Insect biodiversity and dead wood : proceedings of a symposium for the 22nd International Congress of Entomology

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Cover photo: Staphylinids in Tasmanian Forest Insect Collection
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Insect Biodiversity and Dead Wood:
Proceedings of a Symposium for the
22" 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
posBA
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LC
CS ATA LCO GIi l G_PREP
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
22" 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
eBi e
Contents
Page Page
aS heata ka UNTER ES 4 The Role of Dead Wood in Maintaining Arthropod
Pa Re teech Sie eka OunceF is Diversity on the Forest Floor..................... 57
Eis su James L. Hanula, Scott Horn, and
Dale D. Wade
Aquatic Wood — An Insect Perspective............ 9
tee cnsion and Brendam 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
UNG ECTG T be vO) oe oe On ee ee PS 15
Robert J. Higgins and B. Staffan Lindgren
Ryan C. Garrick, Chester J. Sands,
SUES UanUCKS 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
NO WISOULLVVAICS;9.7 cnr de ae wns WA ed Ea aly 5 Fas}
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.C obb
Nicki J. Whitehouse
A Research Agenda for Insects and Dead Wood .... 98
Brown Rot in Inner Heartwood: Why Large Logs SmeGravs
Support Characteristic Saproxylic Beetle
Assemblages of Conservation Concern ........... 42
IndexofAULNOIS tee tec ee aie sco eee ernie 109
Marie Yee, Simon J. Grove,
Alastair MM. Richardson, and
Caroline L. Mohammed
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EVOLUTION OF SAPROXYLIC AND MYCOPHAGOUS
COLEOPTERA IN NEW ZEALAND
Richard A.B. Leschen'
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. | believe that the time is ripe do so (L6bl 1995). The relationships among the four suborders
for ecologists and systematists to build a common research are controversial (See reviews in Lawrence 1999, Kukalova-
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 Kukalova-
taxonomists are not just folks who are interested in building Peck and Lawrence 1993, 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 1969, 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 Kukalova-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 | 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, | phylogeny of beetles is the presence of many derived charac-
briefly review data that indicate that the earliest beetles may teristics in Archostemata (Lawrence 1999), a group that has
have been saproxylic. Then | focus on two phylogenetic ques- undergone considerable radiation (even though it is rather
tions about saproxylic beetles in New Zealand: (1) are New depauperate at present). The suborder includes rostrate forms
Zealand taxa basal with respect to other members of their from the Triassic (Obrieniidae was described by Zherikhin and
clade, and (2) do they have longer branches compared to Gratshev (1993) as belonging among the weevils (Polyphaga),
their sister taxa located in other areas? Both questions relate but were later transferred to Archostemata (Zherikhin 2002)),
to the uniqueness of beetle faunas in the geologically diverse and unusual life histories, including thelytokous development
and geographically isolated archipelago of New Zealand. with viviparous, larviform females in the family Micromalthidae
(Pollock and Normark 2002).
ANCIENT SAPROXYLIC BEETLES Nevertheless, the antiquity of Archostemata is unmatched by
Beetles are holometabolous insects defined by a suite of any other beetle group, with extant genera dating back to the
adult synapomorphies that include hardened forewings or early Jurassic (Omma Newman and Tetraphalerus Water-
elytra that enclose the abdomen and flight wings while resting. house; Zherikhin 2002). At the Permian dawn of Coleoptera,
1 Richard A.B. Leschen, New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand.
Citation for proceedings: Grove, Simon J.; Hanula, James L., eds. 2006. Insect biodiversity and dead wood: proceedings of a symposium for the
224 International Congress of Entomology. Gen. Tech. Rep. SRS-93. Asheville, NC: U.S. Department of Agriculture Forest Service, Southern
Research Station. 109 p.
conifers were on the rise while the first ginkgos and cycads exists only during periods of high sea-levels. During periods
emerged, and so Ponomorenko (1969) and others (Crowson of glaciation and low sealevels the islands are linked. The
1975, Kirejtshuk 2003) speculate that the early Archostemata North Island landscape, by contrast, is a result of the offshore
had xylomycetophagous larvae, based on modern feeding subduction that creates north-south trending axial ranges
habits. Though there are clear indications of ancient wood that absorb plate compression, a volcanic arc, with lines of
boring in fossil wood (Weaver and others 1997) that may have active volcanoes and a backarc region that is being thinned
been caused by beetles, trace and fecal fossils are not affirma- and is subsiding. The elevations of North Island ranges are
tive (Labandeira 1998), and differentiating between species lower than the Southern Alps. The north terminates in a long
that feed on decayed or live wood (Zherikhin 2002) is proble- peninsula pointing towards New Caledonia, while the broad
matic. Meanwhile, the most primitive groups of extant beetles middle portion is dominated by East and West Capes, making
have mouthparts that are associated with feeding on materials the North Island appearing spindle-shaped.
