Where did all the spiderlings go? A story about egg-sac parasitism in Arctic wolf spiders

This week we are in a deep freeze in the Montreal area, so it seems somewhat fitting to discuss Arctic spiders.  I’ve discussed the life-history of Arctic wolf spiders (Lycosidae) before, specifically in the context of high densities of wolf spiders on the tundra.    Much of this work was done with my former PhD student Joseph Bowden.  The latest paper from his work was published last autumn, and was titled ‘Egg sac parasitism of Arctic wolf spiders (Araneae: Lycosidae) from northwestern North America‘. In this work we document the rates of egg sac parasitism by Ichneumonidae wasps in the genus GelisThese wasps are fascinating, and we have found them to be very common on the tundra.  There are often multiple wasps in a single egg sac, and as is typical with Gelis, they leave nothing behind: all eggs within an egg sac are consumed.  After fully developed, the adult wasps pop out of the egg sac; the Gelis adults we encountered had both winged forms and wingless females, the latter superficially resembling ants.

A Gelis emerging from a wolf spider egg sac. Photo by Crystal Ernst, reproduced here with permission.

A Gelis emerging from a wolf spider egg sac. Photo by Crystal Ernst, reproduced here with permission.

The rates of parasitism of Pardosa egg sacs (by Gelis) were, at some sites, extremely high.  In some cases over 50% of the wolf spider egg sacs were parasitized.  Stated another way,  half of all the females encountered with egg sacs had zero fecundity because the female was  carrying around wasps within the egg sac instead of spider eggs.

It’s quite interesting to think about these wingless Gelis femalesafter emerging from egg sacs, they end up wandering around the tundra in search of hosts.  Spiders with egg sacs must be encountered frequently enough for the wasps to grab on to a passing wolf spider in order to parasitize the egg sac.  Recall, densities of wolf spiders can be very high in the Arctic (4,000 per hectare, at least).  Hmmm…. this is all starting to fit… high densities of wolf spiders support high rates of egg parasitism and these wasps can ‘afford’ to be wingless since their hosts are frequently encountered:  an interesting feedback loop!   We can also speculate about large-scale gradients in diversity – many Ichneudmonidae show high diversity in northern regions.  Within Gelis, it’s a good bet that they will find many suitable spider hosts in these environments.

Looking down the microscope - all those Gelis!

Looking down the microscope – all those Gelis!

So, how extreme are these rates of egg parasitism?  Looking at some of the literature, there are certainly a number of papers about  wasps that parasitize spider egg sacs.  Cobb & Cobb (2004) studied two Pardosa species in Idaho, and recorded a egg parasitism rate of about 15% (by Gelis wasps and wasps in the genus Baeus [Sceleonidae]). Van Baarlen et al (1994) studied egg parasitism in European Linyphiidae spiders and their maximum rates of parasitism were about 30%.   Finch (2005) did a detailed study of four spiders species (non-Lycosidae) and rates of egg parasitism varied between 5% up to as high as 60% in an Agroeca species.

Our documented parasitism rates for Arctic wolf spiders are certainly quite high (for Lycosidae), but not out of the range of other published studies for non-Lycosidae.  I do wonder whether we will continue to find high egg parasitism rates if more species were examined in detail – certainly a fertile area of study.  Related to this, what are the population-level consequences of this interaction?  What is the relationship between spider densities and parasitism rates?  Although Joe and I did try to speculate on this, our data are preliminary – again, a key area for future research.

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In the Arctic context, we will continue to uncover fascinating food-web dynamics.  Our research group has already been thinking seriously about this – Crystal Ernst has written a nice post about the idea of an ‘inverse trophic web’ (i.e., predator-dominated) in the Arctic, and a fair amount of my future research will pursue this avenue of research.

Pique your interest…?  Why not think about graduate school in my lab, and study Arctic arthropod biodiversity?

References:

Bowden, J., & Buddle, C. (2012). Egg sac parasitism of Arctic wolf spiders (Araneae: Lycosidae) from northwestern North America Journal of Arachnology, 40 (3), 348-350 DOI: 10.1636/P11-50.1

Cobb, LM & Cobb VA (2004). Occurrence of parasitoid wasps, Baeus sp and Gelis sp., in the egg sacs of the wolf spiders Pardosa moesta and Pardosa sternalis (Araneae: Lycosidae) in southeastern Idaho. Canadian Field Naturalist 118(1); 122-123.

Baarlen, P., Sunderland, K., & Topping, C. (1994). Eggsac parasitism of money spiders (Araneae, Linyphiidae) in cereals, with a simple method for estimating percentage parasitism of spp. eggsacs by Hymenoptera Journal of Applied Entomology, 118 (1-5), 217-223 DOI: 10.1111/j.1439-0418.1994.tb00797.x

Finch, O. (2005). The parasitoid complex and parasitoid-induced mortality of spiders (Araneae) in a Central European woodland Journal of Natural History, 39 (25), 2339-2354 DOI: 10.1080/00222930500101720

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Taxonomic sufficiency in biodiversity research: Is it always necessary to identify species?

