Tag Archives: Cinara piceae

Not all aphids get lost

Although aphids are very good at kicking, we know that aphids would not be very good at football as they are very short-sighted (Doring et al., 2008) but does that mean that they are not very good at finding their host plants? There is a common misperception, and not just confined to non-entomologists, that aphids are no more than aerial plankton. In 1924 Charles Elton

Lost 1

whilst on an expedition to Nordaustlandet* (the second largest of the Spitsbergen group and almost entirely covered by ice) reported finding large numbers of aphids, many still alive, later identified as Dilachnus piceae (now known as Cinara piceae) (Elton, 1925).

Lost 2

Cinara piceae the Greater Black Spruce Aphid –big and beautiful.

 

He suggested that the aphids came from the Kola Peninsula, a distance of about 800 miles (almost 1300 km) due to the strong south and south-east winds blowing at the time. He estimated that they would have made the journey within twelve to twenty-four hours. This was regarded as being an example of totally passive migration and used as one of many examples of aerial plankton** (Gislen, 1948). This is, however, probably not giving aphids credit for what they are capable of doing when it comes to flight. Berry & Taylor (1968), who sampled aphids at 610 m above the grounds using aeroplanes, implied that the aphids, although using jet streams, were flying rather than floating (page 718 and page 720) and that they would descend to the ground in the evening and not fly during the night.

Lost 3

Aphids don’t usually fly during the night. (From Berry & Taylor (1968)).

Dixon (1971) interprets this somewhat differently and suggests that the “movement of the air in which it is flying determines the direction of its flight and the distance it will travel” but then goes on to say “after flying for an hour or two aphids settle indiscriminately on plants”. So yes the speed of the air in which the aphid is flying will determine how far it flies in a set time, but as aphids can fly much longer than an hour or two, active flights of from between 7-12 hours have been recorded (Cockbain, 1961), this rather suggests that the aphids are making a “decision” to stop flying and descend from the jet stream. That said, in the words of the great C.G. Johnson “aphids are weak flyers”, they cannot make progress against headwinds of more than 2 km per hour (Johnson, 1954), although Trevor Lewis gives them slightly more power and suggests that the can navigate against winds of up to 3 km per hour (Lewis, 1964).

Whatever the upper limit is, it doesn’t mean that they are powerless when it comes to ‘deciding’ when to stop flying. In the words of Hugh Loxdale and colleagues, “aphids are not passive objects” (Loxdale et al, 1993). Aphidologists, were until the 1980s (Kennedy, 1986), generally somewhat sceptical about the ability of aphids to direct their flight in relation to specific host finding from the air and not just flying towards plants of the right colour (Kennedy et al., 1961), or at all after take-off (Haine, 1955). The general consensus now, is that aphids control the direction of their flight in the boundary layer*** but that it is determined by the wind at higher altitudes (Loxdale et al., 1993).   Whilst we are discussing viewpoints, another point of debate is on whether aphids migrate or not. Loxdale et al., (1993) state that “migration can be viewed ecologically as population redistribution through movement, regardless of whether deliberate of uncontrolled or from the behavioural viewpoint of a persistent straightened-out movement affected by the animal’s own locomotory exertions or by its active embarkation on a vehicle”. In the case of aphids the vehicle could be the wind. Under both definitions, aphids can be defined as undertaking migrations. Long-distance migration by aphids is defined as being greater than 20 km and short-distance (local) migration being less than this (Loxdale et al., 1993). Long-distance migration is likely to be the exception rather than the rule with most aphids making local flights and not venturing out of the boundary layer, sometimes travelling distances no more than a few hundred metres (Loxdale et al., 1993).

There are different types of winged aphids (morphs) and these show different angles of take-off and rates of climb.  In Aphis fabae for example, which host –alternates between spindle and bean, the gynoparae which migrate from the secondary host to the primary host, have a steeper angle of take-off and climb more rapidly than the alate exules which only disperse between the secondary host plants (David & Hardie, 1988).

Lost 4

http://influentialpoints.com/Images/Rhopalosiphum_padi_emigrant_alate_departing_from_primary_host_c2013-05-21_11-25-12ew.jpg

The gynoparae are thus much more likely to end up in the jet stream and be carried longer distances, with, of course, a greater chance of getting lost (Ward et al., 1998). The alate exules however, may only land in the next field or even in the same one, and easily find a new host plant (Loxdale et al., 1993). These differences between the morphs of host alternating aphids are also seen in the bird cherry-oat aphid Rhopalosiphum padi (Nottingham et al., 1991).  Once safely air-borne, the aphids then have another set of problems to overcome.

