Showing posts with label nectar robbing. Show all posts
Showing posts with label nectar robbing. Show all posts

Wednesday, September 27, 2017

Cheating in mutualisms? Nectar-robbing and nectar-thieving

As a pollination ecologist, I study the mutualistic relationship between plants and their pollinators. Mutualism is roughly defined as an interaction between organisms in which both partners benefit in some way. Pollination is often treated as a classic case: the pollinator gets nectar (i.e., food), and the plant gets pollination (i.e., reproduction).

However, it's worth remembering that neither participant is engaging in the interaction for the benefit of the other: in broad terms, the plant doesn't offer nectar to help out the pollinators, it does it because offering nectar improves the plant's success; the pollinator doesn't pollinate in order to help out the plants, it does it because foraging for food in flowers improves the pollinator's success. I've talked about plants that are jerks a few times before (1, 2, 3); today I'm going to talk about the flip-side of that, the 'pollinators' who are jerks. In this case, when I say that a plant or a pollinator is a "jerk", I mean that it has developed an adaptation that allows it to gain the benefits of the plant-insect interaction without offering the interaction partner any benefit -- in other words, it's a cheater. Deceptive plants fall under this umbrella, because they deceive pollinators by seeming to offer a reward, but without actually doing so and thus without incurring the cost of making the reward (usually nectar). There's some really cool research that has been done on the evolution of cheating in mutualistic relationships, which I may talk about another time. For today, I'm going to focus more narrowly on floral larceny.

So what is floral larceny? The general idea is that the putative pollinator is obtaining the reward without offering the service. So a floral visitor gets nectar without moving any pollen. Nectar-robbing is a frequently-studied form of floral larceny: the visitor, rather than trying to get in through the regular opening of the flower, just cuts a hole near the nectary and sucks up the nectar, avoiding contact with the reproductive parts of the flower and consequently providing no pollination service. In principle, certain floral shapes are adaptations that exclude bad pollinators and improve the fit of good ones by orienting them in particular ways in the flowers, but at least some of these flowers, particularly with long, tubular flowers, and especially the more rewarding ones, are more likely to be the targets of nectar robbers (Rojas-Nossa et al. 2016), so the extent to which they're actually excluding bad pollinators, as opposed to converting them into nectar robbers (which might be worse? Or not, see below), is unknown. Actual measured consequences of floral larceny on floral fitness actually range from negative to positive, which further complicates interpretation of nectar-robbing behaviour (see Irwin et al. 2010 Annual Reviews of Ecology, Evolution, and Systematics).

I managed to get some footage of a nectar robber on Impatiens capensis (jewel-weed). You can see that the robber bites a hole in the flower to get at the nectar; it's quite clear that the nectar-robbing wasp is bypassing the reproductive parts of this flower, but I witnessed several wasps engage both in nectar-robbing, and then in the more standard foraging that involved entering the flower and possibly transporting pollen. 



Similarly, if you look at the Xylocopa virginica (carpenter bee, notably one of the largest insect pollinators in this region; there are several shown in the videos below) on the hostas in the video below, first you can really clearly see in the slow-motion video as she pushes her tongue through the flower into the nectary, but in the next video in real-time, you can see that in some instances she may be brushing her very large abdomen over the reproductive parts of the plants anyway, and you can actually see some pollen grains on her shiny abdomen as she engages in this nectar-robbing behaviour, so it's not really clear whether she's truly failing to provide pollination services here, even while she engages in fairly classic nectar-robbing.





Another form of floral larceny is nectar thievery: the visitor enters through the normal floral opening, but does not make contact with the reproductive parts of the flower and consequently transfers no pollen. 

I have some footage of visitors engaging in nectar thievery on the hostas in my back yard. The video shows a relatively classical case, where the nectar thievery arises because of a morphological mismatch (i.e., the shape of the insect and flower don't match up correctly);this nectar thief is just too small to contact the reproductive parts of the flower, but of course that doesn't prevent it from foraging for nectar on the flowers.