It’s been a successful few weeks in the lab!  Two weeks ago I promoted an exciting paper about spider silk and herbivory and just after that paper come out, another publication from our lab was published, titled: “Does species-level resolution matter? Taxonomic sufficiency in terrestrial arthropod biodiversity studies“.  This paper evolved out of a past graduate-level class in Forest Entomology at McGill, and was re-worked and re-written by post-doc Laura Timms, former Phd student Joseph Bowden, and my colleague Keith Summerville.

Let me provide a plain language summary of this work and I will also touch upon some of the controversy that has arisen because of this paper:

Biodiversity science is about the discovery and description of all the different kinds (species) of organisms living on our planet.  It is a vitally important area of research because different species play important roles in our ecosystems, and as a consequence, are important to us.  The different number of species in an area can also inform us about how we might be harming or helping ecosystems.  This is an active area of study in the context of forestry, since some forest practices (for example, cutting all the trees down in an area) can cause changes in the number of species (and whether they are rare or common) and these changes can inform us about whether our forestry practices are harming our ecosystems.  All of this kind of work, however, depends on the ability of scientists to collect, sort, and identify different kinds of species.  Since most described species on the planet are Arthropods (e.g., spiders, insects, and their relatives), these animals are often used as a way to indicate how biodiversity might be affected by environmental change.  However, there is a problem: it takes a very long time to identify different arthropods, and it is costly and difficult – requiring highly specialized training, by people known as taxonomists.  In our research project, we asked whether not you always need to know the exact differences between insects and spiders  in order to tell if a disturbance is affecting biodiversity.  We did this by looking at a series of data-sets about beetles (Coleoptera), moths & butterflies (Lepidoptera), and spiders (Araneae). These data-sets were from past research projects about how forest disturbance affects biodiversity.

Here is how we did the work: Different kinds of organisms are classified using a two-part name:  the genus and the species.  There can be many different species within one genus.  You can then classify different genera (the plural of genus) into grouping called Families.  For example, all wolf spiders are in the Family Lycosidae.  A common genus within this family is Pardosa – there are dozens of species of Pardosa in Canada; Pardosa mackenziana, Pardosa moesta, Pardosa hyperborea, etc.  We first took our big data-sets and using the lowest level of naming (the species) we asked whether forest disturbance affected biodiversity.  We then grouped all our species into their respective genera -this meant that the data-sets got smaller (i.e., there are necessarily fewer genera than species).  We did the same analysis to see if we could still get a signal about the effects of disturbance on biodiversity, but now with the ‘reduced’ data.  We did this again at the family level.  We did this because we wanted to know if you could take a short-cut. Stated another way, if you don’t have the time or ability to figure out all the species in your research project, can you still see if there is an effect of forestry on biodiversity?

A wolf spider (Lycosidae)

A wolf spider – do you need to know its name?

Our results showed that in most cases, you do not need to know the species identity to see the effects of forestry practices on the biodiversity of spiders, beetles and moths & butterflies – you do not get as clear answers when things were grouped into Families, but the datasets with species grouped into genera were almost as good as when you group things into species.  This was surprising, because an assumption in biodiversity science is that species-level identifications are necessary and should be the ‘gold standard’ for this kind of research.  We showed that in many cases, you can get your answer by identifying arthropods to the generic level:  this can save you a lot of time (and money).   Some researchers (including taxonomists) may not be thrilled with this result as it might suggest that species are not important, and specialized taxonomic knowledge is not essential to complete biodiversity research.  This is certainly not the case, which leads me to the caveats:

1) Our results do not mean species are not important!  Instead, we are saying that if there are logistical and financial constraints, you might be able to answer your research question without having to identify all the species.   If you have a project about large-scale disturbance and are looking to see whether there are any broad affects on biodiversity, our approach might work.   However, you might miss some subtle effects, so this approach must be taken with caution.  Although our suggestion is a short-cut, it would still be important to save all the samples, and at a later time (as money and expertise permits) the species could be determined.

2) Our study is specifically geared towards research about insects and spiders in relation to large-scale forestry disturbances.  We are not saying that this will work in all situations and with all different kinds of organisms! The context is important.  Related to this, if an overarching research question is about species in an ecosystem, species-level identifications are essential.  Everything depends on the research question and the research context.

3) This general approach that we have discussed is highly dependent on what kind of organisms you are studying.  If you are working with a group of organisms that do not have too many different species within a genus, our approach may work.  If, however, there are many species within a single genus, our suggestion will not work as well.  Therefore, a researcher should look at the general relationship between the number of species per genus for their study organisms and use this ratio as a guide when thinking about taking the short-cut that we discussed in the research.

In sum, we are quite excited about this research – we think it will provide more opportunities for biodiversity projects to get done, and will help answer certain research questions when there are substantial constraints on time and money.  This is one way to be pragmatic about biodiversity research.

