Patterns of insect flower visitation in Lavandula buchii Webb (Lamiaceae), an endemic shrub of Tenerife (Canary Islands)

The foraging ecology of insect visitors of the endemic lavender Lavandula buchii (Lamiaceae) was studied in a relict population of the plant in north-west Tenerife (Canary Islands). Timed observations were used to describe this pollination system in terms of the structure of the visitor assemblage, foraging patterns and efficiencies of the main insect species. The visitor pool was dominated by anthophorid solitary bees. Furthermore, it was characterized by a high level of endemicity and a small number of species compared with continental relatives of Lavandula. The main visitors, two anthophorids and a bombyliid fly, clearly differed in their foraging behaviour, i.e. visitation frequency, visit duration, rate of flower probing, and in anatomical traits (body size and proboscis length). In these terms, the bees were by far more efficient foragers than the fly. These results are discussed in the context of insect-flower interactions in insular pollination systems.     

Satellite flies (Leucophora personata,Diptera: Anthomyiidae) and other dipteran parasites of the communal bee Andrena agilissima (Hymenoptera: Andrenidae) on the island of Elba,Italy

Solitary and presocial aculueate Hymenoptera are parasitized by a range of dipteran species in the families Anthomyiidae, Bombyliidae, Conopidae, Phoridae, and Sarcophagidae that are likely to impact on their hosts. We undertook a study over several years of a univoltine and communal bee, Andrena agilissima, and its main dipteran parasites, in particular the satellite fly Leucophora personata (Diptera: Anthomyiidae). Behavioural and ecological data were collected from one nesting aggregation of the host bee on the island of Elba, Italy, from 1993 to 2003, and from a foraging site of the bee, ca 5?km from the nesting aggregation. Other Diptera associated with A. agilissima at the field site were the bee fly Bombylius fimbriatus (Bombyliidae), the conopid fly Zodion cinereum (Conopidae), and the scuttle fly Megaselia andrenae (Phoridae). The phenology of the Diptera broadly overlapped with that of their host across the season of activity (end of April and all of May). Diurnal activity patterns differed slightly; L. personata in particular was active at the host's nesting site before A. agilissima. Female satellite flies also showed a range of behaviours in gaining entry to a host nest. We summarize published data on this and other Leucophora species that parasitize Andrena host bees. Host bees returning to their nests occasionally undertook zig‐zag flight manoeuvres if followed by a satellite fly that were generally successful in evading the fly. Satellite flies that entered a nest, presumably to oviposit, were less likely to remain therein if another host bee entered the same nest, suggesting that one advantage to communal nesting for this host is a reduction in brood cell parasitism by L. personata. We provide the first clear evidence for parasitism by a Zodion of any Andrena host. Both L. personata and M. andrenae concentrated their parasitic activities in the zone of the host nesting aggregation with highest nest densities. Three of the Diptera, L. personata, B. fimbriatus, and Z. cinereum, seemed to have extremely low rates of parasitism whilst that of M. andrenae appeared low. Though they have refined parasitic behaviour that allows them to gain entry into host nests (L. personata, B. fimbriatus, and M. andrenae) or to parasitize adults (Z. cinereum), these parasites seem not to impact upon the dynamics of the host A. agilissima at the nesting aggregation, and the host possesses traits to reduce parasitism.     

Taxonomic diversity,niche width and similarity in the use of plant resources by bees (Hymenoptera: Anthophila) in a cerrado area

Most of the studies on the bee fauna of cerrado have shown incomplete approaches, focusing on species diversity or on a fraction of the assemblage. This work presents similarities among bee species based on visited plants and also shows taxonomic diversity of bees for plant resource sets offered by species in a cerrado area (Pé-de-Gigante), one of the few conservation areas of cerrado in the São Paulo State, Brazil. A total of 737 specimens, belonging to 71 bee species, were collected visiting 52 plant species. Species clusters based on visited plants show that species of Epicharis, for example, are more similar among themselves than from species of other genera. In addition, bee assemblage showed a nested structure, indicating the presence of specialist species exploiting resources subsets used by generalist species under strong intra-specific and weak inter-specific competitions.     