There are also forms of floral larceny relating to pollen-robbing (which causes damage in the course of pollen removal, and in which the pollen isn't transmitted elsewhere), and pollen-thieving (no damage, but pollen is not transferred). There is very little information on these phenomena, probably because they would probably be extremely hard to confirm. Unfortunately, I don't have any footage of these. Pollen-thieving in particular might actually be quite common, depending on how we define it: here's some footage of X. virginica (carpenter bee) grooming pollen off herself; grooming is a common bee behaviour. Bees collect pollen to stock their nest cells with it (i.e., it's food for developing larvae), so a large quantity of the pollen they collect ends up not on other flowers but instead in the bees' nests. This might be considered a form of pollen theft, depending on how you want to define it. Pollen thieving is a rather understudied area, but there's an interesting review for those interested (Hargreaves et al. 2009).



Bonus, partial answer to one of the questions I raised in my first post about I. capensis (why are they shaped like this), here's Apis mellifera (honeybee) grooming herself after visiting I. capensis. Notice that the big patch of pollen between her wings isn't getting removed. Possibly, then, the shape of I. capensis helps to ensure that pollen is deposited on a part of the pollinator where it's less likely to get groomed off and therefore lost as food for bee larvae. The shape could also be at least partially driven by improved accuracy of pollen deposition onto stigmas; if the pollen ends up just anywhere on the pollinator, it might not be very accurately transmitted onto the stigmas of other flowers.



For fun, here's some footage of a Bombus sp. (bumblebee) worker who is definitely picking up pollen as she goes, though it's less obvious whether she's successfully depositing it on stigmas. Look at all that pollen on her abdomen! Pretty much whenever she enters and leaves the flower, she's brushing right up against the reproductive parts of the hosta:


Saturday, July 4, 2015

Leave This One to the Butterflies: Asclepias syriaca - Common Milkweed

Today we have a widely-known plant, Asclepias syriaca (common milkweed). I have posted briefly about this species before. It is native to North America (US range map here, Canada range map here). This species is weedy [1], which likely explains why it has managed to become established in Nova Scotia and PEI [2], which are not originally part of its native range.

Asclepias syriaca whole plant
Asclepias syriaca is a member of the Apocynaceae (dogbane family) [2]; several genera of this family produce latex, including Asclepias spp. [3]. Asclepias syriaca also produces some chemicals (specifically cardiac glycosides) which can be quite toxic to humans and livestock, so although parts of the plant are edible, the seed pods and mature leaves are not to be consumed [4]. Seriously, the effects go all the way up to coma [5] (although it would take a very large dose to experience such serious effects [4]), don't just go chowing down on this plant despite it being listed as edible with various sources. The edible portions include the young shoots and the flower buds [4,5,6,7]. Given that they are edible, I had a taste of the flower buds. They were acceptable, but sort of bland with a vague hint of generic 'green' taste; they may be better cooked than they were raw, but some other reports suggest that though edible and nutritious, they're not really much to write home about flavour-wise [5].

Asclepias syriaca inflorescence
Asclepias syriaca has a long history of use in traditional medicine for a wide range of ailments [5].

Asclepias syriaca inflorescence
This plant is evidently much more palatable food to monarch butterflies (Danaus plexippus); Asclepias syriaca is the primary food source of the monarch butterfly larvae [4,5,7,8,9,10,11]. The cardiac glycosides apparently help confer a protection to the larvae and caterpillars by making them toxic [5], which is pretty cool. So I guess if you can't make a poison yourself, get poison elsewhere and incorporate it into your flesh. Pretty hardcore. Some other insects that favour this plant as a food source include the large milkweed bug, common milkweed bug, red milkweed beetle, blue milkweed beetle, bees, wasps, butterflies, moths [5,6,8,9,10]. Anecdotally, I frequently see ants collecting nectar from this plant, although I am probably witnessing nectar-robbing, as ants usually don't provide pollination services (pollen doesn't stick to their armour much, so it doesn't get transferred).

