a major mechanism of lineage diversification in taxa with low dispersal and small geographical ranges in tropical ecosystems. However, the impact of these climatic events remains questionable for the diversification of species with high levels of gene flow and large geographical distributions. In this study, we investigate the impact of Pleistocene climate change on three Neotropical orchid bee species (Eulaema bombiformis, E. meriana and E. cingulata) with transcontinental distributions and different physiological tolerances. We first generated ecological niche models to identify species‐specific climatically stable areas during Pleistocene climatic oscillations. Using a combination of mitochondrial and nuclear markers, we inferred calibrated phylogenies and estimated historical demographic parameters to reconstruct the phylogeographical history of each species. Our results indicate species with narrower physiological tolerance experienced less suitable habitat during glaciations and currently exhibit strong population structure in the mitochondrial genome. However, nuclear markers with low and high mutation rates show lack of association with geography. These results combined with lower migration rate estimates from the mitochondrial than the nuclear genome suggest male‐biased dispersal. We conclude that despite large effective population sizes and capacity for long‐distance dispersal, climatic instability is an important mechanism of maternal lineage diversification in orchid bees. Thus, these Neotropical pollinators are susceptible to disruption of genetic connectivity in the event of large‐scale climatic changes.     

Long Tongues and Loose Niches: Evolution of Euglossine Bees and Their Nectar Flowers1

I examined relationships between tongue length of orchid bees (Apidae: Euglossini) and nectar spur length of their flowers in the genera Calathea, Costus, and Dimerocostus using phylogenetically independent contrasts. Long‐tubed flowers have specialized on one or several species of long‐tongued euglossine bees, but long‐tongued bees have not specialized on long‐tubed flowers. Whereas long tongues may have evolved to provide access to a wider variety of nectar resources, long nectar spurs may be a mechanism for flowers to conserve nectar resources while remaining attractive to traplining bee visitors.     

Specialized pollination by carpenter bees in Calanthe striata (Orchidaceae), with a review of carpenter bee pollination in orchids

Calanthe striata has nectarless flowers that are self‐compatible, but pollinator dependent. Field observations showed that the flowers were pollinated exclusively by the carpenter bee Xylocopa appendiculata circumvolans, although the bees occasionally wasted pollen by delivering to the stigmatic surface pollinaria that retained their anther caps. Fruit set ratios at the population level varied spatiotemporally, but were generally low (8.3–17.3%). Calanthe striata blooms in spring when post‐overwintering carpenter bees have not yet started foraging for brood production. It can therefore exploit an abundance of opportunistic/naïve foragers. This timing may also increase the possibility of pollinator visits, because no rewarding co‐flowering plants are available in the orchid habitats. A literature review of Orchidaceae pollinated by carpenter bees revealed that at least 14 species of Orchidoideae and Epidendroideae have evolved flowers specialized for carpenter bee pollination. They typically have shallow pink/magenta flowers with a foothold for pollinators; pollinaria are attached to the head, ventral thorax or base of the middle legs of carpenter bees when they insert their heads and/or proboscises into flowers; pollination success is generally low, a probable consequence of the deceptive pollination systems. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 730–743.     

