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New Paper Friday

Foraging areas of Leach's storm petrels

Patricia Jones

 Petrel burrow on Kent Island, photo by Fred Field

Petrel burrow on Kent Island, photo by Fred Field

New paper out with data from Kent Island! April Hedd from Environment and Climate Change Canada has published a paper this week in PLOS One. In 2013 and 2014 Dr. Hedd and her team put global location sensors (GLS) on petrels from 7 breeding sites in the Canadian Maritimes including Kent Island. In total, they obtained data from 133 petrels. From Kent Island, 17 petrels produced data (out of 20 tagged) in 2013 and 15 (out of 20 again) in 2014. On average, petrels made 4 day foraging trips which were 400 to over 800 km away from their burrows. Most of the petrels headed out to forage in deep oceanic shelves off the coast. The Kent Island petrels were different, foraging mostly over shallow water closer to shore. As you can see in the map below, they cover the entire Gulf of Maine and Georges Bank! Kent Island birds tended to have longer foraging trips than birds from other sites, but they covered smaller areas. Their foraging ranges overlapped by over 60% across years. Kent Island falls in the southernmost part of the range of storm petrels, and their different foraging behavior is likely to be influenced by their geographic location. It's wonderful to think about the birds we see on Kent Island having foraged off of Cape Cod a few days before!

 Figure 2 from Hedd et al. 2018.

Figure 2 from Hedd et al. 2018.

Swallow populations across the decades

Patricia Jones

 Swallow boxes on Kent Island. Photo by John Lichter.

Swallow boxes on Kent Island. Photo by John Lichter.

New paper lead by graduate student Tara Imlay from the University of Dalhousie in Nova Scotia (and including Professor Nat Wheelwright from Bowdoin and Professor Marty Leonard from Dalhousie) is out in Ecosphere. Imlay and her colleagues are searching for explanations for the dramatic declines that occurred in many swallow populations in the 1980's. They therefore examined breeding success and whether it was related to phenology or climate change for bank, barn, cliff, and tree swallows in datasets (including Kent Island!) that spanned 57 years. Surprisingly, given the declines in swallows, only bank swallows showed consistent declines in breeding success over the decades. In addition Bank swallows were the only species that did not advance their egg laying timing earlier across decades, all other species are laying eggs 8-10 days earlier than they were in the 1960's. This research indicates that for three of these four swallows, population declines do not seem to be driven by declines in breeding success. This highlights the need to examine factors that may be causing bird mortalities during migration or over the winter season. We only had one breeding pair of tree swallows on Kent Island last summer, but the boxes are up and will continued to be monitored. 

Mason Bees and Bee Hotels

Patricia Jones

 One of our native mason bee species,  Osmia lignaria , the blue orchard bee. Photo by  Brian Buckner

One of our native mason bee species, Osmia lignaria, the blue orchard bee. Photo by Brian Buckner

I have been asked a lot of questions the past couple of weeks about mason bees, and specifically mason bee hotels that you can put up in your yard. This coincides with an article about mason bees in Scientific American about the attempts of the almond industry to start commercially producing (and pollinating with) mason bees. The colony collapse disorder in honeybees has hit almond farmers particularly hard. This has led the industry to search for alternative pollinators, and the blue orchard bee, Osmia lignaria, emerged as a likely candidate. Mason bee is a general term usually referring to bees in the genus Osmia which use mud and clay in building their nests (thus the "mason" reference). Here in Maine we have 16 species of mason bee, only one of which, Osmia caerulescens, is non-native. They are often metallic green or blue like the blue orchard bee in the photo above, but are not to be confused with sweat bees which are also usually metallic but in a totally different family (Halictidae). Mason bees will sting if you squeeze them too hard, but the sting is very minor. Mason bees live in holes in wood the stems of reeds and are solitary, which is to say they do not have a single queen and lots of workers, rather each female lays her own eggs and provisions for them. They are however, fairly gregarious, with females laying their eggs in holes right next to other females. The photo below shows a cut-away bird's-eye-view of what the nests of individual females look like in a man-made mason bee home. A female will select a hole, and at the furthest point from the entrance she will put a ball of collected pollen, lay a single egg on that pollen ball, and then build a mud wall to close off that "cell". She will repeat this over and over until it fills up her hole like so:


Each cell therefore contains a mason bee larva feeding on the pollen ball its mother provided.

In Europe and Japan orchards are effectively using mason bee species for pollination of crops like plums, apples, and nuts. They are highly effective pollinators, with some reports of one mason bee female being able to pollinate as many flowers as 100 honeybees. This has to do with differences in their behavior. Mason bees visit more flowers than honeybees in each foraging visit, travel further between trees, and make more contact with the reproductive parts of the flowers than honeybees. The effort to commercial rear blue orchard bees for almond pollination is colossal, as detailed in the Scientific American article. But if you are interested in interacting with some mason bees in your own yard, it's pretty feasible. Mason bee "hotels" or "condos" have become popular recently. These are easy to make, and there are lots of places you can buy them. They do require some annual maintenance to keep pests under control. 

 Homemade mason bee house note that this kind is not great (see below)

Homemade mason bee house note that this kind is not great (see below)

 Mason bee house for sale from gardening shops

Mason bee house for sale from gardening shops

The best mason bee houses have removable/replaceable tubes (often made out of reeds or cardboard). This is because pests, parasites and molds can build up in the houses, so you need to replace the reeds on an annual basis. Mason bees overwinter as cocoons and emerge as adults in the spring, so you would want to wait for the bees to emerge in the spring and then replace the wooden blocks or tubes. If you want to be more involved in management of mason bees you can actually remove the cocoons from the tubes in the fall, sort and clean them from infected cocoons, and overwinter them in your fridge (see lots of details on how to do this here). You are likely to get a mixture of native and non-native mason bee species, as well as some leafcutter bee species (Megachile sp):

 Leafcutter bee. Photo by Kim Taylor

Leafcutter bee. Photo by Kim Taylor

Leafcutters cut circular pieces out of plant leaves to line their nests. They are also good pollinators, and we have 11 native Megachile species in Maine and two non-natives. They may, however, leave your roses with circular holes in the leaves, but the roses will survive. 

Bee viruses in bee-mimicking flies

Patricia Jones

 The hoverfly (or syrphid fly)  Eristalis interrupta  (or  nemorum ), photo by James Lindsey in Belgium.