An overview of female reproductive traits in South American Mabuya (Squamata,Scincidae), with emphasis on brood size and its correlates

The viviparous New World Mabuya (Reptilia, Scincidae) are characterized by a peculiar suite of reproductive traits associated with an extreme degree of matrotrophy and a specialized placenta. We gathered information on female reproductive traits of South American Mabuya based on a compilation of literature data plus original information on two species (Mabuya dorsivittata and Mabuya macrorhyncha). Considering brood size, we identified one “large-brooded” group (brood sizes up to eight to ten) and one “small-brooded” group (brood sizes up to four to six). Brood size tended to be positively correlated with female body size among species, although the correlation is not as strong as it is within species. There were some general trends for the reproductive ecology of New World Mabuya, such as early sexual maturation, brood sizes of usually four to six young, and parturition occurring around the dry–wet season transition.     

Time-lapse photography reveals the occurrence of unexpected bee-pollination in Calanthe izuinsularis,an endangered orchid endemic to the Izu archipelago

Calanthe izuinsularis is a rare, beautiful and fragrant orchid endemic to the Izu archipelago. Although it is known that mainland populations of closely related Calanthe species are pollinated by medium- to large-sized bees, it is likely that C. izuinsularis has been forced to alter its floral biology to attract alternative pollinators, as large-sized bees are rarely found on the Izu Islands. Indeed, the unusual floral characteristics of C. izuinsularis, which produces pale flowers that emit a strong and fragrant scent during the night, are considered the adaptation to a moth-pollinator syndrome. As expected, our time-lapse photography using a digital camera revealed that the inflorescences of C. izuinsularis were most frequently visited by nocturnal moths such as Noctuidae, Geometridae and Crambidae, which accounted for 50% of all insect visits (in terms of both the number of frames with captured visitors and the number of times visited). However, our study could not provide evidence of a pollinator shift towards moths because none of the moth species was observed with pollinaria. On the other hand, our study found that the pollinaria of C. izuinsularis could become attached to the mesothorax of the small-sized sweat bee Lasioglossum occidens. Therefore, our study suggests that C. izuinsularis is still somewhat dependent on bee pollination, while the orchid has changed its floral colouration and scent. The occurrence of bee-pollination in C. izuinsularis would provide an explanation for how the natural hybridisation between C. aristulifera, C. discolor and C. izuinsularis can occur.     

Breeding in the gray-headed tody-flycatcher (Aves: Tyrannidae) with comments on geographical variation in reproductive traits within the genus Todirostrum

The gray-headed tody-flycatcher Todirostrum poliocephalum is a passerine endemic to the Atlantic Forest of Brazil. We describe the nest and nesting behaviour of this species and investigate geographical variation in breeding traits in the genus using data from the literature, museum collections, and citizen science projects. We located 21 nests of the gray-headed tody-flycatcher, 13 of which we monitored. Nest height above the ground averaged 3.3 ± 3.4 m (n = 21). Nests were built over a 16.8 ± 5.6-day period (n = 6) by both adults. Clutch sizes ranged from two to three eggs, with a mean of 2.9 ± 0.3 eggs (n = 10). Eggs measured 16.6 ± 0.5 × 12.0 ± 0.5 mm (n = 23) and weighed 1.1 ± 0.1 g (n = 19). Mean incubation period was 17 days (n = 3) and mean nestling period was 15.5 days (n = 2). Apparent reproductive success was 30.8%, with predation being the primary cause of nest failure (46.1%). Mayfield’s reproductive success was 25.9%, and daily survival rates for eggs and nestlings were 0.957 and 0.971, respectively. Clutch sizes increased with latitude, but temperature and precipitation seasonality had very low importance in explaining clutch size variation.     

