Abstract. Male Neotropical orchid bees (Euglossini ) collect volatile substances (fragrances) from flowers and other sources (e.g., decaying wood) and store them in specialized hind tibial pockets. The accumulated chemicals are later emitted during courtship display, presumably to lure conspecific females for mating. We analyzed tibial fragrances of males of 15 sympatric Panamanian species in the genus Euglossa to test whether communities of euglossine bees are chemically structured, and to elucidate whether male fragrance signals evolve to convey premating isolation. Our analysis revealed substantial chemical disparity among all lineages. Disparity was mediated by compounds that were exclusive to certain species but also by differences in relative quantity of shared compounds. We mapped tibial fragrance compounds present in each species on a DNA-based phylogeny (reconstructed using partial sequences of COI, EF1-a, ArgK, and Pol-II ) and found that most dominant compounds were highly homoplasious. In an analysis of chemical differentiation in relation to phylogenetic divergence through time, disparity was greater than expected from a null model at any point during evolutionary history, suggesting that diversifying selection has shaped fragrance phenotypes. Notably, chemical disparity was greater within recently diverged lineages than among them, suggesting that chemical preferences in orchid bees evolved rapidly in the early stages of species divergence. We postulate communication interference as the possible mechanism behind the observed fragrance differentiation, which may be the product of reproductive character (fragrance) displacement. Our findings are consistent with the hypothesis that male fragrance signals evolve to convey premating isolation.

Saltational changes may underlie the diversification of pheromone communication systems in insects, which are normally under stabilizing selection favoring high specificity in signals and signal perception. In orchid bees (Euglossini), the production of male signals depends on the sense of smell: males collect complex blends of volatiles (perfumes) from their environment, which are later emitted as pheromone analogs at mating sites. We analyzed the behavioral and antennal response to perfume components in two male morphotypes of Euglossa cf. viridissima from Mexico, which differ in the number of mandibular teeth. Tridentate males collected 2-hydroxy-6-nona-1,3-dienyl-benzaldehyde (HNDB) as the dominant component of their perfume. In bidentate males, blends were broadly similar but lacked HNDB. Population genetic analysis revealed that tri- and bidentate males belong to two reproductively isolated lineages. Electroantennogram tests (EAG and GC-EAD) showed substantially lower antennal responses to HNDB in bidentate versus tridentate males, revealing for the first time a mechanism by which closely related species acquire different chemical compounds from their habitat. The component-specific differences in perfume perception and collection in males of two sibling species are in agreement with a saltational, olfaction-driven mode of signal perfume evolution. However, the response of females to the diverged signals remains unknown.

Male orchid bees (Euglossini) collect fragrances from flowers and other natural sources, a behaviour that has shaped the euglossine pollination syndrome. Males store such chemicals in hind leg pouches and later expose them during courtship display. In the present study, we show that complex bouquets of two sympatric species of Eulaema, E. meriana and E. bombiformis, are chemically distinct. When exposed during bioassays at display perches individual hind leg extracts rapidly and consistently attracted other males of the correct species, even if derived from males of disparate localities (French Guiana and Panama). Conspecific males as well as females of E. bombiformis arrived at natural perch sites only from downwind, and two copulations were observed. Our findings demonstrate that acquired odours mediate exclusive attraction within species and support the idea that such fragrances are pheromone analogues. Their role in acquiring matings and during male-male interaction is discussed.

In orchid bees, males signal their availability as mates by fanning ‘perfumes’, i.e. blends of volatiles that are collected from environmental sources and stored in hind leg pouches. The chemical composition of such perfumes in males with either two or three mandibular teeth has previously led to the discovery of two sympatric, cryptic lineages within Euglossa viridissima Friese on the Yucatan peninsula, Mexico. Here, we combine chemical, morphological, and genetic data for an integrated characterization of the two lineages. The lectotype of E. viridissima Friese in the Museum of Natural History in Vienna has two mandibular teeth, and the species name viridissima must thus be assigned to the predominantly bidentate lineage, whereas the completely tridentate lineage is described as a novel species, Euglossa dilemma sp. nov. Bembé & Eltz. Chemical profiling and microsatellite genotyping revealed that E. viridissima males can occasionally (∼10% of individuals) express a third mandibular tooth, but this tooth is not positioned centrally on the mandible as in E. dilemma, but is displaced towards the tip. Thus, males of the two lineages can be unambiguously diagnosed by mandibular characters alone. Based on 889 bp of CO1 sequence data, we confirm that E. viridissima and E. dilemma constitute a monophyletic group within the genus Euglossa. However, CO1 alone failed to separate these two lineages due to the lack of parsimony‐informative sites. Both species occur in broad sympatry across Central America, but the orchid bees recently introduced to Florida have three mandibular teeth in males, i.e. belong to E. dilemma. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, 1064–1076.

Habitat degradation and fragmentation are widespread phenomena in tropical regions. Negative effects on the biota are numerous, ranging from interruption of gene flow among populations, to the loss of genetic diversity within populations, to a decline in species richness over time. Orchid bees (Hymenoptera: Apidae: Euglossini) are of major conservation interest due to their function as pollinators of numerous orchid species and other tropical plants. Here, we used microsatellite markers to investigate the effects of geographic distance and habitat fragmentation on gene flow among populations. Populations of Euglossa dilemma in three geographic regionsthe Yucatán peninsula (Mexico), Veracruz (Mexico), and Florida (USA)-were genetically structured predominantly across the regions, with the strength of differentiation among populations being positively correlated with geographic distance. Within geographic regions only little substructure was found, suggesting that dispersal is substantial in the absence of geographic or ecological barriers. In a second study, patterns of genetic differentiation among eight species of Euglossa were not related to habitat fragmentation following deforestation in southern Mexico (Veracruz). Specifically, most bee populations in the 9,800 ha forest remnant of Los Tuxtlas (Volcano San Martin) were neither differentiated from, nor had less genetic diversity than, populations in near-continuous forest separated from Los Tuxtlas by 130 km of agricultural land. Either occasional long distance dispersal across open areas has buffered the expected genetic effects of fragmentation, or the history of fragmentation in southern Mexico is too recent to have caused measurable shifts in allelic composition.

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Transcription factors of the SoxD protein family have previously been shown to prevent precocious specification and terminal differentiation of oligodendrocyte progenitor cells in the developing spinal cord. Using mice with specific deletion of the SoxD proteins Sox5 and Sox6 in the central nervous system, we now show that SoxD proteins additionally influence migration of oligodendrocyte progenitors in the spinal cord as well as in the forebrain. In mutant mice, emigration of oligodendrocyte progenitors from the ventricular zone and colonization of the mantle zone are significantly delayed probably because of reduced expression of Pdgf receptor alpha and decreased responsiveness toward Pdgf-A as a main migratory cue. In addition to this direct cell-autonomous effect on Pdgf receptor alpha expression, SoxD proteins furthermore promote oligodendroglial migration by keeping the cells in an undifferentiated state and preventing a premature loss of their migratory capacity. This indirect effect becomes particularly important during late embryonic and early postnatal phases of oligodendroglial development. Finally, we show that Sox5 and Sox6 cooperate with Sox9 and Sox10 to activate Pdgf receptor alpha expression and thereby maintain oligodendrocyte progenitors in the immature state. This contrasts with their behavior on myelin genes where they antagonize the function of SoxE proteins. It argues that SoxD proteins can function either as repressors or as co-activators of SoxE proteins thereby modulating their function in a stage-specific manner.

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