The hoverfly (or syrphid fly) Eristalis interrupta (or nemorum), photo by James Lindsey in Belgium.

This week's paper is in Biology Letters, led by post-doc Emily Bailes from Royal Holloway University of London. Dr. Bailes examined the presence of honeybee viruses in hoverflies (also called syrphid flies as they are in the family Syrphidae). Syrphid flies first deserve some introduction of their own. The Syrphidae family is comprised of flies whose adult lifestages predominantly feed on flower nectar and pollen. What is particularly striking about them is their mimicry of other pollinators including a variety of bees and wasps. As a small example, the insects in the photos below are all flies!

 Syrphid flies. Photos and layout by Joaquim Alves Gaspar. 

Syrphid flies. Photos and layout by Joaquim Alves Gaspar. 

Although syrphid flies fill the same niche as bees (foraging from flowers), and look like bees, there has been little investigation of their role in the spread of pollinator diseases. It has been previously demonstrated that viruses originally found in managed honeybees can spread to wild native bees through visits to shared floral resources . The role of very ecologically similar, but evolutionarily distant, syrphid flies in disease dynamics had remained unexplored. Dr. Bailes and her colleagues collected 20 individuals each of honeybees and four species of syrphid fly at a long-term study site in England called Wytham Woods (famous for its research with great tits). Of the four syrphid flies, two species are honeybee mimics in the genus Eristalis that not only look like honeybees (see the first photo above) but also mimic honeybee behavior at flowers. The other two species were in the genera Episyrphus and Platycheirus and although they forage for nectar and pollen from flowers they do not look or act like honeybees. Dr. Bailes and her colleagues used RNA sequencing to determine the presence or absence of six common bee viruses in the honeybees and flies. They found bee viruses present in the two honeybee mimicking Eristalis, but not in Episyrphus and Platycheirus even though those flies are visiting flowers in the same habitat! The viruses that occured in Eristalis were in lower numbers than in honeybees, and had almost identical RNA sequences to the viruses from honeybees, indicating that the viruses may not be replicating in flies, rather the flies are picking them up from honeybees. The flies, however, may still be able to serve as vectors of the viruses even if they don't infect the flies. The extent to which these viruses are replicating in the flies (and negatively impacting the flies) is an area in need of further investigation. Also I really want to know more about why the viruses only occur in Eristalis. Are they more likely to have the viruses because they share more individual flowers with honeybees? Or is it because of something else about how they behave? I am fascinated by how behavior could be shaping disease transmission. 

More field borders lead to more bees & why we need more bee diversity

Patricia Jones

 A native bee montage by Sam Droege. 

A native bee montage by Sam Droege. 

Bee double-header this week! You may be overwhelmed by the bee landscape ecology papers, and honestly so am I. But this is an attempt to keep up with the literature, so here it goes. Two papers this week. The first one I will cover is in Proceedings of the Royal Society of London B, lead by Annika Hass a PhD student at the University of Göttingen in Germany, and examines how field edges and crop diversity affect bee communities and pollination in Western Europe. The second paper is in Science, lead by Rachael Winfree, a professor at Rutgers University, examines the number of bee species necessary for successful crop pollination in the field. 

In the Proc R Soc B paper, Hass and her colleagues examined the pollinator communities in 229 fields in 4 European countries. To examine the pollination services of those pollinators they planted rows of radish, Raphanus sativus, along the edges of the fields and measured seed set. It's a little counter-intuitive to think about radish pollination, as we mostly eat the roots of radishes rather than the fruits. But people do sometimes eat the radish seed pods, and sometimes use the seeds for oil (and they are definitely used for making more radishes). For each field they mapped the field-border density (they call this configurational heterogeneity), as well as the diversity of crops in the field. Hass and her colleagues found that field-border densities were positively correlated with wild bee abundance, and wild bee abundance was positively correlated with radish seed production on the field edges. Interestingly, crop diversity was actually negatively correlated with wild bee abundance. The authors propose that this is because increased crop diversity in their European fields meant more highly managed crops (with associated pesticide applications). It would be interesting to see if these results are the same when you increase diversity without highly managed crops. It's a little hard for me to envision what higher field border densities (configurational heterogeneity) would look like, and I wish that they had included aerial photographs of fields with high and low field border densities. But generally it seems that interrupting monocultures with natural field borders enhances numbers of bees, and that makes intuitive sense. 

In the Science paper, Winfree and her colleagues measured pollination of watermelon, cranberry, and blueberry, by wild bees on 16 farms in New Jersey and Pennsylvania. They collected bees visiting flowers in each of these crops. In addition they bagged fresh flowers until they opened, and then removed the bags. They recorded the first bee species to visit the flower and then recorded the number of pollen grains deposited by that bee, giving them data on the pollination services of different bee species (I think this is very cool!). They found that in order for the crops to receive target levels of 50% pollination at one site alone might take less than 20 bee species, but to achieve that level of pollination over 15 different sites (spread over thousands of kilometers and thereby varying in many different factors) would require to ecosystem to maintain more like 50 different bee species, and 75% pollination would require 80 different species. When people think of crop pollination they mostly think of honeybees, and this study is so important because it highlights that to effectively pollinate our crops we cannot rely on honeybees but rather need a large diversity of bees.

One that note, the honeybee is a managed livestock species introduced from Europe and is not a good solution for loss of our native bees, and in fact is part of the problem (see recent NPR story on this). I love honey as much as I suspect you do, and we currently are dependent on managed honeybees for many of our crops (because of the way we plant them in huge monocultures - but that's for another time, and see the Proc R Soc B paper above). But we need to be clear when we are talking bee conservation we are not talking about honeybees, honeybees have management problems, but that is a livestock management challenge, not a conservation issue. I have made the analogy that focusing on honeybees when we talk about bee conservation is as if a bird conservation discussion was focused on chickens. There is a lot of research needed on honeybee issues (plus they have very cool biology), and they are an important part of our food chain and economy, but these are not conservation issues. 

Floral Shape and bee communities

Patricia Jones

 A honeybee on an open-shaped flower. Photo by George Gray. 

A honeybee on an open-shaped flower. Photo by George Gray. 

 A bumblebee on a tube-shaped flower, photo by Brendan Zwelling.