How do they ‘decide’ when to land? How do they ‘know’ that there are host plants below them? Aphids have two main senses that help them locate their host plants, vision and smell (odour recognition) (Kring, 1972; Döring, 2014). Generally speaking, aphids respond positively to what we perceive as green or yellow light and negatively to blue and red light (Döring & Chittka, 2007) although this is not an absolute rule. Some aphids are known to preferentially choose yellowing leaves (sign of previous infestation) e.g. Black Pecan Aphid Melanocallis caryaefoliae (Cottrell et al., 2009) which indicates a pretty sophisticated host finding suite of behaviours. Aphids in flight chambers will delay landing if presented with non-host odours even in the presence of a green target (Nottingham & Hardie, 1993) and conversely can be attracted to colourless water traps that have been scented with host plant odours (Chapman et al., 1981). Aphids are thus using both visual and olfactory cues to locate their host plants and to ‘decide’ when to descend from the jet stream or boundary layer (Kring, 1972; Döring, 2014). They are not merely aerial plankton, nor are they entirely at the mercy of the winds, they do not deserve to be described as passive (Reynolds & Reynolds, 2009).

Once at ground level and on a potential host plant, aphids go through a complicated suite of behaviours to determine if the host is suitable or not; if the plant meets all the required

Lost 5

From air to plant – how aphids chose their host plants – after Dixon (1973).

 

criteria, then the aphid will start feeding and reproducing. It is interesting to note that although there may be a lot of aphids in the air, the number of plants on the ground that

Lost 6

Settled safely and producing babies 🙂

http://beyondthehumaneye.blogspot.co.uk/2012/06/aphids.html  https://simonleather.files.wordpress.com/2016/04/cd0a4-aphidbirth2small.jpg

 

are infested with them is relatively low, about 10% in a diverse landscape (Staab et al., 2015), although in a crop, the level of infestation can approach 100% (e.g. Carter et al., 1980). The fact that in some cases less than 1% of those that set off will have found a host plant (Ward et al., 1998) is not a problem when you are a member of clone; as long as not all of the members of a clone gets lost the journey has been a success.

They may be small, they may be weak flyers, but enough of them find a suitable host plant to keep the clone alive and kicking; not all aphids get lost.

 

References

Carter, N., Mclean, I.F.G., Watt, A.D., & Dixon, A.F.G. (1980) Cereal aphids – a case study and review. Applied Biology, 5, 271-348.

Chapman, R.F., Bernays, E.A., & Simpson, S.J. (1981) Attraction and repulsion of the aphid, Cavariella aegopodii, by plant odors. Journal of Chemical Ecology, 7, 881-888.

Cockbain, A.J. (1961) Fuel utilization and duration of tethered flight in Aphis fabae Scop. Journal of Experimental Biology, 38, 163-174.

Cottrell, T.E., Wood, B.W. & Xinzhi, N. (2009) Chlorotic feeding injury by the Black Pecan Aphid (Hemiptera: Aphididae) to pecan foliage promotes aphid settling and nymphal development. Environmental Entomology, 38, 411-416

David, C.T. & Hardie, J. (1988) The visual responses of free-flying summer and autumn forms of the black bean aphid, Aphis fabae, in an automated flight chamber. Physiological Entomology, 13, 277-284.

Dixon, A.F.G. (1971) Migration in aphids. Science Progress, Oxford, 59, 41-53.

Dixon, A.F.G. (1973) Biology of Aphids, Edward Arnold, London.

Döring, T.F. & Chittka, L. (2007) Visual ecology of aphids – a classcial review on the role of colours in host finding. Arthropod-Plant Interactions, 1, 3-16.

Döring, T., Hardie, J., Leather, S.R., Spaethe, J., & Chittka, L. (2008) Can aphids play football? Antenna, 32, 146-147.

Döring, T. (2014) How aphids find their host plants, how they don’t. Annals of Applied Biology, 165, 3-26.

Elton, C.S. (1925) The dispersal of insects to Spitsbergen. Transactions of the Entomological Society of London, 73, 289-299.

Gislen, T. (1948) Aerial plankton and its conditions of life. Biological Reviews, 23, 109-126.