Long-term ecology of euglossine orchid-bees (Apidae: Euglossini) in Panama

Summary Abundance patterns during 6–7 years and orchid visitation were determined for 51 species of the 57 local euglossine bees. Male bees were counted at 3 chemical attractants presented in the same manner each month. Sites were separated by 75 km but included wet Atlantic forest at 500 m elevation, moist forest at 180 m near Barro Colorado Island, and cloud forest at 900 m near the Pacific ocean. 1. From 15 to 30 euglossine species of 4 genera were active in each month and site; monthly species number and general bee abundance were positively correlated. Many species had 3 annual abundance peaks (range 1–4) and were active throughout the year, but peak annual abundances rarely occurred during late wet or early dry seasons. In contrast, Eufriesea generally were present as adults only 1–2 months in a year. 2. Euglossine populations were exceptionally stable. Species at each site were more stable than any known insect population, and stability and abundance were positively associated. However, year-to-year population stability and the degree of seasonality were not correlated. Among the three sites, the more diverse (species rich) bee assemblages displayed lower stability; these were the wetter and upland sites. 3. The most abundant bees visited more orchid species. Eg. and El. each visited and average of 4 orchid species (range 0–13); Ex. and Ef. visited 0–3. Stable populations did not visit more or fewer orchid species than did unstable populations. 4. Less than 68% of species at each site visited orchid flowers; less than a few dozen of the 100–800 bees counted in a day carried orchid pollinaria. Over 20% of the euglossine species never were seen with pollinaria at any site and probably seldom visit orchids in central Panama. 5. Most bee species visited 1 or no fragrance orchids in a given habitat. Orchids tended to utilize common pollinators that seldom included more than 1 species, and they utilized stable or unstable, seasonal or aseasonal bees. However, the most stable and abundant bee, Eg. imperialis, rarely pollinated orchids; fewer than 10 of ca. 20000 bees carried pollinaria. 6. Orchids may interact primarily with discrete seasonal bee population peaks-probably the emerging adults. Although specialized orchid preferences are implicated for species that visit few or no local orchids but pollinate other species and carry pollinaria in other areas, euglossine bees do not need orchids to survive or reproduce.     

Visual signalling of nectar‐offering flowers and specific morphological traits favour robust bee pollinators in the mass‐flowering tree Handroanthus impetiginosus (Bignoniaceae)

Mass flowering is a widespread blooming strategy among Neotropical trees that has been frequently suggested to increase geitonogamous pollination. We investigated the pollination ecology of the mass‐flowering tree Handroanthus impetiginosus, addressing its breeding system, the role in pollination of different visitors, the impact of nectar robbers on fruit set and the function of colour changes in nectar guides. This xenogamous species is mainly pollinated by Centris and Euglossa bees (Apidae) seeking nectar, which are known to fly long distances. The flowers favour these bees by having: (1) a closed entrance in newly opened flowers which provides access only to strong bees capable of deforming the flower tube; and (2) a nectar chamber that is accessible only to long‐tongued bees. Only first‐day flowers with yellow nectar guides produce nectar. Pollinators prefer these flowers over second‐ and third‐day flowers with orange and red nectar guides, respectively. Nectar robbers damage two‐thirds of the flowers and this robbing activity decreases fruit set by half. We attribute the low fruit set of H. impetiginosus to the intense nectar robbing and hypothesize that visual signalling of nectar presence in newly opened (receptive) flowers reduces geitonogamy by minimizing bee visits to unrewarding (non‐receptive) flowers. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 396–407.     

Environmental determinants of genetic diversity in Caragana microphylla (Fabaceae) in northern China

Non‐rewarding orchids rely on various ruses to attract their pollinators. One of the most common is for them to resemble flowers sought by insects as food sources. This can range from generalized food deception to the mimicry of specific sympatric food plants. We investigated the basis of pollinator deception in the European food‐deceptive orchid Traunsteinera globosa, which has unusually compact flowerheads resembling those of sympatric rewarding species of Knautia and Scabiosa (Dipsacaceae), and Valeriana (Caprifoliaceae). Visual signals of T. globosa are similar in both fly and bee vision models to those of the sympatric food plants used in the choice experiments, but scent signals are divergent. Field experiments conducted in Austria and the Czech Republic showed that both naive and experienced (with respect to visitation of T. globosa) insect species approached the orchids at the same rate as food plants, but direct contact with orchid flowers was taxon specific. Flies were most easily duped into probing the orchid, and, in doing so, frequently received and deposited pollinaria, whereas most bees and butterflies avoided landing on orchid flowers. We conclude that T. globosa is a mimic of a guild of fly‐pollinated plants, but the ecological dependence of the orchid on its models remains to be fully tested. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 269–294.     

Solitary invasive orchid bee outperforms co-occurring native bees to promote fruit set of an invasive Solanum

Our understanding of the effects of introduced invasive pollinators on plants has been exclusively drawn from studies on introduced social bees. One might expect, however, that the impacts of introduced solitary bees, with much lower population densities and fewer foragers, would be small. Yet little is known about the potential effects of naturalized solitary bees on the environment. We took advantage of the recent naturalization of an orchid bee, Euglossa viridissima, in southern Florida to study the effects of this solitary bee on reproduction of Solanum torvum, an invasive shrub. Flowers of S. torvum require specialized buzz pollination. Through timed floral visitor watches and two pollination treatments (control and pollen supplementation) at three forest edge and three open area sites, we found that the fruit set of S. torvum was pollen limited at the open sites where the native bees dominate, but was not pollen limited at the forest sites where the invasive orchid bees dominate. The orchid bee’s pollination efficiency was nearly double that of the native halictid bees, and was also slightly higher than that of the native carpenter bee. Experiments using small and large mesh cages (to deny or allow E. viridissima access, respectively) at one forest site indicated that when the orchid bee was excluded, the flowers set one-quarter as many fruit as when the bee was allowed access. The orchid bee was the most important pollinator of the weed at the forest sites, which could pose additional challenges to the management of this weed in the fragmented, endangered tropical hardwood forests in the region. This specialized invasive mutualism may promote populations of both the orchid bee and this noxious weed. Invasive solitary bees, particularly species that are specialized pollinators, appear to have more importance than has previously been recognized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Distribution,habitat disturbance and pollination of the endangered orchid Broughtonia cubensis (Epidendrae: Laeliinae)

The geographical distribution, population structure and pollination ecology are key aspects in the conservation and management of rare orchids. Here, we address these aspects and the main threats affecting the endangered Cuban orchid Broughtonia cubensis. This rewardless orchid is self‐compatible, but pollinator dependent. However, seed production can be negatively affected by insect‐mediated selfing. Three species of small bee (genera Ceratina and Lasioglossum) act as pollinators. As in the case of other nectarless orchids, we detected two species of plant producing large amounts of nectar in the area, the floral morphology of which closely resembles that of B. cubensis. The simultaneous flowering of these species could positively affect the reproductive success of B. cubensis. Nonetheless, the fitness of this orchid in natural conditions is low, possibly related to strong pollen limitation. To the problems arising from reduced fitness is added the fact that its historical distribution range has been greatly reduced in recent years. Throughout this study, we have detected dramatic reductions in the population sizes, in some cases as a result of human plundering, but also as a consequence of hurricanes. Based on the results of this study, we propose some guidelines to manage and conserve this orchid. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 345–357.     

Seasonal cycles,phylogenetic assembly,and functional diversity of orchid bee communities

Neotropical rainforests sustain some of the most diverse terrestrial communities on Earth. Euglossine (or orchid) bees are a diverse lineage of insect pollinators distributed throughout the American tropics, where they provide pollination services to a staggering diversity of flowering plant taxa. Elucidating the seasonal patterns of phylogenetic assembly and functional trait diversity of bee communities can shed new light into the mechanisms that govern the assembly of bee pollinator communities and the potential effects of declining bee populations. Male euglossine bees collect, store, and accumulate odoriferous compounds (perfumes) to subsequently use during courtship display. Thus, synthetic chemical baits can be used to attract and monitor euglossine bee populations. We conducted monthly censuses of orchid bees in three sites in the Magdalena valley of Colombia – a region where Central and South American biotas converge – to investigate the structure, diversity, and assembly of euglossine bee communities through time in relation to seasonal climatic cycles. In particular, we tested the hypothesis that phylogenetic community structure and functional trait diversity changed in response to seasonal rainfall fluctuations. All communities exhibited strong to moderate phylogenetic clustering throughout the year, with few pronounced bursts of phylogenetic overdispersion that coincided with the transition from wet‐to‐dry seasons. Despite the heterogeneous distribution of functional traits (e.g., body size, body mass, and proboscis length) and the observed seasonal fluctuations in phylogenetic diversity, we found that functional trait diversity, evenness, and divergence remained constant during all seasons in all communities. However, similar to the pattern observed with phylogenetic diversity, functional trait richness fluctuated markedly with rainfall in all sites. These results emphasize the importance of considering seasonal fluctuations in community assembly and provide a glimpse to the potential effects that climatic alterations may have on both pollinator communities and the ecosystem services they provide.     

CONSERVATION GENETICS OF NEOTROPICAL POLLINATORS REVISITED: MICROSATELLITE ANALYSIS SUGGESTS THAT DIPLOID MALES ARE RARE IN ORCHID BEES

Allozyme analyses have suggested that Neotropical orchid bee (Euglossini) pollinators are vulnerable because of putative high frequencies of diploid males, a result of loss of sex allele diversity in small hymenopteran populations with single locus complementary sex determination. Our analysis of 1010 males from 27 species of euglossine bees sampled across the Neotropics at 2–11 polymorphic microsatellite loci revealed only five diploid males at an overall frequency of 0.005 (95% CIs 0.002–0.010); errors through genetic nondetection of diploid males were likely small. In contrast to allozyme‐based studies, we detected very weak or insignificant population genetic structure, even for a pair of populations >500 km apart, possibly accounting for low diploid male frequencies. Technical flaws in previous allozyme‐based analyses have probably led to considerable overestimation of diploid male production in orchid bees. Other factors may have a more immediate impact on population persistence than the genetic load imposed by diploid males on these important Neotropical pollinators.     

Buzz-pollination in Neotropical bees:genus-dependent frequencies and lack of optimal frequency for pollen release

Over 50 genera of bees release pollen from flower anthers using thoracic vibrations,a phenomenon known as buzz-pollination.The efficiency of this process is directly affected by the mechanical properties of the buzzes,namely the duration,amplitude,and frequency.Nonetheless,although the effects of the former two properties are well described,the role of buzz frequency on pollen release remains unclear.Furthermore,nearly all of the existing studies describing vibrational properties of natural buzz-pollination are limited to bumblebees(Bombus)and carpenter bees(Xvlocopa)constraining our current understanding of this behavior and its evolution.Therefore,we attempted to minimize this shortcoming by testing whether flower anthers exhibit optimal frequency for pollen release and whether bees tune their buzzes to match these(optimal)frequencies.If true,certain frequencies will trigger more pollen release and lighter bees will reach buzz frequencies closer to this optimum to compensate their smaller buzz amplitudes.Two strategies were used to test these hypotheses:(i)the use of(artificial)vibrational playbacks in a broad range of buzz frequencies and amplitudes to assess pollen release by tomato plants(Solarium Ivcopersicum L.)and(ii)the recording of natural buzzes of Neotropical bees visiting tomato plants during pollination.The playback experiment indicates that although buzz frequency does affect pollen release,no optimal frequency exists for that.In addition,the recorded results of natural buzz-pollination reveal that buzz frequencies vary with bee genera and are not correlated with body size.Therefore,neither bees nor plants are tuned to optimal pollen release frequencies.Bee frequency of buzz-pollination is a likely consequence of the insect flight machinery adapted to reach higher accelerations,while flower plant response to buzz-pollination is the likely result of its pollen granular properties.     

Generic recircumscriptions of Oncidiinae (Orchidaceae: Cymbidieae) based on maximum likelihood analysis of combined DNA datasets

Phylogenetic relationships within the orchid subtribe Oncidiinae sensu Chase were inferred using maximum likelihood analyses of single and multilocus DNA sequence data sets. Analyses included both nuclear ribosomal internal transcribed spacer DNA and plastid regions (matK exon, trnH‐psbA intergenic spacer and two portions of ycf1 exon) for 736 individuals representing approximately 590 species plus seven outgroup taxa. Based on the well resolved and highly supported results, we recognize 61 genera in Oncidiinae. Mimicry of oil‐secreting Malpighiaceae and other floral syndromes evolved in parallel across the subtribe, and many clades exhibit extensive variation in pollination‐related traits. Because previous classifications heavily emphasized these floral features, many genera recognized were not monophyletic. Our classification based on monophyly will facilitate focused monographs and clarifies the evolution of morphological and biochemical traits of interest within this highly diverse subtribe. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 117–146.     

Revision and phylogeny of the bee genus Paratetrapedia Moure,with description of a new genus from the Andean Cordillera (Hymenoptera,Apidae, Tapinotaspidini)