A bumblebee on a tube-shaped flower, photo by Brendan Zwelling.

This week's paper is published in Biology Letters, single-authored by Katherine Urban-Mead, a graduate student in the Department of Entomology at Cornell University. Urban-Mead used network analysis to examine how the shapes of flowers impacted bee species abundance in floral communities. In particular, Urban-Mead compared the availability of flowers with open corollas where the nectar is easily accessible to bees (especially short-tongued bees, but more on that later) with flowers with tube-shaped corollas where the nectar is at the base of the corolla requiring bees to enter the flower (and may therefore be easier for long-tongued bees). 

Urban-Mead worked in six meadows in Connecticut containing diverse flowering plants. In each meadow she selected three 3 x 6 m plots that had similar floral communities. In one plot she removed all the open flowers from one half of the plot, in the second plot she removed all the tube-shaped flowers from one half of the plot, and the third plot was left undisturbed. Urban-Mead then watched these floral plots and recorded the visits made be visiting bees of different morphospecies (bees are quite hard to identify to species in flight - so they are often lumped into identifiable groups). In total she saw more than 500 bees visit more than 3500 flowers. 

The bee community that visited open flowers in all plots was more diverse than the bee communities that visited tube-flowers. In the plots with the open flowers removed, the bees on the remaining tube flowers were not different from the bees on the tube flowers in the undisturbed plots. In the plots with the tube flowers removed, however, the bees on the open flowers were different from the bees on open flowers in undisturbed plots. Part of this shift appears to be due to tongue-length. Bees with longer tongues can more easily access nectar in tube-shaped flowers, and correspondingly, there is a higher proportion of long tongued bees in the control plots and plots with open flowers removed than in plots with tube-shaped flowers removed. It appears, therefore, that the availability of tube-shaped flowers is a more important predictor of bee communities than open flower shapes, but open flowers host higher bee diversity in general. 

Task complexity and social learning in bumblebees

Patricia Jones

 A buff-tailed bumblebee,  Bombus terrestris . Photo by  Kathleen Purser.  

A buff-tailed bumblebee, Bombus terrestris. Photo by Kathleen Purser. 

Baracci et al. 2017.jpeg

This week's paper is in Behavioral Ecology, and is lead-authored by David Baracchi, a post-doctoral researcher at the University of Toulouse in France. Baracchi and his colleagues have examined how the difficulty of a foraging discrimination task (being able to tell which flowers are rewarded) affects the use of social information in bumblebees. In bumblebees, one source of social information is the presence of another bee on a flower. The presence of that bee could indicate to an observer that a flower has available resources. Social information comes with costs and benefits (see my previous post about social learning here). For example, the presence of that bee on the flower also could indicate that the observer bee would have to compete with that bee for those floral resources. Researchers have therefore proposed that individuals should only use social information selectively, and should rely on information they have acquired through their own experience the rest of the time. This has lead to the development of "social learning strategies" which are predictions about when animals might be expected to use social information. For example, you might expect an individual to be more likely to use social information when they are inexperienced, or uncertain about the environment. Baracchi and his colleagues have examined how difficulty in discriminating rewarding from unrewarding flowers affects social information use. 

The difficulty in the floral discrimination task was determined by the difference in the number of bars displayed by rewarded and unrewarded flowers (see figure above from Baracchi's paper), such that one bar versus four bars is an easy discrimination task, whereas five bars versus four bars is a hard discrimination task. Bees were first trained to two different stimuli. They were trained to associate the presence of four bars with sugar solution, and to associate the presence of another bee with sugar solution. Then bees were given a range of tests and Baracchi et al. examined the percentage of errors (landings on unrewarded flowers) made by bees. Bees received the flowers alone, the flowers with social information consistent with the floral information (the bee was present on the four bar flower they had been trained was rewarding), or a conflicting information scenario in which the bee (which they had been taught was rewarding) was now present on the other, non-four bar, flower (either one or five, which they had been taught was not rewarding). 

Baracchi et al. show that when the task is easy, bees perform similarly regardless of whether the social information is available, consistent, or in conflict. In contrast, when the task is hard, errors are substantially reduced when the social information is consistent. These results highlight that while social information may not be important to bees when faced with easy discrimination tasks, in the cases of hard discrimination tasks, bees may be able to use additional sources of information to increase their foraging success. 

Fungicides, pathogen infections, and bumblebee declines

Patricia Jones

 The western bumblebee,  Bombus occidentalis , whose populations are currently in decline. Photo by Diane Wilson.

The western bumblebee, Bombus occidentalis, whose populations are currently in decline. Photo by Diane Wilson.

This week's paper is lead-authored by Professor Scott McArt at Cornell University, and is published in Proceedings of the Royal Society B. McArt and his colleagues have examined the factors associated with declines in populations of four declining bumblebee species in the US: Bombus occidentalis, Bombus affinisBombus pensylvanicus and Bombus terricola. McArt and his colleagues collected over 10,000 bumblebees from 284 sites in the USA. Bees belonged to these species known to be in decline as well as bumblebee species whose populations are currently stable (Bombus bimaculatusBombus impatiens, Bombus vosnesenskii). At each of the 284 sites they quantified 24 landscape variables including: latitude and longitude, habitat types, total human population densities, and pesticide usage including insecticides, herbicides, and fungicides. They also screened the collected bees for two parasites: Crithidia bombi and Nosema bombi. These two parasites are suspected to be an important component of bumblebee population declines. McArt and his colleagues then used multi-variate model selection to assess the impact of individual habitat and pesticide variables on infection prevalence and population declines in bumblebees. They found that the best predictor of both N. bombi infection and bumblebee population declines was use of the fungicide Chlorothalonil. Chlorothalonil is frequently used to combat fungi, molds and mildew, on trees, turf, and fruits. The maps below (from McArt et al. 2017) show the prevalence of Chlorothalonil usage overlaid with parasite infection and population declines in bumblebees. 

McArt et al. figure.jpg

Chlorothalonil has already been shown to be associated with honeybee parasite infection, and has negative impacts on colony growth in the common eastern bumblebee, Bombus impatiens. The mechanism by which chlorothalonil affects bees is uknown, but one possibility is that it disrupts the bee's normal gut microbiota, making them more susceptible to parasite infection. Other interesting findings from this study include a negative relationship between parasite infection and urbanization. McArt and his colleagues suggest that the diversity of flower resources in urban gardens may actually enhance bumblebee nutrition, buffering them against pathogen infection. 