Haine, E. (1955) Aphid take-off in controlled wind speeds. Nature, 175, 474-475

Johnson, C.G. (1951) The study of wind-borne insect populations in relation to terrestrial ecology, flight periodicity and the estimation of aerial populations. Science Progress, 39, 41-62.

Johnson, C.G. (1954) Aphid migration in relation to weather. Biological Reviews, 29, 87-118

Kennedy, J. S., Booth, C. O. & Kershaw, W. J. S. (1961). Host finding by aphids in the field III Visual attraction. Annals of Applied Biology, 49, 1-21.

Kring, J.B. (1972) Flight behavior of aphids. Annual Review of Entomology, 17, 461-492.

Lewis, T. (1964) The effects of shelter on the distribution of insect pests. Scientific Horticulture, 17, 74-84

Loxdale, H. D., Hardie, J., Halbert, S., Foottit, R., Kidd, N. A. C. &Carter, C. I. (1993).The relative importance of short-range and long-range movement of flying aphids. Biological Reviews of the Cambridge Philosophical Society, 68, 291-312.

Nottingham, S.F., Hardie, J. & Tatchell, G.M. (1991) Flight behaviour of the bird cherry aphid, Rhopalosiphum padi. Physiological Entomology, 16, 223-229.

Reynolds, A.M. & Reynolds, D.R. (2009)  Aphid aerial desnsity profiles are consistent with turbulent advection amplifying flight behaviours: abandoning the epithet ‘passive’. Proceedings of the Royal Society B, 276, 137-143.

Staab, M., Blüthgen, N., & Klein, A.M. (2015) Tree diversity alters the structure of a tri-trophic network in a biodiversity experiment Oikos, 124, 827-834.

Ward, S.A., Leather, S.R., Pickup, J., & Harrington, R. (1998) Mortality during dispersal and the cost of host-specificity in parasites: how many aphids find hosts? Journal of Animal Ecology, 67, 763-773.

 

Post script

Political and geographic borders are not factors that deter aphid migrants, Wiktelius (1984) points out that aphids regularly make the journey across the Baltic in both directions to and from Sweden.

Wiktelius, S. (1984) Long range migration of aphids into Sweden. International Journal of Biometeorology, 28, 185-200.

 

*Elton refers to it as North-East Land

** Johnson (1951) objects to this terminology in no uncertain terms. That said, as there are records of non-winged aphids being caught by aircraft (Kring, 1972), it does suggest that there may be some accidental migration going on.

*** The UK Met Office defines the boundary layer as “that part of the atmosphere that directly feels the effect of the earth’s surface” and goes on to say that depending on local conditions it can range in depth from a few metres to several kilometres.

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A Christmas Aphid

A few weeks ago I was contacted by a researcher from the One Show.  They were interested in the possibility of doing a festive piece about what people bring into the house with them on Christmas trees with the idea that George McGavin would shake a Christmas tree over a piece of white paper and tell the audience all about the insects that fell out;  a typical media “how gross nature” is piece.

The researcher was somewhat disappointed when I told her that being winter  that there would be relatively little hiding in the tree, especially if it was a commercially reared cut tree bought from a garden centre or other retail outlet.  Cut Christmas trees in the UK tend to be harvested from October onwards so the chances are that your tree has lain about for at least a month before you bring it into your house and by that time, any sensible winter active herbivore has long departed for fresher trees.  Although conifer trees have a large number of insect species associated with them, most of them spend the winter either off the tree or as inactive eggs hidden under the bark or as eggs actually laid inside the needles e.g. the pine sawfly Neodiprion sertifer.  You would probably find a few opportunistic spiders and possibly some mites and bark lice, but not much else unless you had a potted tree or one that had only recently been felled.  The other thing that would influence what you would find is of course what species of tree you had bought.  Gone are the days when the Christmas tree and Norway spruce (Picea abies) were one and the same.  I guess my caveating and pessimistic reply proved too much for the researcher as I never heard back from her.

The one insect I had waxed lyrical about was of course an aphid, the green spruce aphid, Elatobium abietinum to be precise.   There are a number of aphid species that make a living on spruce trees, some of them quite large and spectacular such as the greater black spruce aphid, Cinara piceae, but like most aphids, they overwinter as eggs (Leather, 1992).

