The Green Meshweaver gets to Hull

 

Another post on a rapidly expanding species, this time a small spider, the Green Meshweaver, Nigma walckenaeri. All started on Saturday, during the YNU AGM in the University of Hull, where spider expert  Geoff Oxford, showed us a specimen he had just collected from the university grounds, a tiny green spider that is currently expanding its range across the UK and now colonising Yorkshire. We all admired the individual, which had been collected with its web on a leaf. 

The first Nigma walckenaeri record from Hull.

After returning from the meeting, I searched the garden ivy, and since I've been searching ivy and holly, which are favoured leaves to weave it's mesh. Nigma walckenaeri  chooses curled leaves, so it can take advantage of this to hide under its little web. As other regular character in this blog, Amaurobius, Nigma is a cribellate silk weaver. 

Today, I took a local walk and came across a magnificent mature ivy, south east facing. I started searching its leaves, wondering if I had the wrong search image in my mind. It didn't take long to find a Nigma.

Nigma walckenaeri  habitat.

There it was! A light-touch web of blueish threads with the green spider sitting, well camouflaged, underneath.

I moved part of the web aside to have a closer look (also top shot). This is a female, with her whole body green. Once I had seen one, I found another, and another, almost every ivy leaf had it's little Nigma in it. 

A male, with its brown cephalothorax.

This individual was busy weaving its web.

I even spotted a male (with reddish-brown opistosoma) on the edge of a female's web.

Until 1993 it was only found in London and the home counties, and it is associated to parks and gardens. Since then it has been steadily spreading north. I'll keep a look for it in the garden!

Urban Purple Hairstreaks

I had a most unexpected and delightful walk to work this morning. As I was about to cross Westbourne Avenue, where a water leak has been repair, a passer-by flushed a small butterfly from the barriers. I thought it was probably a Holly Blue, but when it settled with wings closed on a privet hedge nearby, I realised it was a hairstreak! I fumbled for my camera as it fluttered along the hedge into a front garden, where it sat with wings open. With shaking hands, I was able to take a photo (below), confirming the id as Purple Hairstreak! 

The first Purple Hairstreak

The Avenues is a very leafy part of Hull, with large gardens often holding large trees. In fact, there is a mature, spectacular oak at the back of the Adult Education Centre on Park Avenue not far away, and, as I walked past, I wondered if the wind might have blown the small butterfly away. I carried onto Jack Kaye fields. A Speckled Wood by the entrance to the fields, by an oak, reminded me to do a butterfly count, and I set an alarm on my phone to 15 min. I counted six Speckled Woods, as they squabbled around the trees. Then I moved onto the first glade. A small butterfly settled on a Whitebeam to bask in the sunshine. As I focused my camera, I got the feeling of being in a dream: it was another Purple Hairstreak! It has its wings closed and I took plenty of photos. 

Tree canopy butterflies

Purple Hairstreaks are canopy specialists, they spend much of their time on tree tops, around oaks (the larval food plant) and nearby trees, where they feed on honeydew. Although other species that obtain most of their food from honeydew, like Speckled Wood and Holly Blue, can occasionally be found feeding on flowers, or resting or fluttering low down, Purple Hairstreaks rarely come down to the lower level of trees, but they have been seen feeding on bramble or hemp-agrimony. They live up in the tree tops, and it is for this reason that they are easily missed. There are only 18 records of this species in East Yorkshire, mostly in nature reserves, but also at Snuff Mill Lane, just outside Hull, where I saw them in July and in Beverley Westwood, which has a sizeable population. They are not only found in woodland, but also along lanes and in parks with oaks. But, as these lovely, tiny butterflies, spend their adult lives up in tree canopies gorging on honeydew, they are likely to be overlooked and under-recorded. In the last few weeks I've been looking for them at Jack Kaye, staring into that small oak by the entrance, so although I was beyond pleased to find it, I was a bit astounded that it had actually happened. 

I will keep an eye on this area to check if these are isolated individuals or if there is a colony, and I plan to keep looking at oaks, wherever they are!

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).

Budapest slug mating

It's the last day of October: damp, windy and mild. As I go out into the garden, pondering upon the impending winter lockdown, I find two Budapest slugs circling on a cherry leaf on the ground. Budapest slugs remind me of hedgehog poo, dark and shiny and all the right shape. If they had been on the path pebbles I would have easily missed them. I watch them for a few minutes. They keep slowly circling, head-to-tail for a while as they follow each other's mucus trails. This is slug courtship and it happens at slug pace. It's 10:40 and it doesn't look like much is happening any time soon. I regularly go out to check on them for the rest of morning and afternoon. As I'm typing this, I grab a torch and decide to check on them again. They are still there, exactly in the same position as 3 hours ago. 

10:42

11:06. The pair have moved onto the path pebbles now, the circling has stopped. They are now finding each others genital openings at the right side of their heads.

11:20. Copulation proper appears to start.

12:20

14:31

17:20. Very little change in the last few hours. The slugs are now entwined and practically immobile.

I pick the Slugs of Britain and Ireland and check the species account for info on their mating behaviour. Of note is that they are mainly subterranean and active year round.

This quote is fitting: 

"in winter adults are often found almost motionless, mating. Mating lasts many hours and involves the production of elaborate spermatophores"

I wonder if they'll be there tomorrow.

More information

Rowson, Ben, James Turner, Roy Andreson and Bill Symondson Slugs of Britain and Ireland. 2014. FSC Publications. AIDGAP.