Kent Island Special! Effects of Climate Change on Reproduction in Storm-Petrels

Patricia Jones

 Bowdoin student and Kent Island fellow, Claire Goffinet ('19) with a Leach's storm-petrel chick summer of 2017. Photo by Fred Field for Bowdoin College

Bowdoin student and Kent Island fellow, Claire Goffinet ('19) with a Leach's storm-petrel chick summer of 2017. Photo by Fred Field for Bowdoin College

This week's paper is in Global Change Biology and is co-authored by three former Kent Island Directors. It is lead-authored by Bob Mauck of Kenyon College, with Don Dearborn of Bates College, and (posthumously) Chuck Huntington of Bowdoin. The three directors used long-term data collected on Kent Island to study the effects of global temperature on reproductive success in Leach's storm petrels. Data on temperature has been collected on Kent Island since 1939, and the storm-petrel dataset was initiated by Chuck Huntington in 1955. In this week's paper Mauck et al. examined the effects of local (Kent Island) temperature, global mean temperature (a combination of air and sea temperatures), and sea surface temperature on reproduction of storm-petrels using 56 years (1955-2010) of the Kent Island long-term databases on weather and petrel reproduction. The best fitting model for predicting storm-petrel reproductive success included both global mean temperature and local temperature on Kent Island, but global mean temperature is the most important predictor of reproductive success in storm-petrels.

Mauck et al. found that as temperatures rose between 1955 and 1988 storm-petrel hatching success increased (likely due to decreased incubation costs), until in 1988 it appears that global temperatures reached a critical point, and since 1988 hatching success as declined as global temperatures have continued to rise. The decline in hatching success since 1988 is most likely due to the negative impacts of increasing temperatures on food availabilities for petrels as fish move further northward and into deeper waters. 

The effects of increasing temperature, however, were not the same on all birds. Birds with more breeding experience have higher hatching success in general, and appeared to be somewhat buffered against these effects of climate change. The effects of increasing temperatures on hatching success were more strongly observed in less experienced petrels (both positively before 1988 and negatively afterword). Mauck et al. propose that this is because less experienced petrels are less effective foragers, and therefore have benefitted more from reduced incubation costs with increased temperatures, but suffered more from reduced food availabilities. 

 Two authors and Kent Island Directors in 2007, Bob Mauck (L) and Chuck Huntington. Photo by another Kent Island Director, Ed Minot.

Two authors and Kent Island Directors in 2007, Bob Mauck (L) and Chuck Huntington. Photo by another Kent Island Director, Ed Minot.

Kleptopredation in nudibranchs

Patricia Jones

 The nudibranch (a kind of mollusk),  Cratena peregrina,  on it's prey, a hydroid (which are related to jellyfish - yes! I know it looks more like a plant). Photo by Jean-Marc Kuffer.

The nudibranch (a kind of mollusk), Cratena peregrina, on it's prey, a hydroid (which are related to jellyfish - yes! I know it looks more like a plant). Photo by Jean-Marc Kuffer.

This week's paper is in Biology Letters, lead authored by Dr. Trevor Willis who is a Senior Lecturer at the University of Portsmouth in the UK. Willis and his co-authors studied the relationship between nudibranchs and their hydroid prey off the coast of Sicily. Nudibranchs are spectacular little animals. Often referred to as "sea slugs" (which does them no justice) they comprise about 2,300 species of shell-less mollusk distributed worldwide. They vary enormously in color, including bright polka-dots and stripes, as well as in shape. The plumes on their backs are called cerata and are where much of their gas exchange occurs (gills) and also where they defend themselves through nematocysts, stinging cells that they acquire from their prey which includes anemones and the hydroids (more on those soon). Many nudibranch species are also toxic, either via the consumption of toxic sponges or de novo synthesis of toxins. These defenses likely explain the extraordinary warning colorations of many nudibranchs. 

Many nudibranch species, including Cratena peregrina, the subject of this paper, consume hydrozoans. Hydrozoans are small animals related to jellyfish. They have different life stages, one of which is the polyp stage in which they are called hydroids. Hydroids are most commonly colonial, which is to say they live in groups. Hydroids are a bit like upside-down jellyfish that live in little tubes all connected to each other in a branching pattern (which makes them look a bit like a plant). 

 The hydroid  Eudendrium racemosum . Photo by Fernando Herranz.

The hydroid Eudendrium racemosum. Photo by Fernando Herranz.

Hydroids use the stinging nematocysts on their tentacles to capture zooplankton prey. Hydroids themselves are then prey to nudibranchs. In this week's paper Willis and his colleagues examined this relationship between nudibranchs, hydroids, and the hydroids' feeding on zooplankton. In particular, they tested the hypothesis that nudibranchs prefer to feed on hydroids that have just captured a zooplankton, than on hungry hydroids. To do this they brought nudibranchs and into flowing seawater tanks in the lab and provided them with the choice between feeding on hydroids that were starved versus hydroids that had just been fed zooplankton. They measured nudibranch preferences for fed and starved hydroid polyps as well as how long it took the nudibranchs to consume fed and starved polyps. They also examined the stable isotope ratio of nudibranchs, hydroids, and zooplankton (see my post on "Smashing & Spearing Stomatopods" for more explanation of stable isotope analysis) to assess what nudibranchs are eating in the wild. 

Willis and his colleagues found that nudibranchs preferred to attack hydroid polyps that had just consumed zooplankton over starved hydroids, and it took nudibranchs longer to consume those fed hydroids than starved hydroids. In addition, Willis showed with stable isotope analysis that nudibranchs were not solely consuming hydroids, but that zooplankton made up at least half of their diet. This data suggests that not only are nudibranchs preferentially consuming hydroids that have just captured prey in the lab, they also do so in the field. Willis and his colleagues introduce a novel term for this behavior: "kleptopredation". Kleptopredation is a combination of kleptoparasitism (when animals steal food from other individuals) and predation, as the nudibranch in this case is doing both. It is not clear why nudibranchs do this. The most obvious hypothesis is that polyps with captured zooplankton deliver more calories (and potentially nutritional diversity?) than starved polyps. Willis also suggests that the consumption of fed polyps results in nudibranchs consuming less polyps total, and may prevent them from driving their hydrozoan prey locally extinct. 