Cinara_piceae_aptera_on_Picea_abies_at_Selwyn_Wood

The greater black spruce aphid, Cinara piceae (Photograph courtesy of http://influentialpoints.com/Gallery/Aphids_on_spruce_Picea_in_Britain.htm)

The green spruce aphid, E. abietinum or Elatobium as it is commonly known, (there is only one species in the genus), overwinters in the UK and most other parts of the world, as an adult or immature stage (nymph) (Nicol et al., 1998).

The adult is small, green and inconspicuous, and quite difficult to see unless you are actually looking for them.

Elatobium and nymphs

The green spruce aphid, Elatobium abietinum and nymphs.

The green spruce aphid is a native of Europe and normally attacks Norway spruce.  They avoid current year needles as these tend to be distasteful to them (the chemistry of young spruce needles is pretty nasty and makes them unsuitable hosts for the aphids) and feed on the previous year and older needles.  Spruce needles, even older ones, are not particularly nutritious, so the aphid injects a toxic material in its saliva that makes the needles more nutritious by encouraging nitrogen mobilisation (Kloft & Erhardt, 1959).  Their populations build up during the spring and towards the end of May and beginning of June, they take flight and the trees seem relatively free of aphids (Bevan, 1966).  As they are so small, they are most obvious after they have gone, either by the damage they cause, premature senescence of the needles as shown in the photograph above, premature needle drop or by the presence of a large number of ladybird larvae.  When I worked for the Forestry Commission as an entomologist, I quite often received phone calls from distressed foresters who had sprayed the blue beetles damaging their spruce trees!

Although they are difficult to find during the summer months they are still there; this summer collapse of singe-host aphids is quite common (Karley et al., 2004).  In the autumn,  Elatobium populations begin to build up and as they do not overwinter as eggs, they are able to continue reproducing through the winter months (Powell & Parry, 1976). Sitka spruce, Picea sitchensis, the most commonly grown conifer in the UK, is a native of North America and as such has very low resistance to Elatobium and displays an almost hypersensitive response to the toxic saliva produced by the aphid.

If it is a particularly mild winter then the spruce trees are likely to show severe signs of damage by June and July.  After several mild winters spruce trees may end up with only current year needles present, which has a severe effect on their growth and appearance.

Elatobium damage needles

Branches of Sitka spruce with only current year needles present after a severe Elatobium abietinum infestation

Elatobium damage trees

Sitka spruce trees showing discoloured needles after attack by Elatobium abietinum.

It may be small, inconspicuous and not worth a TV appearance, but  Elatobium abietinum is now a pest with a world-wide distribution and an international reputation.

References

Bevan, D. (1966). The green spruce aphis Elatobium (Neomyzaphis) abietinum Walker. Scottish Forestry 20, 193-201.

Karley, A. J., Parker, W. E., Pitchford, J. W. &Douglas, A. E. (2004). The mid-season crash in aphid populations: why and how does it occur? Ecological  Entomology 29, 383-388.

Kloft, W. & Ehrhardt, P. (1959). Unterschungen uber Saugtatigkeit und Schadwirkung der Sitkafichtenlaus, Liosomuphis abietina (Walk.), (Neomyzaphis abietina Walk.).  Phytopathologie Zeitzschrqt 35, 401 – 410.

Leather, S. R. (1992). Aspects of aphid overwintering (Homoptera: Aphidinea: Aphididae). Entomologia Generalis 17, 101-113.

Nicol, D., Armstrong, K. F., Wratten, S. D., Walsh, P. J., Straw, N., Cameron, C. M., Lahmann, C. & Frampton, C. M. (1998). Genetic diversity of an introduced pest, the green spruce aphid Elatobium abietinum (Hemiptera: Aphididae) in New Zealand and the United Kingdom. Bulletin of Entomological Research 88, 537-543.

Powell, W. & Parry, W. H. (1976). Effects of temperature on overwintering populations of the green spruce aphid, Elatobium abietinum.  Annals of Applied Biology 82, 209-219.

Sullivan, C.R. (1965) Laboratory and field investigations on the ability of eggs of the European Pine Sawfly, Neodiprion sertifer (Geoffroy) to withstand low winter temperatures.  Canadian Entomologist, 97, 978-993

 

Postscript

During the 1980s when ‘Acid Rain’ was very much in the news, Elatobium damage was often mistaken as a symptom of acid rain in the UK.

 

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