Asclepias syriaca being visited by ants - photo I posted in the previous post about this species
Anyway, my point here is that although Asclepias syriaca is edible, it's not all that tasty, so you're better off leaving it to the monarch butterflies and the other insects that prefer it -- especially considering that the monarch butterfly is potentially under threat and currently being assessed for potential endangered status [12]. I will say (again, purely anecdotally), that I have seen none of the usual monarch butterflies that I am accustomed to seeing at my family's land in the Upper Gatineau; in spite of this, I am reserving judgement on the question of the species' endangerment until the assessment report is released.

Asclepias syriaca inflorescences
Asclepias syriaca is a serious nectar-producer, and it certainly announces that fact loudly; it has a very strong, sweet, pleasant scent that is strong enough to be easily perceptible to humans. Given the large quantities of nectar it produces, it's no surprise that it is a very attractive plant for a whole lot of insects.

Asclepias syriaca - perfect flower close-up - photo posted in my previous post about this species
The flowers of Asclepias syriaca are bisexual or perfect, meaning that they have both male and female parts [6]. This species is primarily outcrossing, meaning that it relies on pollen being brought from another individual in order to reproduce successfully [6]. Good thing it attracts so many willing assistants with the nectar reward it offers for their trouble!

Saturday, May 31, 2014

Dutchman's Breeches - Dicentra cucullaria - Dicentre à capuchon

(English below)

Cette fleur fait parti de la famille Papaveraceae (pavots). La plante entière est toxique et grâce à cela elle évite de se faire consommé par les chevreuils. La prédation des chevreuils a des effets nuisibles aux fleurs du printemps.

D. cucullaria
La toxicité de Dicentra cucullaria est grâce aux alcaloïdes qui s'y retrouvent.

D. cucullaria est pollinée par une espèce unique d'abeille: Bombus bimaculatus. La reine de B. bimaculatus sort de la terre le printemps en même temps que D. cucullaria commence à fleurir et elle est la seule insecte capable d'obtenir le nectar. Le nectar se retrouve dans les éperons de la fleur, qui sont en haut; pour y accéder, il faut séparer les deux pétales extérieures de la corolla. Ceci exige plus de force que la majorité de pollinateurs sont capable d'atteindre. Seule la reine de B. bimaculatus est capable. Les deux espèces, par conséquence, ont des impactes l'un sur l'autre sur le trajet évolutionaire. Ceci agit de coévolution.

D. cucullaria inflorescence; inflorescence de D. cucullaria

Comme le nectar est très attirant et vaut la peine d'obtenir, il y a d'autres insectes qui on une autre stratégie. Il y a des insectes qui percent le corolla pour y voller le nectar, en ce faisant évitant l'exigance de force pour le prendre.

D. cucullaria with evidence of nectar robbing (note the holes in the corolla); évidence du vol de nectar sur D. cucullaria (noter les trous dans le corolla)
This flower is part of the Papaveraceae (poppy) family. The whole plant is toxic and because of this it has avoided deer predation. Deer predation has had serious negative impacts on spring ephemerals in many places.

D. cucullaria's toxicity comes from a number of harmful alkaloids found in its tissues.

This species is pollinated by Bombus spp. queens, primarily a single species: Bombus bimaculatus. The B. bimaculatus queen emerge from underground in the spring at the same time that D. cucullaria starts to flower and she is the only insect able to get the plant's nectar. The nectar is stored in the nectar spurs of the flower, which are pointing upward in this flower. In order to get the nectar, it is necessary to separate the two outer petals of the corolla; this requires considerable strength, which the majority of pollinators simply do not have. Only the B. bimaculatus queen does (of the pollinators out during the blooming season of D. cucullaria). The two species have thus impacted each others' evolutionary processes. This is called coevolution.

Given that the nectar is attractive and is worth some effort to collect, there are also insects who use a different strategy to get it. There are nectar robbers as well, who pierce the corolla to get the nectar, thus avoiding the strength requirement. Naturally, this provides no benefit to the flower (no pollination occurs).