The mutualism between bats and pitcher plants

Patricia Jones

 The Bornean bat  Kerivoula hardwickii  approaching the pitcher plant  Nepenthes hemsleyana. 

The Bornean bat Kerivoula hardwickii approaching the pitcher plant Nepenthes hemsleyana. 

 Bat inside a pitcher plant roost. Photos by Merlin Tuttle.

Bat inside a pitcher plant roost. Photos by Merlin Tuttle.

This week's paper is in Scientific Reports, lead authored by Michael Schöner at the University of Greifswald in Germany. The research focuses on an extraordinary relationship between the woolly bat, Kerivoula hardwickii, and a pitcher plant, Nepenthes hemsleyana. Pitcher plants are believed to have evolved in areas with low soil nitrogen content. To achieve their necessary nutrients these plants have turned to carnivory. The pitcher part is a liquid trap, usually filled with digestive juices. The pitchers emit odors that attract insects who become trapped in the pitcher where they are digested, providing nutrients for the plant. Nepenthes hemsleyana, however, is different. Rather than emitting attractive odors or containing digestive juices, instead the pitcher of N. hemsleyana is a perfect roost for bats. Woolly bats sleep in the pitcher plant during the day, and poop there, providing nitrogen for the plant. These pitchers have an unusual shape that makes them not only ideal for a sleeping bat, but also easily detected by echolocation. Additionally, the inner wall of the pitcher has a waxy texture that deters egg-laying insects, providing a pest-free home for the bats. This is considered a mutualism, because the bat provides poop to the plant, and the plant provides a roost for the bat. 

The pitcher plant is highly dependent on the bat, as this pitcher is not particularly good at catching insects. Woolly bats, however, will roost other places than these pitchers. They will roost in the old pitchers of other pitcher plant species, and in furled leaves. So how can the pitcher plant depend on a bat that does not depend on it? In other words, what stabilizes this mutualism?

The authors speculate that the mutualism is stabilized by the quality of the pitcher plants as roosts for the bats. The pitchers of this species make such superior roosts that bats will continue to use them even if they have the option to go elsewhere. To test this they had to examine whether bats preferentially used pitchers of this species (Nepenthes hemsleyana) in the field. The authors first radiotracked bats in the field to examine which of the available roosts the bats were using. At one site pitchers of N. hemsleyana made up only 5% of the available roosts but were all occupied by bats. Interestingly, they did not see any bats switching between different types of roosts in the field (if a bat was roosting in furled leaves they continued to roost in furled leaves and did not necessarily switch to pitcher roosts). To confirm this roost fidelity they brought the bats into flight arenas and gave them a range of options. Schöner and his colleagues found that bats which had been found in pitchers in the wild always selected to roost in pitchers. But 21% bats that had been found in furled leaves in the field switched to pitchers in the lab. This asymmetry in roost selection indicates that bats preferentially roost in pitcher plants over furled leaves. To confirm that bats roosting in pitchers versus in leaves in the field were not genetically different (excluding the possibility that they were different subspecies of bats roosting in different places), Schöner and his colleagues conducted population genetics on the bats collected from different roosts and confirmed that there were not genetic differences by roost. It appears therefore that bats learn, either socially or individually, a certain roost type and then stick with that roost type over their lives.

The mutualism between woolly bats and pitcher plants is likely stabilized by the high quality of the pitcher plants as roosts, and a general preference of bats to use them when they are available. Interestingly, however, it appears that there are individual bats that will use furled leaves as roosts for their entire lifetimes. This willingness to use other roosts could buffer the populations of woolly bats from environmental perturbations that reduce the availability of pitcher plant roosts. 

What does learning look like in a bumblebee brain?

Patricia Jones

 Figure 1 from  Li Li et al.  a) the arena set-up with bees exposed to 10 flower colors; b) the different experimental treatments; c) the frontal-view of a whole bumblebee brain; d-f) enlarged views of the portions of bumblebee brains involved in vision; g) enlarged so you can see the labeled microglomeruli; h) microglomeruli structure.

Figure 1 from Li Li et al. a) the arena set-up with bees exposed to 10 flower colors; b) the different experimental treatments; c) the frontal-view of a whole bumblebee brain; d-f) enlarged views of the portions of bumblebee brains involved in vision; g) enlarged so you can see the labeled microglomeruli; h) microglomeruli structure.

This week's paper is in the Proceedings of the Royal Society of London B, lead-authored by Li Li, a PhD student in the Chittka group at Queen Mary University London. Neurons connect to (and thus communicate) with other neurons at regions called synapses. Microglomeruli are places were lots of synapses come together (synaptic complexes). This study looked at the link between bee learning of flower colors, and the density of microglomeruli in the bee's brains.

Li Li and her colleagues tested bumblebees in five different treatments. All bees were first pre-trained to forage from colorless (clear) artificial flowers. For some of the bees that was it, they were dissected to establish a baseline of microglomeruli density with no exposure to colors (Treatment 1). The next group of bees were trained to distinguish between 10 different colors, five of which had sugar solution and five had nasty tasting quinine (like what's in your tonic water; Treatment 2). The third group of bees were trained to distinguish between only two colors to examine the effect of the number of colors bees had to learn on microglomeruli density (Treatment 3). Group four continued to forage on clear flowers for as long as the other bees were learning flower colors to control for the effect of amount of foraging time on microglomeruli density (Treatment 4). In the final treatment the bees continued to forage on clear flowers, but they were surrounded by the colored flowers (with no rewards) to control for the effect of exposure to lots of flower colors (Treatment 5). In treatments 2-5 after the bees were trained they were then given memory tests to determine how well they performed at learning the task. You can see all these treatments detailed in Fig 1b above. 

In the ten color discrimination task (Treatment 2), Li Li and her colleagues found that learning rate and memory performance correlated with microglomeruli density in the collar part of the bee brain (Fig 1c above). This result, however, does not distinguish between correlation and causation. That is to say, it could be that better learners brains' increase more in microglomeruli density, or that bees with better learning abilities are the ones that already have higher microglomeruli densities. In comparing bees that had color learning versus those with no color learning (Treatments 2 and 1), they found that bees in the color learning treatment had higher microglomeruli densities and larger brain calyxes (the brain part in Fig 1d) in total. There are possibilities other than color learning directly to explain this pattern, however, and using the other treatments Li Li and her colleagues were able to show that the brain size is simply a product of the foraging activity the bees did (comparing treatments 4 and 2), and that the microglomeruli density is a product of exposure to colors not learning necessarily (comparing treatments 5 and 2). So in summary, there is a correlation between density of these microglomeruli and bee learning and memory. But causation is tricky, learning colors does seem to increase microglomeruli density, but this effect may largely be due simply to exposure to the colors. 