Effects of fragmentation on genetic variation in populations of the terrestrial earthworm Drawida japonica Michaelsen, 1892 (Oligochaeta,Moniligastridae) in Shandong and Liaodong peninsulas,China

Earthworms are excellent indicators of landscape fragmentation due to their widespread distribution and poor dispersal abilities over water. Based on field studies conducted in the Shandong and Liaodong peninsulas and their associated islands during summer 2009, we analysed the genetic variation of Drawida japonica Michaelsen, 1892 (Oligochaeta, Moniligastridae) using DNA sequences of the 16S and 28S ribosomal genes in samples obtained from 13 localities, 12 of which were islands, one artificially connected to the mainland by man (Yangma SC). We identified 55 haplotypes among 79 samples, based on 1094 bp, including 47 unique haplotypes. The phylogenetic tree topologies showed two distinct clades. A multiple-apex Poisson distribution using a mismatch analysis of nucleotides was also detected. Results of a Tajima D neutral test were not significant (p > 0.05). Genetic divergence among populations was much larger than within populations. A positive relationship between genetic diversity and island area was revealed, but no significant correlation between genetic diversity and distance to the nearest larger landmass was found. These results indicate that the intraspecific genetic diversity of D. japonica has been influenced by geographic isolation (i.e. island habitat). Passive transport via anthropogenic activities or nonhuman vertebrates (birds or mammals) may also play a key role in dispersal.     

‘On the rocks’: reproductive biology of the endemic toad Xanthophryne (Anura: Bufonidae) from the Western Ghats,India

Xanthophryne is a toad genus endemic to the northern Western Ghats of India, with two extant sister species – Xanthophryne koynayensis and Xanthophryne tigerina. Both species are local endemics and endangered. We studied reproductive biology of these toads and found that they are specialized to the lateritic rocky outcrops at mid-elevations in high rainfall areas. Xanthophryne toads have sporadic, multiple spawning bouts lasting 2–4 days during early monsoon. In this explosive breeding behaviour, we observed male toads to engage in ‘pelvic thrusts’, a unique and novel behaviour among anurans. Females oviposit in shallow pools in depressions of lateritic boulders where their tadpoles metamorphose. These ephemeral rocky pools have limited resources and they desiccate rapidly with a break in the rains. To mitigate the stochastic risk of desiccation and subsequent large-scale egg/tadpole mortality, females may disperse their reproductive investment spatially and temporally in multiple clutches, and tadpoles metamorphose rapidly. Here, we describe the amplexus, spawning and male advertisement call, and provide a comparative account of the life history traits of the two Xanthophryne species. These toads face numerous threats and are in need of urgent conservation action. These toads, seemingly well adapted to the isolated rocky outcrops, offer an excellent opportunity to understand endemism, mating systems, anuran ecology and behaviour.     

First record of Orbopercularia lichtensteini (Peritrichia: Operculariidae) attached to a hemipteran,Corisella edulis (Nepomorpha: Corixidae), in Hidalgo,Mexico, with notes on its cytology and distributional patterns

Operculariidae includes sessile ciliate species mainly attached to diverse substrates, but records are scarce for their attachment to aquatic true bugs. The goal of this contribution is to provide new cytological data with optical and scanning electron microscopy of Orbopercularia lichtensteini, and some notes about their distribution on a Mexican corixid. We manually collected the corixids in a pond in Mexico during a six-month period, and all individuals were revised to observe the peritrichids. We calculated the abundance and prevalence, with an emphasis on the specific micro-location of the epibiotic peritrich on the host Corisella edulis. A total of 36,205 peritrichids were recorded in 158 corixids. The colonies with fine striated zooids were dichotomously branched and mostly attached to left eyes, but also on other body regions, i.e. left wings and rostrum. We conclude that some morphological colony traits bring some advantages for food particle capture, and aspects of the biology and morphology of Corisella edulis play an important role in determining the spatial distribution of O. lichtensteini.     