The bee genus Paratetrapedia represents a commonly collected group of bees and is especially diverse in forested areas of the Neotropics. Its taxonomy has remained poorly understood because of a lack of modern revisionary work and numerous species described as Tetrapedia whose type specimens have not been re‐examined in recent times. Here, a comprehensive study was carried out to review the taxonomy of the genus Paratetrapedia and to investigate cladistically the relationships amongst its species. Eighteen new species of Paratetrapedia are described, giving a total of 32 species in the genus. A phylogenetic analysis of the species of Paratetrapedia was carried out using 61 morphological characters for 41 terminal taxa. The phylogenetic results confirmed the monophyly of Paratetrapedia and allowed the recognition of five species groups: the lugubris, moesta, bicolor, lineata, and flavipennis groups. Nasutopedia gen. nov. , recognized as the sister group of Paratetrapedia and with its distribution restricted to the western forested portions of the Andean highlands, is proposed based on distinct morphology, its placement in the phylogenetic tree, and biogeographical patterns. Species of Paratetrapedia are especially diverse in the Amazon Forest; the eastern Brazilian Atlantic Forest contains four endemic species, and one species is endemic to the Cerrado of central Brazil. Paratetrapedia shows a biogeographical pattern similar to other Neotropical groups of bees and birds, with wide distribution and high diversity in lowland forests and whose sister taxon occurs on highlands of north‐western portions of the Andean cordillera. Identification keys for males and females of all species are provided, as well as distribution maps and illustrations of general external morphology and genitalia. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 351–442.     

Does bee or wasp mimicry by orchid flowers also deter herbivores?

General visual bee mimicry and specific chemical mimicry by flowers to solitary female bees or wasps are well known in several orchid genera, for example, the Mediterranean genus Ophrys, the Australian genera Cryptostylis and Chiloglottis, and the South-African Disa. This mimicry has been shown to attract solitary male bees or wasps, which are their species-specific pollinators. The visual and chemical signals are considered to be a type of deceptive pollination mechanism based on mimicry for the exploitation of perceptual biases of animals. We propose that in addition to this unique pollination mechanism, these plants exhibit another, rarely mentioned and practically forgotten, non-exclusive function of bee or wasp mimicry (Batesian mimicry). This mimicry may deter large mammalian herbivores, and possibly also insects from the plants and especially from their flowers by a type of visual and olfactory deceptive aposematism. While visiting the flowers, bees and wasps may add a Müllerian effect to this defense. We extend this hypothesis to many other rewarding flowers that are bee or wasp pollinated and propose that abundance of pollinating bees or wasps may deter herbivorous mammals and insects from the plants during their peak flowering season.     

Host range evolution in a selected group of osmiine bees (Hymenoptera: Megachilidae): the Boraginaceae‐Fabaceae paradox

Bees are extraordinarily diverse with respect to host plant choice and adaptation. Recent findings suggest that bee host range might be largely governed by evolutionary constraints related to pollen digestion or flower recognition and handling. In the present study, we applied phylogenetic inference to investigate whether such constraints underlie host plant choice in bees of the Annosmia‐Hoplitis group (Megachilidae) and to what extent these bees have evolved specialized adaptations for pollen collection. We demonstrate that most pollen specialist species exclusively exploit either Boraginaceae or Fabaceae, whereas all pollen generalists harvest pollen from both Boraginaceae and Fabaceae. The counterintuitive affinity towards these two plant families, which are neither closely related nor share similar flower morphologies, demonstrates that pollen host choice is considerably constrained in this group of bees. We hypothesize that this Boraginaceae‐Fabaceae paradox might be the result of (1) similar secondary metabolites in the pollen of both families; (2) metabolites that can be detoxified by the same physiological tools; or (3) similar pollen nutrient composition. Contrary to the widely held belief that specialized adaptations for pollen collection are rare among bees, such adaptations are common in the Annosmia‐Hoplitis bees, where they have evolved several times independently to exploit flowers of widely different morphologies. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Genome size variation in Orchidaceae subfamily Apostasioideae: filling the phylogenetic gap

With more than 160‐fold variation, Orchidaceae are currently the most diverse angiosperm family with respect to the amount of nuclear DNA. This study provides first genome size estimates for approximately 50% of species currently recognized in subfamily Apostasioideae, which is sister to the other four orchid subfamilies. The estimated 1C‐values range from 0.38 pg in Apostasia nuda to 5.96 pg in Neuwiedia zollingeri var. javanica, a nearly 16‐fold range. The two genera show non‐overlapping genome sizes, with those in Apostasia being distinctly smaller than those in Neuwiedia. In fact, most Apostasia spp. are at the lower end of the range of orchid C‐values. Observed discontinuities in DNA amounts in genera most probably reflect interspecific variation in ploidy. In addition to ploidy heterogeneity in N. zollingeri var. javanica, intraspecific variation in genome size (up to 17.7%) was also detected in some species; this can be plausibly related to the incidence of different geographical variants or unrecognized taxonomic heterogeneity. The AT content varied from 62.6 to 66.0%, which is in the upper range recorded for angiosperms. The genome size data obtained in this study fill a major phylogenetic gap in Orchidaceae and show that (very) small genomes prevail in subfamily Apostasioideae. © 2013 The Linnean Society of London     