Please share your thoughts!

Reference:

Timms, L., Bowden, J., Summerville, K., & Buddle, C. (2012). Does species-level resolution matter? Taxonomic sufficiency in terrestrial arthropod biodiversity studies Insect Conservation and Diversity DOI: 10.1111/icad.12004

Life History of Arctic Wolf Spiders: Part 1

For those of you who follow my blog, you will notice I’m somewhat obsessed with the Arctic – in part because of our large Northern Biodiversity Program, but also because it’s an ideal  system for studying the ecology of arthropods.    It also doesn’t hurt that the Arctic is a beautiful place to work!

The northern Yukon landscape: spider habitat

I am very excited to write about the latest paper published from our laboratory, titled Life history of tundra-dwelling wolf spiders (Araneae: Lycosidae) from the Yukon Territory, Canada.  This has just recently been published in the Canadian Journal of Zoology, with Dr. Joseph Bowden as the lead author.  Dr. Bowden graduate from my laboratory just over a year ago, and is now living in California with his family.  Although the climate is somewhat warmer in California compared to the Yukon, he’s still actively working on research related to the biology of Arctic arthropods.   Dr. Bowden was a terrific student in my laboratory, and has already published some work about the community ecology of Arctic spiders: he has one paper in the journal Arctic and another in Ecoscience.

Dr. Joseph Bowden, working in the Yukon and ready for the biting flies!

In the CJZ paper, Joseph studied three species of tundra-dwelling  wolf spiders (family Lycosidae) and asked whether body size or condition better explained variation in fecundity and relative reproductive effort (defined as the ratio of female body mass to clutch mass).  He also tested whether  a trade-off exists between investment in offspring size and number.  The field work for this research was really enjoyable, as it involved collecting spiders by visual surveys and dry pitfall traps – after collection, Joseph set up a laboratory in a local campground shelter to do measurements on the species:

Dr. Joseph Bowden in a Northern “laboratory”

One of the main findings was that body size explained well the variation in offspring number.  Stated another way, larger female wolf spiders produced more eggs, a finding well supported in the literature.   A second main finding was that females with a lower condition allocated relatively more to offspring production than did females in better condition. This makes some sense – if the going is tough (i.e., poor condition), the females primary objective (from a fitness perspective) is to invest in offspring.  A third key finding was that  we found a negative relationships between egg size and number.    These trade-offs may in part be because of variation in resource availability at some of the study sites in the Yukon tundra.

An Arctic Pardosa (Lycosidae) female, with egg sac

Joseph also calculated tundra wolf spider densities.  Here’s the text of the CJZ paper that describes the methods (straightforward but time consuming):

Densities of the three focal species were estimated using a ring of hard plastic measuring 1.13 m in diameter (1 m x 1 m area) and about 12 cm high. The ring was haphazardly and firmly placed on the tundra surface in each site and all wolf spiders collected inside the ring were identified and counted. This protocol was adapted from Buddle (2000).

Results? Well… the most common species Pardosa  lapponica averaged about 0.4 spiders per square metre.  Some simple calculations will tell you just how common wolf spiders are on the Tundra:  4000 wolf spiders per hectare.  Don’t forget – wolf spiders are only part of the Arachno-fauna in the Arctic.  With confidence, this estimate of 4000 spiders per hectare represents a minimum.  There are a LOT of Arachnids living on the tundra!

In sum, this paper by Joseph is about studying some good old-fashioned natural history of a fascinating group of animals.  The methods are straightforward, but the findings are significant.  It’s pretty difficult to progress in ecology without a deep understanding of a species’ biology and life-history.  Life-history studies are the cornerstone of biology, and I’m thrilled that Joseph recognized that fact and did this research on Arctic wolf spiders.

    You will see that this post is titled Part 1:  some more work will hopefully be published soon – stay tuned for Part 2…

References

Bowden, J., & Buddle, C. (2012). Life history of tundra-dwelling wolf spiders (Araneae: Lycosidae) from the Yukon Territory, Canada Canadian Journal of Zoology, 90 (6), 714-721 DOI: 10.1139/z2012-038

Bowden, J., & Buddle, C. (2010). Determinants of Ground-Dwelling Spider Assemblages at a Regional Scale in the Yukon Territory, Canada Ecoscience, 17 (3), 287-297 DOI: 10.2980/17-3-3308

Buddle, C. (2000). LIFE HISTORY OF PARDOSA MOESTA AND PARDOSA MACKENZIANA (ARANEAE, LYCOSIDAE) IN CENTRAL ALBERTA, CANADA Journal of Arachnology, 28 (3), 319-328 DOI: 10.1636/0161-8202(2000)028[0319:LHOPMA]2.0.CO;2

Bowden, J. & Buddle, C. (2010). Spider assemblages across elevational and latitudinal gradients in the Yukon Territory, Canada.  Arctic 63(3): 261-272 http://arctic.synergiesprairies.ca/arctic/index.php/arctic/article/view/1490

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