Morphology and composition of the internal axis in two morphologically contrasting deep-water sea pens (Cnidaria: Octocorallia)

Umbellula encrinus Linnaeus, 1758 and Anthoptilum grandiflorum (Verrill, 1879) are two species of deep-water sea pens (Cnidaria: Octocorallia) commonly found in the Northwest Atlantic. These sea pens are frequently caught as fishing bycatch, but little is known about their biology. Here we characterise the internal skeleton (axis) of these two morphologically contrasting species, in terms of axis carbonate composition and proportion, morphology and metrics (i.e. cross-sectional area and weight) through gradients along their lengths. Using the conspicuous growth rings present in the axis of U. encrinus we also estimated intra-colonial variation in radial growth rates for this species. X-ray diffraction (XRD) analyses revealed that the carbonate portion of the axis is composed of magnesian calcite (Ca, Mg)CO₃ in both species. On average, the axis is composed of 71% carbonate material in U. encrinus, and 65% carbonate in A. grandiflorum. The carbonate proportion was greatly reduced from base (71%) to tip (33%) in the axis of U. encrinus, while in A. grandiflorum changes were less abrupt (~6% difference from base to tip). In U. encrinus, the axis is quadrangular in shape through its whole length, twisting longitudinally. Twisting can occur in both clockwise and anti-clockwise directions. The axis of A. grandiflorum varied from elliptical near the base to cylindrical through most of the remaining length, with no twisting observed. Axes’ cross-sectional areas also decreased with distance from the base in both species, being the largest near the base. Estimated radial growth rates in U. encrinus were variable within a single colony, ranging between 0.04 and 0.08 mm.yr^?1. This is the first study to describe changes along the axes of sea pens, and to characterise the axes in the studied species. It provides a baseline for studies considering sea pens’ skeletons (e.g. ocean acidification and palaeoceanography).     

The occurrence of facultative paedomorphosis in a lacustrine population of the Pyrenean newt (Calotriton asper): morphology and age structure

Facultative paedomorphosis is a common polyphenism in newts where two alternative phenotypes can coexist: metamorphs and paedomorphs (mature with larval traits, i.e. retaining gills). This phenomenon has been reported in some lacustrine populations of the Pyrenean newt (Calotriton asper). The morphology and life-history traits were studied in a lacustrine population of this species with the occurrence of facultative paedomorphosis. The results showed that 24.3% of adults retained gills at different levels of reabsorption and had a smoother skin compared with the other populations, which were fully metamorphic. The body size and shape showed significant sexual differences, with males being larger than females. The presence of paedomorphic traits also affected the body size and shape, revealing a complex pattern of growth in which metamorphic adults were larger than paedomorphs. The age structure was different between sexes, with a median age of 10.5 and 7 years and longevity of 19 and 14 years in males and females, respectively. Considering each sex separately, the age structure was not significantly different between metamorphic and paedomorphic newts. In addition, metamorphosed and branchiate juveniles were found with a maximum age of 13 and 20 years, respectively. Remarkably, long-lived larvae with an exceptional individual of 18 years were found, which has not previously been reported in any other newt species. The mechanisms that can explain the evolution of paedomorphosis are complex and comprise the interaction of multiple factors related to the cost of reproduction and growth.     

A new Stelis (Dolichostelis) from northern Colombia (Hymenoptera: Megachilidae): first records for South America and a synopsis of the bee fauna from the Caribbean region of Colombia

The cleptoparasitic anthidiine bee genus Stelis Panzer is primarily Holarctic in distribution, extending as far south as Costa Rica. A new species of Stelis subgenus Dolichostelis Parker and Bohart is described and illustrated based on females collected in dry forests of the Caribbean region of Colombia. Stelis (Dolichostelis) costaricensis Friese is also recorded from dry forests but in western Colombia. These are the first records of the genus, as currently recognized, for South America. An updated key to the species of Dolichostelis and a synopsis of the diversity and biogeography of the bee fauna from the Caribbean region of Colombia are provided. Based on reports from the literature and data from collections, we provide a checklist consisting of 125 species in 48 genera of five families. The complex composition of this bee fauna parallels that of the flora, with taxa having the following origins or distributions: West Indian, North or Central American, Andean, xeric or temperate South American, and Neotropical widespread. A brief account on the contribution of George Salt to the knowledge of the biology of this Colombian bee fauna is also provided.     