Single mating in orchid bees (Euglossa, Apinae): implications for mate choice and social evolution

Neotropical orchid bees (Euglossini) are conspicuously different from other corbiculate bees (Apinae) in their lack of advanced sociality and in male use of acquired odors (fragrances) as pheromone-analogues. In both contexts, orchid bee mating systems, in particular the number of males a female mates with, are of great interest but are currently unknown. To assess female mating frequency in the genus Euglossa, we obtained nests from three species in Mexico and Panama and genotyped mothers and their brood at microsatellite DNA loci. In 26 out of 29 nests, genotypes of female brood were fully consistent with being descended from a singly mated mother. In nests with more than one adult female present, those adult females were frequently related, with genotypes being consistent with full sister–sister (r = 0.75) or mother–daughter (r = 0.5) relationships. Thus, our genetic data support the notions of female philopatry and nest-reuse in the genus Euglossa. Theoretically, single mating should promote the evolution of eusociality by maximizing the relatedness among individuals in a nest. However, in Euglossini this genetic incentive has not led to the formation of eusocial colonies as in other corbiculate bees, presumably due to differing ecological or physiological selective regimes. Finally, monandry in orchid bees is in agreement with the theory that females select a single best mate based on the male fragrance phenotype, which may contain information on male age, cognitive ability, and competitive strength.     

Cover Caption

Tomato flowers are pollinated by bees that vibrate their thorax for pollen collecting, behavior known as buzz‐pollination. The phenomenon is common and a specimen of the Neotropical stingless bee Melipona quadrifasciata is here depicted visiting a tomato flower for pollen collecting. While testing whether flower anthers exhibit optimal frequency for pollen release and whether flower bees tune their buzzes to match these frequencies, we recognized that neither bees nor plants are tuned to optimal pollen release frequencies (see pages 133–142). Photo provided José Lino‐Neto.     

A humped latitudinal phylogenetic diversity pattern of orchid bees (Hymenoptera: Apidae: Euglossini) in western Amazonia: assessing the influence of climate and geologic history

Amazonian rainforests are among the most species‐rich terrestrial habitats on Earth. The aim of this study was to analyze phylogenetic diversity (PD) patterns of orchid bee assemblages along a latitudinal gradient of 15° from northern Peru to central Bolivia and to relate them to climatic factors and geological history. We expanded an existing phylogeny of orchid bees and analyzed the PD of 15 orchid bee assemblages along a latitudinal gradient using mean pair‐wise phylogenetic distance. The resulting pattern was correlated to climatic factors and elevation. We found a hump‐shaped pattern of PD that peaked in central Peru and decreased towards the equatorial and especially towards the southern, subtropical sites. The decrease in PD towards higher latitudes is a common pattern found in many taxa, which in our case correlates with increasing climatic seasonality. However, the decrease towards the equatorial sites is unusual and may be related to a particular historic event: the northern sites with low PD are situated in the area of the former Lake Pebas, which covered western Amazonia until 3 mya. After the lake disappeared orchid bees mainly belonging to two distantly related species groups apparently colonized the region and diversified, which led to the comparatively low observed PD. In contrast, in central Peru, no in situ radiations were detected, hence the assemblages were composed of species from diverse phylogenetic lineages. Additionally, we identified multiple phylogenetically independent radiations of regionally restricted Euglossa species along the latitudinal gradient that, according to a published, dated phylogeny, diversified roughly 3–1 mya. The hump‐shaped latitudinal pattern of PD of the orchid bees of western Amazonia thus appears to have resulted from a preponderance of early divergent lineages in central Peru and of young radiations from distantly related clades colonizing higher latitudes, possibly triggered by historic climate fluctuations and major geological events.