Insect Biodiversity and Dead Wood: Proceedings of a Symposium for the 22Nd International Congress of Entomology
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United States Department of Insect Biodiversity and Dead Wood: Agriculture Proceedings of a Symposium for Forest Service the 22nd International Congress of Entomology Southern Research Station Brisbane, Australia General Technical August 15–21, 2004 Report SRS–93 Grove, Simon J.; Hanula, James L., eds. 2006. Insect biodiversity and dead wood: proceedings of a symposium for the 22nd International Congress of Entomology. Gen. Tech. Rep. SRS–93. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 109 p. In August 2004, the city of Brisbane, Australia, was host to one of the largest recent gatherings of the world’s entomologists. The 22nd International Congress of Entomology featured a multitude of symposia covering a wide range of entomology-related topics. This general technical report is based on papers presented on one such symposium, “Insect Biodiversity and Dead Wood.” It features contributions from around the world. Dead wood is anything but dead. It is the lifeblood of an intricate web of life in which insects feature prominently. The papers presented here consider both the basic ecology and evolutionary history of saproxylic (dead-wood dependent) insects and how such insects can be affected by management of the forests where most species live. Past management has not always been beneficial to saproxylic insects and in some parts of the world has harmed them greatly. With continued quality research, however, there is some prospect that future land management will be more accommodating to saproxylic insects. Keywords: Arthropod diversity, coarse woody debris, fossil insects, funnel web spider, invertebrate conservation, New Forestry, saproxylic insects. Preface In August 2004, the city of Brisbane, Australia, was host to one of the largest recent gatherings of the world’s entomologists. Several thousand delegates attended the 22nd International Congress of Entomology, which featured a multitude of symposia that together covered a wide range of entomology-related topics. This special General Technical Report is based on papers presented in a symposium entitled “Insect Biodiversity and Dead Wood.” It features contributions by scientists from around the world, and these contributions clearly illustrate our growing understanding of the entomological importance of dead wood. Dead wood is anything but dead. It is the lifeblood of an intricate web of life in which insects figure prominently. The papers presented here consider the basic ecology and evolutionary history of saproxylic (dead-wood dependent) insects, and show how these species can be affected by human management of the forests where most of them live. Past management has not always been kind to saproxylic insects, and in some parts of the world this has harmed the fauna significantly. However, there is reason to think that continued research may in the future give land managers the technical tools to accommodate saproxylic insects. Whether societies will adopt these friendlier techniques remains to be seen, but we hope the information presented in this volume will encourage them to do so. Simon Grove, Co-convenor Forestry Tasmania Jim Hanula, Co-convenor USDA Forest Service Southern Research Station ii Contents Page Page Evolution of Saproxylic and Mycophagous The Role of Dead Wood in Maintaining Arthropod Coleoptera in New Zealand . 1 Diversity on the Forest Floor . 57 Richard A.B. Leschen James L. Hanula, Scott Horn, and Dale D. Wade Aquatic Wood — An Insect Perspective . 9 Peter S. Cranston and Brendan McKie The Fine Scale Physical Attributes of Coarse Woody Debris and Effects of Surrounding Stand The Use and Application of Phylogeography Structure on its Utilization by Ants (Hymenoptera: for Invertebrate Conservation Research Formicidae) in British Columbia, Canada . 67 and Planning . 15 Robert J. Higgins and B. Staffan Lindgren Ryan C. Garrick, Chester J. Sands, and Paul Sunnucks Saproxylic Beetles in a Swedish Boreal Forest Landscape Managed According to ‘New Forestry’ . 75 Phylogeography of Two Australian Species of Stig Larsson, Barbara Ekbom, Funnel Web Spider (Araneae: Mygalomorphae: L. Martin Schroeder, and Hexathelidae) in Tallaganda State Forest, Melodie A. McGeoch New South Wales . 23 Amber S. Beavis and David M. Rowell Maintaining Saproxylic Insects in Canada’s Extensively Managed Boreal Forests: A Review . 83 What Can Forest Managers Learn from Research on David W. Langor, John R. Spence, Fossil Insects? Linking Forest Ecological History, H.E. James Hammond, Joshua Jacobs, Biodiversity and Management . 30 and Tyler P. Cobb Nicki J. Whitehouse A Research Agenda for Insects and Dead Wood . 