Is the occluded corolla of Antirrhinum bee-specialized?

This paper provides the first results of pollinator fauna associated to the reproductive biology of Antirrhinum charidemi, Antirrhinum graniticum and Antirrhinum braun-blanquetii. Censuses of over 16 hours spanning the flowering phenology of each species were drawn up. As previously hypothesized, the characteristic occluded (personate) corolla of Antirrhinum was exclusively pollinated by bees, but by a higher number (four) of bee families than predicted. Seven bee species (Bombus hortorum, Anthidium manicatum, Chalicodoma lefebvrei, Anthidium sticticum, Anthophora dispar, Xylocopa violacea and Anthidium cingulatum) account for over 90% of flower visits (2098). The flower visitation index varied between species (0.0126?0.0320), reaching relatively medium values compared to Mediterranean plants. In contrast, reproductive success was found to be high when estimated as both fruit (45.4–84.2%) and seed (75.4–98.4%) sets, suggesting that these bee guilds are effective pollinators of the three self-incompatible Antirrhinum species. Specialization of the personate flower of Antirrhinum is therefore interpreted for bees with both large bodies and high visitation indices.     

Diversity and life-history traits of wild bees (Insecta: Hymenoptera) in intensive agricultural landscapes in the Rolling Pampa,Argentina

The decline of bees is a major concern due to their vital role in pollinating many crops and wild plants. Some regions in South America, and especially the Pampas, are amongst those parts of the world where stressors of bee populations have been little studied. The Pampas has been intensively transformed for agriculture, being presently one of the most productive areas of agricultural commodities in the world. Here, we aim to provide first insights on the taxonomic and functional composition of bee assemblages in the Rolling Pampa, the most intensively managed part of the Pampas. Soybean (herbicide-tolerant genetically modified varieties) is the predominant crop in this region. Bees were sampled with coloured pan traps. Sampling points were located on field margins in either the cropped or the semi-natural grassland area of a farmland site devoted to annual cropping. A total of 2384 individuals were caught, representing 33 taxa [mainly (morpho)species]. The subgenus Lasioglossum (Dialictus) largely dominated captures (78% of the total abundance) and was relatively abundant in the entire study area, suggesting that some species are likely to reach their ecological requirements in cropped areas. No-till fields and field margins may provide large areas for these below-ground nesting species, while their polylectic food preferences allow them to collect pollen on a wide range of plant species, among them possibly soybean. On the contrary, the richness and the abundance of other taxa were higher in the semi-natural area than in the cropped area. Among them, above-ground nesting or oil-collecting species, which have more specialised nesting and floral requirements, were highly associated with the semi-natural area. Our findings highlight the large dominance of L. (Dialictus) species in this highly intensively managed landscape, and the urgent need of preserving semi-natural habitats to maintain species-rich and functionally diverse bee communities in the Pampas.     

