Do Migrant Hawkers migrate in groups?

This summer it appears to be a bumper year for Migrant Hawkers. Their sudden appearance in several places with several individuals present, and the huge numbers seen around makes me think that these are migrants, rather than locally bred individuals. Last week I took a clip of a large group feeding by a nearby wood, a photo would not have conveyed the feeling of watching these dragonflies hunting in groups, paying little notice to one another, in contrast to the two feisty Brown Hawkers in attendance, who squabbled every time they met.

A sunny spell earlier today brought out at least 10 individuals over the garden, the largest number I've had. They are not aggressive or territorial, but, is there more to it? When roosting, they actually appear to choose to rest near other individuals, despite an abundance of sites nearby with a similar aspect, even when breeding (top shot, two mature males basking close to one another at Hornsea Mere, 24 September 2018. 

The following, much more spectacular image was shared on twitter by @jwood_t on the 2nd of August shows 12 Migrant hawkers roosting side by side:

Given that they appear to be a sociaI species, feeding in groups and roosting close to each other, I wonder if they are attracted to one another during migration. Do they actively seek each other, migrating in a compact flock? And why are they social, is there an antipredator advantage, or a better timing and orientation of migration, a quorum decision on directions, or even an energetic advantage, when migrating in groups?

Migration in groups would not be a new phenomenon in dragonflies. A striking example is this amazing photo of migrating dragonflies shared by Dave Smallshire:

My final offering for #worldodonataweekend @Britnatureguide @BDSdragonflies: a unique image taken high over Gomatong Cave, Borneo showing a world first - 2 of about 10 migrating flocks of dragonflies! These can be identified as Rhyothemis obsolescens from their wing pattern. pic.twitter.com/zLlUez9jEV

— Dave Smallshire (@smallshire) May 10, 2020

The Migrant Hawker is indeed known for migrating in large swarms - hence its name - but little experimental research has been carried out on this species. Recent research conducted at a bird observatory on the Baltic coast in Latvia starts to address this gap. Aline Knoblauch and collaborators took advantage of dragonfly 'by-catch' in Heligoland traps during a few weeks in August and September. These large funnel-like traps covered on wire-netting, widely used to investigate bird migration, also capture dragonflies, unharmed. The researchers also used field 'orientation experiments, releasing freshly caught dragonflies into a closed round arena made of mesh and analysing recorded videos to determined if they had a preferred flight direction. All dragonflies were released after the short trials. Their results showed that individuals captured during autumn migration orient themselves to a southerly flight direction irrespective of the prevailing winds, even though there were more captures in the Heligoland trap when the prevailing wind were northerly wind, indicating that the dragonflies were choosing to fly when wind direction was favourable. These results indicate true migration, rather than random accumulations of feeding individuals in suitable areas. This experiment, however, doesn't answer the question of orientation to other individuals.

An observational study on the related species, Southern Migrant Hawker, Aeshna affinis, provides some intriguing clues about group migration, likely to apply to Migrant Hawkers as well: 

"In the late afternoon, thousands of immature individuals were flying above the swamp, mainly at a height of 5-10 m. They formed a huge compact bubble, which was spatially clearly defined on the outside, but with unorganised bee-like swarming inside. Individuals showed a distinctly jerking and dancing flying style, with a minimum space kept between individuals."

This indicates not just agglomeration due to being in the same area with the same intent of migrating, but active flock formation behaviour. The insects also appeared to follow geographical features, a river, to migrate. 

Another interesting feature of Migrant Hawker migration has been noted in a study recording the numbers of Migrant Hawkers migrating on the Danube delta. A strong male-bias of 2 males to 1 female was found, which was also noted in the first study. Although sex-biased dispersal and migration is well known in many animals, it is unclear if this is due to different timing of migration of males and males, or to different geographical or altitudinal pattern of migration of sexes. Note that observational studies are limited to dragonflies flying low, and Heligoland traps only capture dragonflies flying just above ground (within about 3 m).

We are starting to understand Migrant Hawker migration, but there is still a lot of questions to answer.

More information

Knoblauch, A., Thoma, M. & Menz, M. H. M. Autumn southward migration of dragonflies along the Baltic coast and the influence of weather on flight behaviour. Anim. Behav. 176, 99–109 (2021).

Schröter, A. A mass migration of Aeshna affinis in southern Kyrgyzstan: attempt to provide a spatial and temporal reconstruction (Odonata: Aeshnidae). Libellula 30, 203–232 (2011)

Dyatlova, E. S. & Kalkman, V. J. Massive migration of Aeshna mixta and Sympetrum meridionale in the Ukrainian Danube delta (Odonata-Anisoptera: Aeschnidae, libellulidae). Entomol Bericht 68, 188–190 (2008).

Dictynidae: mesh weavers

Credit: Drriss & Marrionn CC BY-NC-SA 2.0 via Flickr.