such as microfungi and algae (Crowson 1981, Lawrence
1989), and this type of microphagy (Betz and others 2003, The geological history of New Zealand, and its contrasting
Leschen 1993) may have been a primitive feeding mecha- landforms, make it an ideal model system for biogeographic
nism. The presence of microphagous feeding mechanisms studies, so it is no surprise that the discipline of panbiogeog-
suggests that the diets of ancient beetles were not strictly raphy has its most fervent supporters here (Craw and others
wood feeding and beetles may have occurred in any decaying 1999, Hull 1988). New Zealand is home to a unique fauna
habitats that promote fungal growth (Personal communication. (like the panbiogeographers). Ratite birds have undergone
2005. John F. Lawrence, Systematist, CSIRO Entomology, major extinction in New Zealand, and the once diverse group
GPO Box 1700, Canberra, ACT, 2601, Australia). Because of is now reduced to three species of kiwi that at are the delight
the rise of conifers with their cork-cambium layer that is easily of conservationists, lay people, and scientists. Because of
separated from heartwood after death, it is quite likely that the presence of these ratite birds, and of tuatara, tailed-frogs
the subcortical situation may have been one of the micro- (Leiopelmatidae), and primitive plants like kauri (Agathis
habitats that ancient beetles first inhabited (Crowson 1984, australis (D. Don) Lindl. (Araucariaceae)) and whisk-ferns,
Ponomorenko 2002). most biologists consider New Zealand a refuge for ancient
lineages that evolved prior to the fragmentation of Gondwana.
Likewise, many of the beetles may be relictual and should be
ANCIENT PHYLOGENETIC PATTERNS IN
of interest from an evolutionary and ecological (conservation)
NEW ZEALAND
perspective.
One way to gain an appreciation of the antiquity of saproxylic
beetles is to study their history in New Zealand, an ancient
So, is the dead wood beetle fauna relictual? One way to test
land with an interesting mix of old endemic species and those
this question is to determine the phylogenetic position of New
that have colonized more recently. There are approximately
Zealand beetles in reconstructed phylogenies (cladograms)
5000 described New Zealand beetle species (Klimaszewski
containing taxa from other areas. Relatively basal (primitive)
and Watt 1997), though there may be up to 10,000 species in
taxa will be located near the root of trees, while derived taxa
total, which are placed into 86 families (Leschen and others
will be located in higher branches. In figure 1 clade AB is
2003): 58 of the families have species associated with dead
located at the base and is considered primitive relative to,
wood. The total numbers of species rival the number for
say, taxon E, which is more derived. Also, do New Zealand
Britain (just over 4000 species), a country of similar area. But
New Zealand’s location and history are strikingly different
from Britain’s (and its fauna is too!).
New Zealand is an island archipelago in the southwestern
Pacific, chiefly comprised of two main islands that are pre-
sently positioned astride the Australian and Pacific plates.
Active tectonism has produced a diverse and complex land-
scape (Suggate and others 1978). Isolation from Gondwana
began around 82 million years ago when New Zealand was
separated from Australia by the development of the Tasman
Basin (Cooper and Millener 1993) and the rifting of the New
Zealand landmass to its present eastward location. By the
Oligocene period (c. 35 my) most of New Zealand was sub-
merged. Later, plate collision resulted in a mountain building
phase which originated in the Miocene and continues to pre-
sent day. The result is that the South Island is dominated by
the north-south trending Southern Alps, which rise to elevations
of nearly 4000 m and extend around 500 km. To the east in
the rain shadow of the Southern Alps, the landscape consists
of lowlands and basins. The southwest end of the South Island
contains extensive fjiordlands while the northeast of the South
Island terminates in sounds. The South Island is separated
from the North Island by the narrow Cook Strait which was Figure 1—Five-taxon area-cladogram showing distribution of taxa
A-F. The bars across each branch indicate character changes
formed around 0.5 my ago (Lewis and Carter 1994) and
(OZ = Australia, CH = Chile, NZ = New Zealand).