Larger soldiers....have a better sense of smell? A lesson from stingless bees

Patricia Jones

 Neotropical stingless bees,  Tetragonisca angustula , at the entrance to their nest. Photo by  Alex Wild .

Neotropical stingless bees, Tetragonisca angustula, at the entrance to their nest. Photo by Alex Wild.

This week's paper is in Biology Letters, lead-authored by Christoph Grüter at the Universidade de São Paulo in Brazil. The research is with a stingless bee species, Tetragonisca angustula. "Stingless bee" is actually a technical term that refers to bee species in the tribe Meloponini, which is comprised of >500 species distributed throughout the tropics worldwide. Like other social bees they store floral nectar, and stingless bee honey is actually collected by some human cultures (often referred to as melipona or meliponine honey). In many Hymenoptera (the bees, wasps, ants, termites etc.) colonies need to defend themselves from raids on their food or larvae. Raiders might be individuals of the same species from a different colony, or individuals of other species. In ants and termites, colonies often have some individuals that develop as soldiers, who work in nest defense and often are large, with weapons of some kind such as biting mouthparts. In many of the bees and wasps, there is not this need for soldiers because all of the workers can sting. But in the stingless bees there are some individuals that are larger, but they cannot sting and don't have any biting how do these larger workers defend the colony?

Grüter and his colleagues hypothesized that larger workers would be better at identifying intruders. In the Hymenoptera individuals recognize each other by scent, particularly by the cuticular hydrocarbon profile on their bodies, which they detect through pore plates on their antennae. Larger individuals have larger antennae, therefore more pore plates and a better sense of smell! Grüter and his colleagues hypothesized that the larger bees would be better at distinguishing between nest-mates and non-nest-mates. 

Grüter et al. had 10 colonies of T. augustula in the lab on campus at São Paolo. Into each colony they introduced bees that were either nest-mates, non-nestmates, or from a different stingless bee species, and recorded the responses of the guarding soldier bees. They also measured the size of the guarding bees. They found that larger bees were more likely to accurately identify and attack non-nestmates, and that the larger bees also had larger antennal surface areas and more pore plates. This confirms Grüter et al.'s hypothesis that larger bees have better senses of smell and are better able to identify intruders. I would think that the larger bees are also better able to cope with the intruders once they identify them, but that has yet to be tested. 


When to migrate south? When there is a good tail wind.

Patricia Jones

 A common noctule,  Nyctalus noctula . Photo my own.

A common noctule, Nyctalus noctula. Photo my own.

This week's paper addresses an age old question, how do animals decide when to migrate? The paper is entitled "Determinants of spring migration departure decision in a bat" lead-authored by Dina Dechmann at the Max Planck Institute. Pregnant female noctules in the spring migrate hundreds of kilometers Northeast from hibernation caves to insect-rich feeding grounds (this might be from Switzerland to Sweden or Bulgaria to Russia). But how do they decide when to leave? 

Over three years Dechmann and her colleagues fitted 29 female noctules with radiotransmitters and then used a small Cessna plane to search for the signals of bats that were migrating. They also obtained detailed weather information for the sites where the bats were tagged. In songbirds, birds tend to decide to migrate once they have put on enough fat. That was not the case for noctule bats, their decision to migrate was not affected by their body condition. Migration decisions in noctules were best explained by wind direction, wind speed, and air pressure. Bats were most likely to migrate on nights with faster tailwinds in the migration direction, and with higher air pressure. The tailwinds make a lot of intuitive sense, and generally high air pressure is associated with more stable weather conditions which might also be to a migrating bat's benefit. Songbirds generally migrate much further than bats, so the lack of putting on fat determining migration is likely due to the shorter migration distance. 

How poison dart frogs don't poison themselves

Patricia Jones

 The little devil frog,  Oophaga sylvatica , is one of the poison dart frog species in this study. Photo by Lucas Bustamante.

The little devil frog, Oophaga sylvatica, is one of the poison dart frog species in this study. Photo by Lucas Bustamante.

Poison dart frogs (frogs of the family Dendrobatidae, which as far as I can tell comes from "climbs trees" in Greek), are emblematic of the tropics. They are incredibly brightly colored, from blues to reds, to greens and yellows, in stripes as well as spots. But they are as well known for their poisons as for their colors. Their skins contain toxins such as epibatidine, a general toxin that can kill in doses as small as a microgram. They acquire these toxins from consuming poisonous insects (such as centipedes). Extracts from some species of poison dart frogs were used by indigenous Americans to poison the tips of their blow darts. But how does an animal as poisonous as a poison dart frog not poison itself?

This weeks paper, entitled "Interacting amino acid replacements allow poison frogs to evolve epibatidine resistance" is lead-authored by postdoctoral researcher Rebecca Tarvin (UT Austin) and is in Science. Animals that acquire defensive toxins through their diet are generally thought to have three alternative paths available to them to not poison themselves. 1) They can compartmentalize the toxin in their tissues such that toxin is kept separate from the tissues that would be sensitive to it. 2)  They can metabolize the toxin to reduce its toxicity to themselves. 3) They can be insensitive to the toxin's effect. This third option sounds like the safest. There is a hitch, however, because the best toxins are ones with very general effects on many different potential predators. For a toxin to be effective it needs to bind to a receptor. So what types of receptors do many different, distantly related (birds AND rodents AND snakes for instance) all have? Ones that are crucial for biological function. Such is the case with the poison frog, as epibatidine binds to the receptors for acetylcholine, a widespread nerve signaling molecule (neurotransmitter). But how then does the poison frog cope? It needs to be insensitive to epibatidine but still be sensitive to acetylcholine. 