98 Brown Rot in Inner Heartwood: Why Large Logs Simon J. Grove Support Characteristic Saproxylic Beetle Assemblages of Conservation Concern . 42 Index of Authors.............................. 109 Marie Yee, Simon J. Grove, Alastair MM. Richardson, and Caroline L. Mohammed iii iv Insect Biodiversity and Dead Wood: Proceedings of a Symposium for the 22nd International Congress of Entomology Editors: Simon J. Grove, Conservation Biologist Forestry Tasmania Southern Research Station Hobart, Tasmania 7001, Australia James L. Hanula, Research Entomologist U.S. Department of Agriculture Forest Service Southern Research Station Athens, GA 30602-2044 i EVOLUTION OF SAPROXYLIC AND MYCOPHAGOUS COLEOPTERA IN NEW ZEALAND Richard A.B. Leschen1 Abstract—Beetles are an old holometabolous group dating back to the early Permian and associated with sediments contain- ing conifers, ginkgos, and cycads. To determine the antiquity of dead wood beetles the evolution of gondwanan saproxylic and mycophagous beetles was examined in the context of available phylogenies that include New Zealand taxa. Phylogenetic position and branch lengths showed that 50 percent of the New Zealand fauna is basal and may represent old lineages dating to around 82 million years when New Zealand separated from Gondwana. Meanwhile, 60 percent of the New Zealand taxa have relatively long branches relative to overseas outgroups and this phenomenon may have resulted from the adapta- tion of these groups to resource shifts in changing forest communities. The resource shift hypothesis predicts that polyphagy will be widespread in mycophagous and phytophagous insects. Podocarp-broadleaf forest associations are more primitive than associations with Nothofagus based on species level phylogenies and forest associations of long-branched taxa. INTRODUCTION These features are thought to be characters that form part of Though my work as a systematist is geared to develop classi- an adaptive complex that led to the extraordinary number of fications, build keys, and provide monographs that collate beetle species that exists today. There are four suborders, natural history and taxonomic data, most of the contributors Archostemata, Adephaga, Polyphaga, and Myxophaga. The and readers of this volume may be chiefly interested in the first three have many saproxylic species, and the fourth may ecology of dead wood insects. I believe that the time is ripe do so (Löbl 995). The relationships among the four suborders for ecologists and systematists to build a common research are controversial (see reviews in Lawrence 999, Kukalová- program and it may be useful to introduce some of the readers Peck and Lawrence 2004, Leschen and Beutel 2004) with to what can happen in the field of ecological phylogenetics. many studies supporting Archostemata as sister taxon to the Moreover, it may help some ecologists to appreciate that other suborders (compare Beutel and Hass 2000 to Kukalová- taxonomists are not just folks who are interested in building Peck and Lawrence 993, 2004). The Archostemata have a collections, eager to obtain a few specimens in exchange for number of primitive characteristics (plesiomorphies), and the a service identification, but are scientists keen to place their common belief that they are a primitive group in Coleoptera knowledge about natural history in a broader scientific context. is matched by the fossil record. The first true beetles are Archostemata recorded from the early Permian (265 my) and The insect order Coleoptera is one of the most important taxa placed in the family Tshekardocoleidae, a short-lived family associated with dead wood (Grove 2002). It is the largest ranging from the early Permian to early Triassic (Ponomarenko group of described organisms and beetles occupy almost 969, 2002). But the Archostemata, as defined by Pono- every niche available in dead wood (and outside of it). Because merenko (2002), may be paraphyletic (see Kukalová-Peck of their immense diversity, microhabitat associations and and Lawrence 2004) and some of its members, including behaviors, beetles are popular subjects for ecological studies. Tshekardocoleidae, may be placed separately in the order But what about their evolution? How did modern saproxylic Protocoleoptera. communities come into existence? In this essay I report on phylogenetic work in progress and discuss the evolution of Part of the phylogenetic debate surrounding the ordinal saproxylic beetles at deeper evolutionary time scales. First, I phylogeny of beetles is the presence of many derived charac- briefly review data that indicate that the earliest beetles may teristics in Archostemata (Lawrence 999), a group that has have been saproxylic. Then I focus on two phylogenetic ques- undergone considerable radiation (even though it is rather tions about saproxylic beetles in New Zealand: () are New depauperate at present). The