Emperor home ranges, habitat use and differences between sexes

Emperors (Anax imperator) have done very well this year in my local area, which has allowed me to become familiar with the different behaviours of males and females. Females are quite stealthy, entering ponds to oviposit on floating vegetation, often flying low over the water. In contrast, males display an obvious territorial behaviour, patrolling high over the water of the large pond, lake or slow running drains where they breed. Males move towards passing birds, and towards any intruders, with spectacular fights and clashes between the males, some times resulting on individuals falling on the water.

Ovipositing female emperor.

A male patrolling alongside the marginal vegetation of a large ditch.

All this means that we are biased in our understanding to these more obvious behaviours, ovipositing and territoriality. But many other questions remain about Emperor's behaviour and ecology, for example, where are females when they are not ovipositing? or males, when they are not patrolling a pond? how far do individuals roam or disperse? where do they roost? how is their behaviour affected by temperature? do they make use of the habitats around ponds? what about sex differences?

Capture-mark-resighting techniques have be used in dragonflies to study individual movements, behaviour or demographic parameters. Individuals of large dragonflies can be marked using alpha-numeric unique wing codes written with permanent markers, which could be read from a distance, with no need from physical recapture (for example using binoculars). This technique, however, requires an enormous field effort to relocate as many individuals as possible.

Another technique that has been used to study migration is to analyse location-specific isotopes in wing samples. These isotopes have signatures specific to geographic areas, which point at the area where the larval stage took place (as the wing tissue is formed during the larval stage). This method has been used to reveal the multi-generational migrations of the Green Darner (Anax junius) a North American relative of the Emperor that is a long-distance migrant. Hydrogen stable isotopes showed that the migration cycle comprises a north migrating generation, a south migrating generation and a resident generation that develops around the Gulf of Mexico. Another study using stable isotopes on the Global Skimmer, Pantala flavescens, revealed its multigenerational migration steps around the Indian Ocean. Although ranging from South Africa to Sweden, and still involved in natural colonisation towards the north, facilitated by global heating, the Emperor is a resident species, so this technique is not of much use.

Radio transmitters in dragonflies?

As technologies have resulted in the miniaturisation of radio-transmitters  in recent years, they have increasingly been used to study more local movement patterns in large dragonflies, like the North American Tiger Spiketail (Cordulegaster erronea) a relative of our Golden-ringed Dragonfly and Green Darners. The individuals need to be captured and fitted with tiny transmitter before release, and they have to be found in the landscape using scanning receivers fitted with an antenna. Size matters as the transmitter must be less than 30% of the weight of the dragonfly not to impede normal behaviour. The Emperor, one of our largest dragonflies, weighs about 1 g, and can be fitted with such small transmitters (check the photo here of an individual fitted with a transmitter). In an early small scale study (5 tagged individuals), researchers looked at home ranges and local movements between roosting sites and pond territories in male Emperors. The furthest moving male travelled 1.5 km from the tagging pond to another pond.

An article published recently sheds some light on home ranges and habitat use of Emperors using radio-transmitters. Marceau Minot and his collaborators chose five ponds in an urban/rural interface in northern France. Over the summers of 2017 and 2018 they marked 87 mature emperors with unique wing codes and visited the ponds at least once weekly to search for marked individuals. They also fitted 54 individuals with radio transmitters, and tracked them daily for up to 15 days.

Capturing individuals to fit radio transmitters or mark their wings has a cost. Both techniques increase mortality in the day after capture, probably due to the stress of the capture. 

Sex differences in behaviour

Females had larger home ranges than males. The furthest distance a female travelled was 1.9 km while males moved less, with the maximum male movement 0.5 km. This could be related to male's territorial behaviour. Presumably females oviposit in several ponds.

Reproductive behaviour of males, but not females, is positively related to temperature.

Flying behaviour in females is positively related to temperature, while resting high in trees is negatively related to temperature.

Both sexes were mostly found on ponds or pond marginal areas, although males spend more time near water.

Resting happened in ponds and trees. Females tend to roost high on trees, more than males. While males tend to rest low in vegetation.

A male emperor resting on marginal vegetation.

The researchers estimated the effect of marking protocol on survival of the dragonflies. The manipulation of the individuals affected their survival, possibly due to the stress during manipulation: 76% of individuals survived 24 after capture and wing marking, while just 56% survived 24h after being fitted with a radio-transmitter. Individuals with proportionally larger wings (not larger body mass) and younger in age survived better throughout the experiment.

Habitat management

The study also highlighted that both rural and urban ponds will benefit from the presence of neighbouring trees as suitable roosting sites, and emperors will benefit from the presence of a network of ponds, rather than isolated ponds.

More information

Hallworth, M. T., Marra, P. P., McFarland, K. P., Zahendra, S. & Studds, C. E. Tracking dragons: stable isotopes reveal the annual cycle of a long-distance migratory insect. Biol. Lett. (2018).

Hobson, K. A., Anderson, R. C., Soto, D. X. & Wassenaar, L. I. Isotopic evidence that dragonflies (Pantala flavescens) migrating through the Maldives come from the northern Indian subcontinent. PLoS One 7, e52594 (2012).

Knight, S. M., Pitman, G. M., Flockhart, D. T. T. & Norris, D. R. Radio-tracking reveals how wind and temperature influence the pace of daytime insect migration. Biol. Lett. 15, 20190327 (2019).

Levett, S. & Walls, S. Tracking the elusive life of the Emperor Dragonfly Anax imperator Leach. 27, 59–68 (2011).

Minot, M., Besnard, A. & Husté, A. Habitat use and movements of a large dragonfly (Odonata: Anax imperator) in a pond network. Freshw. Biol. 46, 207 (2020)