Tarvin and her colleagues found that in the two groups of poison dart frogs that both make epibatidine, they have a single amino acid substitution that changes an amino acid in their acetylcholine receptor, making them insensitive to epibatidine. Both groups separately evolved this same nucleotide change (millions of years apart!). This change alone would make the two groups also less sensitive to acetylcholine, but each group separately evolved a different secondary amino acid substitution that restores their sensitivity to acetylcholine. This highlights the myriad paths that evolution can travel, parts of the solution are reached by the same paths in different lineages, and other parts of solution are achieved through different paths. 

As molecular, biochemical, and neurophysiology techniques improve we are starting to find insights into how a trait as complicated as resistance to a general toxin can evolve, and this paper is a wonderful example of how exciting those discoveries can be. 


Vocal flexibility in geckos

Patricia Jones

 Tokay gecko ( Gekko gecko ) in Thailand. Photo by Tontan Travel. 

Tokay gecko (Gekko gecko) in Thailand. Photo by Tontan Travel. 

This week's paper is from Proceedings of the Royal Society B, authored by Henrik Brumm and Sue Anne Zollinger at the Max Planck Institute for Ornithology. Brumm and Zollinger are the first to show flexibility in vocalizations by a reptile, and how the geckos do it is pretty cool. When we humans are trying to communicate in a noisy space, we usually yell at each other (i.e. increase our amplitude - technically called the Lombard effect). Tokay geckos, Gekko gecko, are native to Southeast Asia. They have a distinctive call, and in fact both the names "gecko" and "tokay" are onomatopoeia for the way the call sounds. Geckos in noisy environments have a different strategy for being heard. 

What they did

The methods for this study are very simple. They had male geckos in captivity in rooms lined with acoustic absorbing material and a microphone. For 24 hours white noise was broadcast to the geckos in the room, and their calls were recorded, and then for 24 hours there was silence and their calls were recorded. They repeated this to record four total days of calls. They then compared call duration, structure, and amplitude during noise and during silence. 

What they found

Geckos did not increase the amplitude of their calls, but they did increase the duration of the call syllables. This should also make them easier to hear. Additionally, Geckos have two types of calls, a softer cackle and the loud "Gecko!" Geckos in noisy environments make more of the louder gecko and less of the quieter cackle. They therefore are selecting to use more of the call types that are easier to hear when they are in noisy environments. 

The takeaway

Vocal flexibility in the form of the Lombard effect has been demonstrated in birds, mammals and even frogs. This is the first demonstration of vocal flexibility in reptiles, but also importantly, it is a different type of flexibility. Increasing the duration of syllables and perhaps selecting different words are tools that we use to communicate in noisy environments, but have not been reported as much in other animals. I am really interested in the use of these two different strategies, both duration increase and use of different calls. The white noise that the author's used is really broadband noise. I wonder if geckos in different types of noisy environments (next to a rushing stream, or when lots of crickets or cicadas are calling around them) use these two strategies differently in different noise contexts. 

Giving better talks

Patricia Jones

This week's paper is a departure from the normal format. It is by Katie Langin and is in the Ornithological journal Condor. Langin's paper is entitled: Tell me a story! A plea for more compelling conference presentations. In it she gives advice on how we could all improve our public speaking skills. 

And, but, therefore

The first recommendation that Langin makes, made by others before her as well, is to tell a story. It is important to get the audience hooked and intrigued in the question you are asking before you lose them in data. The recommendation for how to do that is in "and, but, therefore". This format lays out two things that are known about a study system: "frog-eating bats hunt their prey by listening for frog calls and have amazing learning abilities". Then highlight something that is unknown: "but almost nothing is known about their natural history". Finally state what you have done to address this: "therefore we radio-tracked wild bats as they moved from roosting and foraging sites in the wild". (This is not a fabulous example, but one that has been on my mind). 


The next piece of advice is don't try to tell your audience too many things! Pick out a few results (3 max?) that are the most central to your story to tell them about, they can't absorb more than that anyway. 

Memorable conclusion

Langin then advises to wrap up your story with at least a couple of minutes of conclusion. Even if the takeaways and future steps seem obvious, lay them out clearly so your audience doesn't miss them. 

Think of the broad audience

Talk not just to the expert in the room that you want to impress, but try to reach as many people as possible. This will help to spread the word about your research, and the expert will not be disappointed by a simplified story. 

Less text more visuals

Basically try to convert as much text as possible to visuals. Show your predicted outcomes with figures instead of text. Diagrams of methods instead of descriptions, graphs instead of written out results etc. 

Check out her appendix

Langin's paper has a long appendix with lots of helpful hints. Check it out! And see you at ESA!

Rational decisions in frog-eating bats

Patricia Jones

 The fringe-lipped (also called frog-eating) bat,  Trachops cirrhosus . Photo by  Christian Ziegler . 

The fringe-lipped (also called frog-eating) bat, Trachops cirrhosus. Photo by Christian Ziegler

This week's paper is lead-authored by Claire Hemingway who (like I was, in full disclosure) is a graduate student with Mike Ryan at UT Austin and Rachel Page at the Smithsonian Tropical Research Institute, both of whom are also authors on the paper. They address the concept of "rationality" in animal decision-making. Rationality in this case is used in the economic sense. In examining human decision-making the field of economics has established principles of rationality. One of the principles of rationality is independence from irrelevant alternatives. This principle is that preference for two alternatives should not change with the addition of other options. If you prefer A to B, the addition of C should not change your relative preferences for A and B. However, humans are highly irrational. This is demonstrated by the "decoy effect" that is frequently used in marketing. Hemingway and her colleagues give a nice example: if there are two cans of vegetables on a shelf one of which is expensive and organic (choice A) and the other of which is inexpensive and conventional (choice B) you may not have a strong preference between them. But if a third can is added that is expensive and conventional (choice C) this may cause you to now prefer the expensive organic over the inexpensive conventional. This irrationality has been demonstrated in many other animals, including hummingbirds, honeybees, and even slime moulds (such a cool paper!). Here it is tested in frog-eating bats. 

What they did

Frog-eating bats hunt frogs by listening for their calls (referred to as "eavesdropping"). In particular, a substantial literature has developed on bat response to calls of the túngara frog. Túngara frogs make simple calls (a "whine" alone) and complex calls (a whine plus "chuck" sounds added to it). Bats prefer complex calls, but they also prefer louder calls. The two options that Hemingway and her colleagues used, therefore, were a complex call (choice A) and a louder simple call (choice B). In the decoy effect test they added in a third stimulus, a simple call but even quieter than the complex call (choice C). This "decoy" should (if bats are irrational) increase the preference for A over B, because C is inferior to A in two different ways, versus inferior to B in only one way. 

What they found

When you pool all the bats together they had no preference for A over B with or without C. BUT that is not to say that all the bats were going to A 50% of the time and B 50% of the time. This was the case for many of the bats, but in the A/B choice test 2 of the 11 bats strongly preferred A, and one bat strongly preferred B. When C was added in, 3 bats preferred A and one bat preferred B. Only for one bat did preference change with the addition of C. That bat, as predicted, had no preference when there was just A and B and preferred A when C was added. It was the only irrational bat of 11. The other ten had no change in preference for A vs. B when C was added. This individual variation is so typical of animal behavior studies! 

The takeaway

Although many other studies have shown irrationality in animal decision-making, this study did not. There are a number of possible reasons why this could be. The authors propose that the high metabolic demands of powered flight in bats may select for more rational decision-making in foraging choices than in other species. BUT, irrationality has also been shown in hummingbirds! The discordance of these results with other studies could also be an artifact of the system, or stimuli chosen. Regardless, the question is very cool and I am glad to see it tested in this system!


Nicotine and floral constancy in bumblebees

Patricia Jones

 A buff-tailed bumblebee,  Bombus terrestris , visiting heather. Photo by  Steve Slater . 

A buff-tailed bumblebee, Bombus terrestris, visiting heather. Photo by Steve Slater

This week's paper is by David Baracchi from his postdoc in Lars Chittka's lab at Queen Mary University. Baracchi tackled the very cool combination of nectar chemistry and bee learning. Research with caffeine has indicated that the presence of caffeine in flower nectar enhances pollinator memory for that flower type, thereby increasing floral constancy (consistency in visits to one floral type over others) and thus pollination. In this paper they examined nicotine, which is present in the flowers of plants such as wild tobacco, Nicotiana attenuata. Baracchi studied the effect of nicotine in flower nectar on bee learning of flower colors, and fidelity to that color even after it was no longer rewarding. 

What they did

Baracchi and his colleagues did three experiments. In the first they assessed how the presence of nicotine at different concentrations affected foraging choices of bees. Individual bees were allowed to forage in an array of artificial flowers, half of which were blue and half violet. They tested 20 bees each with three nicotine concentrations, two natural concentrations (low natural and intermediate natural) and one unnaturally high concentration.

In the second experiment they examined the effect of nicotine on learning. Bees were trained to associate one flower color with a reward (sugar water alone or plus one of the three nicotine concentrations), and another color with just water. They then counted how many visits bees made to the rewarded versus unrewarded flower types.

In the third experiment they used the same bees from the second experiment but reversed the color/reward associations. So the flower color that had been rewarding was now unrewarding and vice versa. They examined how many visits bees made to the two flower types. 

What they found

In the first experiment, bees were actually attracted to nicotine-laced nectar at the low natural concentration, showed no preference at the intermediate natural concentration, and were deterred at the unnaturally high concentration. 

In the second experiment bees with the intermediate natural concentration of nicotine added to their rewarding flowers made more correct choices than bees with the low natural, unnaturally high, or no nicotine in nectar. Nicotine therefore appears to enhance memory (or preference for some other reason, this paper does not distinguish) for rewarding flower types but only at intermediate concentrations. 

In the third experiment when the color/reward pairings were reversed, bees made more visits to the now unrewarding flower types when they had been paired with nicotine. Especially at the highest concentration! Although the unnaturally high concentration did not appear to improve learning, it did cause the bees to remain faithful to that flower color even when it was no longer rewarding. The other two nicotine concentrations also enhanced fidelity to unrewarding flower types, in a dose-dependent manner, with the strongest effect at the highest concentration. 

The takeaway

I am fascinated by how compounds in flower nectar may have psychological effects on pollinators to manipulate their behavior to the plant's advantage. The dose dependent effects in this paper are very interesting. Baracchi and his colleagues show preference but no improved learning at very low concentrations, no preference but improved learning at intermediate concentrations, and no preference or improved learning but strong fidelity at high concentrations. I am interested in the different pollination requirements or conditions that could give a selective advantage to each of these concentrations. 

Plant Spines and Caterpillar Feeding

Patricia Jones

 A tobacco hornworm,  Manduca sexta , struggles with the spines of a purple devil,  Solanum atropurpureum . Photo by Rupesh Kariyat from the cover of  Biology Letters .

A tobacco hornworm, Manduca sexta, struggles with the spines of a purple devil, Solanum atropurpureum. Photo by Rupesh Kariyat from the cover of Biology Letters.

This week's paper is in Biology Letters, lead authored by Rupesh R. Kariyat at ETH Zürich. Kariyat and his colleagues examined the interactions between a moth, the tobacco hornworm, Manduca sexta, and horsenettle plants, Solanum carolinense. In previous research, they had shown that horsenettle fed upon by moth caterpillars grows more spines. In this paper they examine how spines impact the caterpillars. 

What they did

They examined the effect of spines on caterpillar performance three ways. First they used inbred and outbred lines of S. carolinense, because inbred lines have less spines than outbred lines. Second, they used three closely related plant species that vary in the number of spines: S. carolinense, S. aethiopicum, and S. atropurpureum. Finally they experimentally removed spines (cut them off with a razor blade) in both the inbred/outbred experiment and the three species experiment. They then examined how long it took a caterpillar placed on the soil at the bottom of the plant to reach a single leaf at the top of the plant, and how long it took a caterpillar to totally defoliate the plant. 

What they found

Kariyat and his colleagues consistently found that plants caterpillars took longer to travel up plants and defoliate plants when plants had more spines. This effect disappeared when the spines were removed. High spine densities not only appeared to slow caterpillars down, but also made them fall off the plants more often. 

The takeaway

This result is intuitive and supports the function that many of us would have hypothesized for plant spines. But measuring the effect of spines is tricky, because plants that vary in spine number likely vary in many traits, and if you remove spines it may cause the plants to induce defenses (plants can become better defended when damaged as noted above with plants that grew more spines, but they can also increase their concentration of toxic chemicals etc). This paper handles these problems nicely by testing the hypothesis with multiple experimental variations. None of them are perfect, but together they tell a convincing story.