Nesting dynamic and sex allocation of the oil-collecting bee Centris (Heterocentris) analis (Fabricius, 1804) (Apidae: Centridini)

The study of nesting biology is an important biological aspect for both conservation and management, but despite this, only a few of the over 20,000 described bee species have had their nesting biology studied. The aim of this study was to describe and analyse the nesting biology and dynamic of an aggregation of the cavity-nesting bee species Centris analis during one generation. Twenty-eight females were marked and had their nesting activities documented, 64 nests were sampled, and 113 individuals emerged from these nests. An asymmetry in the sex allocation was observed for this species, and the sex ratio was male-biased in the first nests built. Nonetheless, considering an individual-level perspective, the females showed different strategies, with some of them producing more daughters, other females producing more sons, and still others producing an equal proportion. The productivity per female was low compared to that of other solitary bee species, and the females displayed an aggressive usurpation behaviour against conspecific females and performed an intraspecific parasitism. As the substrate used to nest is a limiting factor for this species, the usurpation behaviour may be an important strategy that could increase the chance of nesting. A few males used the nesting site as a resting place and the fidelity observed to this locale was high. This study provides new information about the natural history of this bee species, which is an important pollinator of both crops and native plant species.     

A new genus of Atlantic octocorals (Octocorallia: Gorgoniidae): systematics of gorgoniids with asymmetric sclerites

Gorgonian corals from the families Gorgoniidae and Plexauridae are among the most abundant and diverse shallow‐water cnidarians found along tropical and temperate Atlantic coasts and reefs. Some morphological traits in the Gorgoniidae comprise a taxonomic challenge due to their morphological plasticity as well as a lack of discrete character states among most genera. Some gorgoniids have asymmetrical sclerites in the surface coenenchyme, a character trait that consists of a continuum among several genera and can cause confusion with members of the family Plexauridae. The collection of a unique gorgoniid in Tobago (Trinidad and Tobago), which contains this trait but also maintains affinities to many different genera, stimulated a wider morphological comparison of gorgoniids using scanning electron microscopy (SEM). Using the morphology of asymmetrical sclerites, gorgoniids could be grouped in four monophyletic clades that were compatible with recent molecular phylogenies and provide a provisional subfamilial classification: (1) a derived clade including gorgoniids with scaphoid sclerites, comprising the reef fauna from the western Atlantic (e.g. Gorgonia, Pseudopterogorgia [ = Antillogorgia], Pterogorgia, and Phyllogorgia); (2) a related group with taxa containing capstan sclerites including genera with short capstans and/or disk‐spindles sclerites such as Pacifogorgia and Leptogorgia ( = Lophogorgia); (3) a clade of gorgoniids with asymmetrical spiny sclerites including the Atlantic genus Muriceopsis and the Indopacific Pinnigorgia together with the distant single species from the new genus and species Tobagogorgia hardyi (it is worth noting that Muriceopsis was previously classified in the Plexauridae, but there is overwhelming molecular support that groups it with the Gorgoniidae and morphological evidence that suggests affinity with Pinnigorgia); and finally (4) a basal clade with long and spiny spindles, which includes some African species such as Filigorgia angolana and F. schoutedeni as well as other new combinations (all species previously classified as Leptogorgia but not phylogenetically related). The combined set of sclerites of Tobagogorgia (e.g. both asymmetrical and irregularly bent spindles) separates it from the diagnostic characters of related genera such as Muriceopsis and Pinnigorgia. The latter two genera are also profusely branched, including pinnate patterns, whereas T. hardyi has only one or two branches. Externally, T. hardyi looks like some Leptogorgia colonies that branch minimally but it is clear that the sclerites are very different and thus also their phylogenetic divergence.     

A comparison of the structure and function of the cerotegument in two species of Cryptostigmata (Acarina)

Cerotegument plastron structure and function are compared in two species of non-related Cryptostigmata from different families in two environments prone to flooding, namely soil and leaf litter, as represented by Euzetes globulus (Nicolet) (Euzetidae), and the intertidal zone by Ameronothrus marinus (Banks) (Ameronothridae). Differences in cerotegument structure reflect the degree of exposure to mechanical stresses and hydrostatic pressure encountered in these two environments. Cerotegument plastrons communicate with the tracheal system in heavily armoured species, e.g. E. globulus, but gas transfer may occur across the cuticle in those species having a lightly sclerotized leathery cuticle, e.g. A. marinus.