These are small cribellate spiders, less than 5 mm long, some with characteristic colour patterns. They build an irregular sheet web usually with a central retreat. Many of the 18 British species are uncommon. Some of the species live on plant foliage, others are ground dwellers.
 The top shot shows Nigma walckenaeri a green and white spider that builds sheet webs on the top surface of leaves. This species is mainly found in the Thames and Severn valleys, where it can be common in gardens.

'Unusual friendship' between males and females
 Dyctina arundinacea, the most common species, makes its web atop a dead shoot or flower head. The spider adds more threads as the summer progresses, for, unlike orb weavers, the web is a permanent structure. Bristowe remarked that in this species, the male spends an unusually long time in company of the female in her web (a month or more). Once a male enters a receptive female's web, he signals to her and she signals back with vibrations and tactile signals. The male then goes on to modify the female's web and adding threads to build a canopy, where mating happens. They signal to each other every time they meet. Males and females can even share capture and eating of prey. The female sets her egg sacs inside the web, and she can produce up to six in a season. While a male have to gain from staying around a female, for example, preventing rival males from mating with 'his' female, it is unclear what the female has to gain from accepting the male, unless his cooperation in prey capture helps she build up resources for egg production. More research is needed on these spiders natural history. There are tantalising observations of  males possibly participate in egg guarding. Some non European Dictynidae are social, so it is possible that the roots of social behaviour lay on these unusual male-female associations.

Communal mining bees

A few years ago, a crack in the concrete path at the front of our house became busy with bees coming in and out. We assumed they were honeybees that had settled their hive there, but on closer inspection they turned out to be mining bees instead. Their cleptoparasite bees, Nomada, hung around the nest entrance awaiting an opportunity to get in and lay their eggs. There are many similar species of Andrena mining bees in the UK and several of them looked very similar to mine so I finally gave up with obtaining an ID and I didn't find much more about my front path bee. The bees are back again in the same spot, and a couple of days ago, thanks to tweep @DayMoonRoseDawn I finally knew the identity of these curious bees: they are Andrena carantonica, a large, furry mining bee. Although mining bees are solitary, a handful of species in the UK often nest communally. Communal bees have a simple form of social structure: they share a nest, but individual females will build their own cells, lay eggs and provision their own offspring: they do not have the complex social organisation with workers and division of labour of honeybees. Nesting aggregations of A. carantonica consist of hundreds of females (up to 600) females using the same nest entrance. In my street, cracks on concrete in front garden paths are favoured. Adults emerge in April, with the smaller males doing so just before females and awaiting them inside the nest. In a series of studies on this species, Robert Paxton and collaborators revealed fascinating details of the social structure of this species. In one of these studies they installed nets on nest entrances to be able to capture recently emerged bees: 75% of females were already inseminated when they left the nest for the first time, indicating that copulation happens most of the time inside the communal nests. Although the females in the nest are not highly related, there is a large amount of inbreeding in this species: genetic analysis suggests that 44% of females mate with their full brothers. Emerging females are much more numerous in a given nest (about 1 male to 3 females), although, as Rose Dawn pointed out, it is likely that many males never leave the nest and die inside after having mated, so emergence nets do not capture them. Females must by necessity swap nests after emergence so that the relatedness levels is still low within a given nest, and the bees show a high degree of tolerance to conspecifics, regardless of their nest of origin. If a high level of relatedness is not responsible for the evolution of communality, why do these bees nest communally? A potential benefit is improved nest defence. There are always bees going in or out, and in my own observations, a few bees often sit by the nest entrance (like as in the photo at the top), therefore opportunities for parasitism by cleptoparasites might be reduced. There are still many questions to answer, which probably will have to await to observations being carried out inside nests, but the study of these and other communal species might offer clues as to how and why complex social systems like those of honeybees evolved.

A male Andrena carantonica near the nest entrance

Female Andrena carantonica

A cleptoparasite bee Nomada flava by mining bee nest entrance.

Thanks to Rose Dawn for providing an ID and sharing lots of information on this bee, including results from her own research

More information

Paxton, R., Thoren, P., Tengo, J., Estoup, A., & Pamilo, P. (1996). Mating structure and nestmate relatedness in a communal bee, Andrena jacobi (Hymenoptera, Andrenidae), using microsatellites Molecular Ecology, 5 (4), 511-519 DOI: 10.1046/j.1365-294X.1996.00117.x

PAXTON, R., THOREN, P.A., & GYLLENSTRAND, N. (2000). Microsatellite DNA analysis reveals low diploid male production in a communal bee with inbreeding Biological Journal of the Linnean Society, 69 (4), 483-502 DOI: 10.1006/bijl.1999.0371

Communal daddy long legs

These three daddy long legs (Opiliones), Paroligolophus agrestis, had found a cosy spot at the top of this apple. I am not aware that this species is particularly gregarious, although many daddy long legs do not dislike each others company and some actually form dense aggregations. The function of the social behaviour of daddy long legs is poorly understood and the explanations range from thermoregulation to defence. Although most individuals aggregating are adults, mating does not seem to explain this behaviour. The following video shows an instance of mass aggregation in Daddy long legs: