Posts

, ,

Dung beetles in high mountain landscapes of Oaxaca

By Matthias Rös, Alfonsina Arriaga-Jimenez, Bert Kohlmann

 

Dung beetles (Scarabaeidae) belong, besides ants and butterflies, to the best-studied insect groups in tropical ecosystems. Three subfamilies are considered as true dung beetles: Scarabaeinae, Geotrupinae, and Aphodiinae. There are about 10,000 species of dung beetles around the world known to science, although that number is still rising; montane areas in the tropics are exceedingly rich in species, and new species are regularly discovered. 

High mountain ecosystems in the American tropics have been less studied than the diversity-rich lowland rain forests, which have received greater attention and efforts for conservation purposes. Nevertheless, the significance of temperate ecosystems within the tropics may have been underestimated regarding their importance to explain species distribution patterns in various biodiversity hotspots of the Earth. Mexico, and particularly the state of Oaxaca, will serve us here as an example to explain why. 

Oaxaca is one of the most (if not the most) biodiverse states of Mexico. One reason is the rugged orography, shaped by different geological events, which, accompanied by changing climate, separated and connected animal and plant populations several times, and so turned Oaxaca into a laboratory of species evolution. Oaxaca is situated in the southeast of Mexico and is dominated by three major montane areas (Sierra Norte, Sierra Sur, Mixteca Shield). Eighteen percent of the state has an elevation higher than 2000 m, and around four percent is situated between 2500m and 3700m. 

Typical land-use patterns in Oaxacan mountains. Forest dominated landscapes with traditional milpa system (corn, beans, squash). El Rosario Temextitlan, Chinantla, Sierra Norte de Oaxaca at elevations between 2000 and 2700 m. Photo by Matthias Rös.

In the last two years, we have collected and described new dung beetle species from Oaxaca. All of them were not collected in pristine or remote places, but in mountain forests close to the capital city of Oaxaca. Whereas the state has few large reserves, Oaxaca is known for its high number of community-conserved areas (CCA), and the new species were collected in the CCA La Mesita, in San Pablo Etla, a 3000 ha community-managed forest at altitudes between 1800 and 3200 m, which provides firewood, clean water to the entire watershed, and offers small scale sustainable tourism. In Oaxaca, at lower altitudes, there exists an oak forest, with mostly small trees that lose all their leaves during the dry season, reminiscent of the familiar chaparral vegetation. In Oaxaca, this oak forest is a typical vegetation type of piedmont, mostly surrounding the Central Valley. We named Canthidium quercetorum after this forest type, only known at present from La Mesita. Onthophagus etlaensis, named after the Nahuatl word for bean-fields, sampled by us in the same reserve, had already been collected in the 1970s but was erroneously identified because of its closeness to another, more common species. This is a very typical pattern found in Oaxaca: there abound many endemic sister species of common and more widespread taxa, and they have a small distribution range in the mountains of Oaxaca, which indicates their speciation in situ.  Finally, Phanaeus dionysius, a veritable jewel of a beetle, was also found in this CCA.

Onthophagus etlaensis (left) and Phanaeus dionysius (right), two dung beetle species of the subfamily Scarabaeinae, described from the community-conserved area of La Mesita, San Pablo Etla, near the city of Oaxaca.

Oaxaca belongs to the Mexican Transition Zone, a region ranging between the southern USA down to the Nicaraguan lakes. Its outstanding characteristic is the overlap of Nearctic and Neotropical species distributed here, the former more often at higher elevations with the latter at lower elevations. Both Neotropic and Nearctic faunas have generated a high number of endemic species in Mexican mountains. 

Besides its rich biodiversity, Oaxaca is also one of the most understudied states in Mexico, and regarding plant or animal groups we have only little information. This  might explain why we also found, in addition to the recently described species, some species which were last collected 45 years ago. 

This map shows Oaxaca as depicted by a 3D Digital Elevation Model. Black dots represent sampling sites for Onthophagus anthracinus, the red dot Canthidium quercetorum, and the blue dot Phanaeus dionysius.

AAJ started to work on dung beetle diversity at high-altitude mountains ten years ago when she collected insect material from the alpine prairies of the Trans-Mexican Volcanic Belt (TMVB). For her Ph.D. project, she moved up to high elevations between 2500 and 3500 at four volcanos. One of the most interesting results was that a high variability of diversity patterns between the volcanoes existed. We also found an unexpectedly high diversity, coupled with low abundances and detection probabilities, that in three years of sampling, abundances were still lower than what you collect in one rainy season in a cloud forest. Our next step shall be to compare diversity patterns between the mountains of Oaxaca and the Trans-Mexican Volcanic Belt. Bert Kohlmann has studied for almost three decades the dung beetle communities in the high altitude-mountains of Costa Rica and Mexico, where interesting evolutionary phenomena have been discovered associated with the Last Glacial Maximum. Nevertheless, to detect and understand processes which determine diversity patterns at high altitude mountains in the tropics, more attention, longer sampling periods, and deeper taxonomic knowledge of the species and their phylogenetic relationships covering the whole Neotropics is needed. Matthias Rös studies diversity patterns in natural and human-modified landscapes, looking for biodiversity-friendly land-use patterns. Oaxaca seems to have plenty of these biodiversity-friendly land-use patterns in its mountain landscapes, despite or even because of a human-induced modification history dating millennia. Our research of describing new species is the baseline for further investigations. How can we protect the outstanding biodiversity under scenarios of climate change and land-use intensification? Oaxaca might suggest very interesting answers to many questions related to this topic. Oaxaca and its mountains still have many secrets to unfold, and we want to explore and reveal them.

 

Arriaga-Jiménez, A., Escobar-Hernández, F., Rös, M., & Kohlmann, B. (2020). The establishment of the Onthophagus anthracinus (Coleoptera: Scarabaeidae) species complex and the description of a new species. The Canadian Entomologist, 152:1-17. https://doi:10.4039/tce.2019.62. (Paper made available to read for FREE until March 24, 2020 in cooperation with Cambridge University Press)

 

Related research to dung beetles in high mountains:

Kohlmann B., Arriaga-Jiménez, A., Rös, M. 2018. Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae). ZooKeys743:67-93. https://zookeys.pensoft.net/articles.php?id=23029

Arriaga-Jiménez, A., Rös, M. & Halffter.G. 2018. High variability of dung beetle diversity patterns at four mountains of the trans-Mexican volcanic belt. PeerJ 6:e4468. https://doi.org/10.7717/peerj.4468

Kohlmann, B., Arriaga-Jimenez, A., & Rös, M. 2018. An unusual new species of Canthidium (Coleoptera: Scarabaeidae: Scarabaeinae) from Oaxaca, Mexico. Zootaxa 4378 (2): 273–278. https://doi.org/10.11646/zootaxa.4378.2.7

, , ,

Ancient spiders from an ancient forest

20170528-IMG_20170528_175611_DRO

Ever wish you could travel back through time and see a west coast Vancouver Island rainforest before industrial logging? To see huge old trees, intact soils and life in a climax ecosystem? You do not have to invent a time machine, you only need to travel about an hour out of Port Renfrew to the spectacular Walbran Valley.

20170528-IMG_20170528_181230_DRO

As part of an effort to document the biodiversity of the valley, I traveled with fellow arachnologists Claudia Copley, Darren Copley, Zoe Lindo, and Catherine Scott, along with birders, mycologists, lichenologists and assorted volunteers to spend a day among the giant trees. We were there at the invitation of the Friends of Carmanah-Walbran to lend our expertise to the effort of catloguing the biodiversity of this beautiful, yet still at-risk west coast habitat.

We arrived at the somewhat storied “Bridge to Nowhere”, where in 1991 environmental protesters confronted the logging companies, the RCMP and the government of British Columbia, holding the line against industrial exploitation of a rare ecosystem. What the activists were asking for seems modest: Can’t we have just this one watershed, among all the others on Vancouver Island, be preserved and protected from the clearcutting and degradation that is the fate of every other valley on the Island?

20170528-IMG_00212. Pacheedaht elder Bill Jones walks across the Bridge to Nowhere

While the Friends of Carmanah-Walbran took the other participants deep into the woods on hikes, we arachnologists ventured only short distances into the woods, as our slow and careful sifting through the soil and beating of the bushes is certainly not a thrill ride for everyone. For us, however, it was thrilling, as within 30 minutes of arrival on site, we had found a beautiful and seemingly dense population of Hexura picea, a relative of tarantulas.

20170528-IMG_00803. Hexura picea, a tarantula relative, brought out of its underground silk tunnel complex for a photo shoot.

These little, pretty, but nondescript spiders live in small silk tunnel complexes among the soil and rocks of the forest floor. Each tunnel has a main entrance lined with silk, and several other openings which may facilitate rapid escape or offer alternate exits at which to snare prey. Being members of the suborder Mygalomorphae, they are indeed tarantula relatives, a group of spiders that closely resemble ancient spiders. Many mygalomorphs retain traces of segmentation on their abdomens, unlike the more modern araneomorph spiders. In the Mecicobothriidae (to which Hexura belongs) the terminal spinneret segments bear “pseudosegmentation”

IMG_0366

The section of forest we found this spider in was a real “tangled bank”, in fact the scree slope associated with Walbran creek and a small tributary, which has since been covered with a layer of soil and a stand of hardy trees.

20170528-IMG_00574. Erosion is a gentler process in a forested valley, with trees holding on to what would be a talus slope higher in the mountains. The soils beneath these trees support an extensive food web.

Finding these spiders in the Walbran was not unexpected, as they had previously been found in the Carmanah Valley and at Avatar Grove, but their presence on Vancouver Island is somewhat puzzling, as they represent the only known Canadian population, and are seemingly not present on the BC mainland.

Given the dense population in the Walbran, the valley would be an wonderful place to study their behaviour, which so far is undocumented. We would presume that much of the activity of these spiders takes place at night, although Catherine was able to lure one out of its burrow by tickling the silken doormat with a twig.

20170528-IMG_01115. Hexura picea emerges from its silken tunnel and onto its “doormat” to “kill” a vibrating cedar twig.

The litter sampling we conducted will surely yield many more species, although we have to wait until the Berlese funnels have extracted all of the arthropods. The work of sampling and cataloguing biodiversity takes time, and is not totally congruent with the rapid “bioblitz” ethos.

If you are ever in BC, and want a trip back in time (never mind our politics), please do not hesitate to come out to the Walbran Valley. You may just discover something amazing.

20170528-IMG_02486. Darren and Claudia picking up pan traps beside the Malaise flight-intercept trap.

 

 

 

 

, , , , ,

Missed Mandate, Missed Biology: The ongoing “Mother Canada” debacle in Cape Breton Highlands National Park

Opinion Piece – M. Alex Smith, Department of Integrative Biology, University of Guelph (salex@uoguelph.ca; @Alex_Smith_Ants; www.malexsmith.weebly.com)

—-

Like many Canadians, I have been hearing more and more about the so-called “Mother Canada” development in Cape Breton Highlands National Park (CBHNP). Proposed by a combination of private funding in partnership with the federal government, this enormous 10-storey memorial is meant to “… be a place for remembrance and gratitude” to Canadians who have “fallen as a result of war and conflict”. Parks Canada has expressed direct support for this monument through actual monetary donations. The erection of such a memorial within a Canadian National Park has garnered much recent interest in the Canadian and international press.

Beyond any aesthetic concerns people may have about the specific plans, in my opinion, there are two critical problems with this monument. The first was pointed out in a Globe and Mail editorial of June 24 2015: it is redundant. Every town and city in Canada already has a memorial to those who have served and sacrificed. My second objection is a combined biological and sociological one. It concerns the location of a private funded monument within a Canadian National Park, where it appears very unclear what the ramifications of that action will be on the fauna in and around the proposed site. The mandate of Parks Canada is elegantly expressed in its charter, “To protect, as a first priority, the natural and cultural heritage of our special places and ensure that they remain healthy and whole” while fostering “public understanding, appreciation and enjoyment in ways that ensure the ecological and commemorative integrity of these places for present and future generations”. Indeed, 26 former senior Parks Canada managers wrote an open letter to the Minister of Environment Leona Aglukkaq detailing their objections and that such a plan, “is in violation of the site’s Wilderness Zone designation as detailed in the Management Plan for the Park”.

Beyond the effects of the actual physical construction on the park environment, the monument will potentially increase tourist traffic to the area. How will these changes affect the biota (both animal and plant) of the immediate area? Exactly how well known is that fauna? How was the effect on the sites and the adjacent park environment determined?

A detailed impact analysis was completed by Stantec Consulting Limited who concluded that the effects of the development are, “generally predicted to be negligible to moderate in magnitude”. Conclusions regarding the effect of the construction and development on the “wildlife” of CBHNP were based on a single terrestrial field survey of the locality and a consultation of a CBHNP sightings database. (Stantec is actually listed as a Partner and Supporter of the development). In the Stantec impact analysis, “wildlife” is exclusively mammals and birds. As an ecologist whose professional and personal life is replete with instances of being overwhelmed and delighted by the diversity of arthropods living coincidentally (and cryptically) with their better-studied vertebrate relatives, this raised some concerns.

So what can I offer? Well in 2009, I spent a wonderful time collecting arthropods in CBHNP as part of the BioBus program out of the Biodiversity Institute of Ontario at the University of Guelph. In fact, four colleagues and I spent a night collecting insects at a site only 3 km away from the proposed development (Black Brook and the nearby Jack Pine Trail). The Jack Pine trail was particularly beautiful! The trail goes through a forest of Jack Pine that is more than 200km away from the rest of its range and has survived fire and spruce budworm infestation. At any rate, since all the data is publicly available online (dx.doi.org/10.5883/DS-ASCBHNP), I thought this would be an opportune time to explore those records in light of the planned “Mother Canada” development.

 

Figure 1: A high resolution GigaPan panorama taken at the Black Brook collection site (http://gigapan.com/gigapans/29312).

Figure 1: A high resolution GigaPan panorama taken at the Black Brook collection site (http://gigapan.com/gigapans/29312).

 

Figure 2: The collection team earlier in the trip in Terra Nova National Park Newfoundland.

Figure 2: The collection team earlier in the trip in Terra Nova National Park Newfoundland.

It was a beautiful night in 2009 (Jul-21) at Black Brook where we collected arthropods using two common methods (UV light (which means lots of moths!) and free-hand active search using insect nets). That night, in about four hours of collecting, we came away with 363 specimens from nearly 200 species (191 named and provisional species based on their DNA barcodes). To put this number in context, CBHNP has 200 species of bird – a total nearly matched for arthropods by our single nights work at one location! This diversity is only a small fraction of the diversity of arthropods currently protected by CBHNP. Via these DNA barcodes, (public on BOLD (www.barcodinglife.org, dx.doi.org/10.5883/DS-ASCBHNP) we can compare them to the > 4 million DNA barcode records representing >400,000 species worldwide on this database.

What we find from this comparison is that some of these species may be exceedingly rare. Despite concentrated collections in this and other National Parks before and since this night* there are four species which have been found only once out of these millions of records. While this diversity is currently protected by Parks Canada, it is within 3 km of the proposed “Mother Canada” development. It is unclear how the changes in traffic and construction from the development will affect this protected diversity.

Why bring this up now? What use is a rapid analysis of a single night’s collections? I decided to bring it up to call attention to numerous small and cryptic species in and around the location of the proposed development about which we know very little. Going ahead with an enormous private development within a National Park is a mistake that flies in the face of the mandate of Parks Canada – and does so without good evidence that it would not have negative effects on the diversity of animals that it was created to protect.

 

Figure 3: This neighbor-joining tree is a graphical representation of the diversity of nearly 200 species of arthropods collected at Black Brook in July 2009. The taxa are colour coded and are followed by the number of specimens we caught.

Figure 3: This neighbor-joining tree is a graphical representation of the diversity of nearly 200 species of arthropods collected at Black Brook in July 2009. The taxa are colour coded and are followed by the number of specimens we caught.

John Barber (a freelance journalist from Toronto) closed his recent article in the Guardian newspaper with a marvelous quote from Valerie Bird, a WWII veteran and resident of Cape Breton, “It is vulgar and ostentatious,” she said. “It certainly doesn’t belong in a national park, and I don’t think its going to do a darn thing for veterans.” “I think the idea of this horrible thing offends veterans,” she added. “I find it difficult to find words. This is a monstrosity.”

Not simply a monstrosity – but one contrary to of the principle mandate of Parks Canada, “to protect, as a first priority, the natural and cultural heritage of our special places and ensure that they remain healthy and whole”. Ultimately, this is the essence of the problem. This issue is more than a simple discussion regarding the aesthetics of a >$25 million, >25-metre tall conglomeration of private and corporate citizens (in apparent partnership with our federal government). If a private group wants to erect a memorial on private grounds and can raise the money for their monument – it is certainly their prerogative. This is a critical discussion about the mandate of Parks Canada and specifically how well they protect the natural heritage resident within that Park.

To place this monument in a National Park is not the right of any private group. To consider placing such a monument in a National Park without careful consideration of the most diverse Park residents (insects, spiders and their kin) is not simply poor planning; it’s poor management and should be stopped.

* -Since that evening in 2009, the BioBus has continued to collaborate with Parks Canada in Cape Breton Highlands National Park and now even more is known about the vast diversity of small and important insects from other areas within this National Park. Collections of arthropods have now been made for 3000 species! For more information about those collections visit the reports section at www.biobus.ca. The author has no current association with the BioBus program. All specimens analysed here are publically available via the public data portal of the Barcode of Life Data System (dx.doi.org/10.5883/DS-ASCBHNP).

Useful websites:

Thanks to Morgan Jackson for helpful thoughts on an earlier draft of this post.
Figure 4 – Shareable infographic outlining information & data presented in this article. Please feel free to circulate.

Figure 4 – Shareable infographic outlining information & data presented in this article. Please feel free to circulate.

,

Recreational boating affects dragonflies

—-By Aaron Hall—-

A typical adult dragonfly. Note the spiked legs, which are held in a basket shape to help catch prey while flying.

A typical adult damselfly. Note the spiked legs, which are held in a basket shape to help catch prey while flying.

Dragonflies are charismatic insects, and most of us can probably remember chasing them or watching their acrobatic flights when we were children. But what most of us didn’t realize when we were kids, is that dragonflies spend the majority of their lives as toothy, alien-looking predators living underwater before they become adults. Depending on the species, they can live in the water for several weeks up to several years.

A typical larval dragonfly, which feeds on other aquatic animals - and even other dragonflies!

A typical larval dragonfly, which feeds on other aquatic animals – and even other dragonflies!

By living part of their lives in water, and part on land/in the air, dragonflies represent an interesting conservation challenge. Historically, conservation science has focused on single habitats, such as lakes, streams, forests or grasslands. Little attention has focused on incorporating multiple habitat types, such as those required by dragonflies, into conservation, potentially leaving species like dragonflies in danger.

In the Waubaushene area of Georgian Bay (Lake Huron), recreational boating is very common. These boats create waves that can dislodge both adult and larval dragonflies, affecting their ability to find food and avoid predators. The overall number of boats, the speed of these boats, and how close they are to coastal wetlands are the most important factors that determine how impactful boat-generated waves are on dragonflies. My colleagues and I at the University of Toronto investigated how much influence these recreational boats have, relative to more natural processes, on dragonfly communities in Georgian Bay.

A Google Earth image of an area in Georgian Bay. Note the many waves created by boats as they travel through this region.

A Google Earth image of an area in Georgian Bay. Note the many waves created by boats as they travel through this region.

Taking the lead on this project, I counted dragonflies from 17 islands in Waubaushene. The coastal wetlands around these islands are inhabited by dragonflies. The islands studied in this project were selected to represent a range of influence from boats in the area, determined by their distance and orientation to marked boating channels and area marinas.

Aaron Hall counting adult dragonflies at one of the islands in Waubaushene.

Aaron Hall counting adult dragonflies at one of the islands in Waubaushene.

The results show that boats do have an influence on dragonfly communities, providing a link between recreational boating and dragonfly communities. This research provides important insights that can be applied to the protection and conservation of dragonflies, and suggests that some very simple changes in boater behaviour could have big implications. For example, if boats travel slower or further away from dragonfly habitats, they would have less impact. These two factors might be simple to change. In areas where boats mostly stay within marked boating channels, if these channels were moved or adjusted so they are as far away from dragonfly habitats as possible, impacts would be minimized. Additionally, speed limits could be set in these channels to reduce the size of waves created by boats. These simple measures could have a positive impact on dragonflies, which are a critical component in the aquatic and terrestrial foodwebs of this region.

Want to know more? This research is published in the journal Insect Conservation and Diversity. You can also follow me on Twitter @aarohall.

,

Green and red for the holidays

pentatomidae 1

This season is full of green and red decorations, and exhortations about family, so I thought I would share this lovely family portrait of some 1st instar stink bugs exemplifying the togetherness of the holiday season. I found this family group in Fort Pierce, Florida on the underside of a Brazilian Peppertree leaf. Schinus terebinthefolius is an invasive plant brought in as an ornamental for its beautiful red berries and evergreen leaves (and used in Christmas decorations!). Not surprisingly, entomologists are on the lookout for a insect-based solution for peppertree control!

pentatomidae 2

, , ,

Canadian amber reveals bizarre ants from the Cretaceous – Editor’s Pick for TCE 145(4)

By Dr. Chris Buddle, McGill University & Editor of The Canadian Entomologist

———-

It’s with great pleasure that I announce my pick for the latest issue of The Canadian Entomologist.  Ryan McKellar and colleagues wrote a paper on a new trap-jawed ant from Canadian late Cretaceous amber (freely available during September).  As they write in the Abstract, the new species “….expands the distribution of the bizarre, exclusively Cretaceous, trap-jawed Haidomyrmecini beyond their previous records…”. They truly are bizarre! Facial structures right out of a sci-fi movie!  When reading the paper, I was also surprised that the fossil record for the Formicidae is sparse during the Cretaceous.

Haidoterminus cippus. Figure 1 from McKellar et al. 2013

I asked the lead author a few questions about this work, and am pleased to share the responses with you. It’s truly exciting research, and I am thrilled that the pages of TCE include systematics from amber. This work stirs the imagination, and takes us all back in time.

What inspired this work?

My interest in the Canadian amber assemblage really began when Brian Chatterton (then my M.Sc. supervisor) showed me some of the slides that he had borrowed from the Royal Tyrrell Museum of Paleontology in order to write a book on Canadian palaeontology. The sample set contained insects with bizarre adaptations for life at low Reynolds numbers, and obvious ecological associations, spurring an interest that ultimately led to a research in parasitic microhymenoptera. Michael Engel subsequently introduced me to a much wider array of taxa, and we continue to explore the Canadian assemblage together and with the help of colleagues.

What do you hope will be the lasting impact of this paper?

New records, such as this trap-jawed ant, help to flesh out our picture of the amber-producing forest and its inhabitants. I hope that a comprehensive account of this assemblage will eventually provide insights into terrestrial conditions that are unavailable from other fossil types, and that this will shed some light on changes in diversity and conditions leading up to the end-Cretaceous mass extinction.

Where will your next line of research on this topic take you?  

With any luck, we will be able to complete our coverage of Hymenoptera in Canadian amber soon, and make more of a concerted effort to cover other insect orders and some of the ecological associations found within the deposit. Grassy Lake amber still has a lot to offer, but it is only one of western Canada’s many amber deposits. As a larger-scale project, we are currently part of a team examining the numerous fragile ambers associated with coals in the region. The goal of this research is to create an amber-based record of forest types and inhabitants that spans more than 10 million years within the Late Cretaceous and Paleocene.

Can you share any interesting anecdotes from this research?

Surface-collecting amber can be quite difficult, because unpolished Canadian amber typically has a matte orange-brown colour, and is often covered with a carbon film or weathering crust. If there is no fractured surface visible and the specimen is not translucent, it can be quite difficult to distinguish from the surrounding coal or shale. Furthermore, there is such a range of shapes and sizes that some of the smaller amber droplets are easily confused with modern seeds. One of the quickest ways to see if you are dealing with amber is to wet the specimen and look for amber’s characteristic lustre, or tap the specimen on your teeth (amber feels like plastic compared to most suspect rocks). Naturally, I have licked quite a few samples in the course of my collecting, and have lost a lot of my appreciation for rabbits.

A selection of amber from Grassy Lake. Photo courtesy Brian Chatterton

A selection of amber from Grassy Lake. Photo courtesy Brian Chatterton

Thanks to Cambridge Journals Online for making this month’s Editor’s Pick Freely Available for the month of September!

This post is a regular series highlighting great papers from the pages of the Canadian Entomologist. 
McKellar R.C., Glasier J.R.N. & Engel M.S. (2013). A new trap-jawed ant (Hymenoptera: Formicidae: Haidomyrmecini) from Canadian Late Cretaceous amber, The Canadian Entomologist, 145 (04) 454-465. DOI: