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Evolution and Ecology of Termite Nesting Behavior and Its Impact on Disease Susceptibility

A dissertation presented by
Marielle Aimée Postava-Davignon

to
The Department of Biology

In partial fulfillment of the requirements for the degree of
Doctor of Philosophy

in the field of
Biology

Northeastern University Boston, Massachusetts
April, 2010
2

Evolution and Ecology of Termite Nesting Behavior and Its Impact on Disease Susceptibility

by
Marielle Aimée Postava-Davignon

ABSTRACT OF DISSERTATION
Submitted in partial fulfillment of the requirements for the degree of Doctor of
Philosophy in Biology in the Graduate School of Arts and Sciences of Northeastern
University, April, 2010
3

Abstract

Termites construct nests that are often structurally species-specific. They exhibit a high diversity of nest structures, but their nest evolution is largely unknown. Current hypotheses for the factors that influenced nest evolution include adaptations that improved nest thermoregulation, defense against predators, and competition for limited nest sites. Studies have shown a lower prevalence of pathogens and parasites in arboreal nesting animal species compared to ground nesters. Nest building behavior is plastic and can adapt to changing environments. As termites can detect and avoid pathogens, I hypothesized that the evolution of arboreal termite nests was an adaptation to avoid infection. To test this, bacteria and fungi from nest cores, trails, and surrounding soils of the arboreal nesting Nasutitermes acajutlae were cultured. Abiotic factors such as temperature, relative humidity, and light were measured to elucidate how they influenced the interactions between termites and microbes. Fungi associated with N. acajutlae were identified to determine the potential pathogenic pressures these termites encounter in their nest as compared to the external environment. To determine the effect of nest structure on survival, termites representing the one-piece (Zootermopsis angusticollis),

intermediate (Reticulitermes flavipes) and separate (Nasutitermes corniger) type nests

were exposed to the fungal entomopathogen Metarhizium anisopliae, and the termites’ survival tracked over a 20-day period in five different artificial nest architectures. A protected nest environment and the benefits of socially mediated immunity within those nests have also been implicated in promoting termite eusocial evolution. In order to test whether immune protein production is socially induced in termites, the SDS-PAGE
4protein profiles of naïve Z. angusticollis nymphs, and nestmates directly exposed to M.

anisopliae were examined following social contact. The results presented in this dissertation demonstrate that arboreal termites have lower nest microbial loads and diversity compared to surrounding soils, and that the degree of social contact as influenced by nest architecture can significantly affect termite survival. This research suggests that pathogens played a role in furthering evolution and ecology of termite nesting behavior, and lays the groundwork for future studies in this area.
5
This dissertation is dedicated to my family.
To my parents, who have loved and supported me through all my endeavors. To my sisters, who have given me invaluable guidance, strength, and love. To my brothers-in-law, who watch over and care for me like a true sister.
To my niece, whose laughter and bright smile helped get me through tough times.
I love you all, and could not have done this without you.
6

Acknowledgements

I would like to express my gratitude to the many people without whom this research would not have been possible. My advisor, Rebeca Rosengaus, who introduced me to and led me through the wonderful world of termites, a place to which I otherwise would never have known I truly belonged. My collaborator Claire Fuller for six years of exciting field research, and friendship. My committee members Wendy Smith, Jacqueline Piret and Gwilym Jones for their advice and support. Mark Bulmer for his expertise, and camaraderie. John Stiller and Erica Waddle for their work on fungal identifications. My fellow graduate students Tamara Hartke, Kelley Schultheis, and Lindsey Reichheld for their input and shoulders to lean on. Comfort Chieh and Danny Dijohnson for culturing countless bacterial and fungal samples. Kayla Hamilton and Charlie Ferranti for their work on the Nasutitermes portion of the nest architecture project. Earthwatch SCAP girls 2005-2008 for their hard work in the field on St. John. All other Rosengaus lab members, and graduate students past and present: you are a rare and genuinely fine group of people, and it has been a pleasure working with you.
Thank you to Huddart and Redwood Regional Parks, and the Smithsonian
Tropical Research Institute for termite collections. Kim Seefeld and the R Development Core Team for assistance with statistics. The Northeastern Biology Department and NSF GK-12 program for financial support. This research was funded by an NSF-CAREER grant (DEB 0447316) awarded to Rebeca Rosengaus, an NSF grant (IBN-0116857) awarded to James Traniello and Rebeca Rosengaus, and by the Durfee Foundation through the Earthwatch Student Challenge Awards Program.
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Contents

Abstract

26

Acknowledgements Contents

7

List of Figures List of Tables

9
11 12

  • 1
  • Introduction

1.1 Ancestry and eusocial evolution 1.2 Termites in the fossil record
13 15 16 19 20 22 26 28
1.3 Nest types, classifications, and associated levels of sociality 1.4 Nest building behavior 1.5 Nest evolution 1.6 Selective pressures acting on nest evolution 1.7 Pathogenic pressures and the evolution of eusociality 1.8 Central aims

  • 2
  • Dynamic interactions between the arboreal Caribbean termite

Nasutitermes acajutlae (Holmgren), its associated microbial communities, and the environment

33

2.1 Introduction 2.2 Methods 2.3 Results
34 37 42
8

  • 2.4 Discussion
  • 60

3456
Fungi naturally associated with the arboreal nesting Caribbean termite Nasutitermes acajutlae (Holmgren)

66

3.1 Introduction 3.2 Methods 3.3 Results
67 69 72

  • 73
  • 3.4 Discussion

The effect of nest architecture on termite susceptibility to a fungal pathogen

87

4.1 Introduction 4.2 Methods 4.3 Results
88 95
103

  • 111
  • 4.4 Discussion

Social induction of hemolymph proteins in the dampwood termite Zootermopsis angusticollis (Holmgren)

119

5.1 Introduction 5.2 Methods 5.3 Results
120 123 130

  • 139
  • 5.4 Discussion

Overall discussion and conclusions

144 150

Literature Cited

9

List of Figures

  • 1.1
  • Figure modified from Eggleton and Tayasu (2001) depicting the eight

  • lifeways of termites
  • 30

43 43 44
2.1 2.2 2.3 2.4

Nasutitermes acajutlae arboreal carton nest Collecting Nasutitermes acajutlae in the field

LogTag data loggers Examples of fungi cultured from Hurricane Hole trail material

  • on thiostrepton PDA
  • 49

50 51 52 53 54 55 56 57 58 78
2.5 2.6 2.7 2.8 2.9
Mean log transformed cuticular bacterial CFUs/SA ± SE Mean log transformed substrate bacterial CFUs/g ± SE Mean log transformed cuticular fungal CFUs/SA ± SE Mean log transformed substrate fungal CFUs/g ± SE Correlations between microbial numbers and temperature
2.10 Mean nest (core), ambient (trail) and soil temperatures ± SE 2.11 Example traces from LogTag data loggers 2.12 Correlation between numbers of fungi and amount of light 2.13 Mean nest (core), ambient (trail) and soil % moisture ± SE 3.1 3.2
Examples of fungal genera associated with Nasutitermes acajutlae Fungal genera and number of occurrences in core, trail, and surrounding soil samples of Nasutitermes acajutlae individuals and nests Relative similarities of fungi among Nasutitermes acajutlae nests located in different habitats
80 81
3.3
10

  • 86
  • 3.4

4.1 4.2 4.3

Nodules in a fallen nest of Nasutitermes acajutlae

  • Natural nests of termites
  • 94

  • Example artificial nest architectures
  • 101

Survival distributions resulting from Cox regressions as a function of treatment (a), species (b), and nest architecture (c) Survival distributions as a function of nest architecture Summary of survival across the five nest architectures for

Zootermopsis angusticollis  (Za), Reticulitermes flavipes  (Rf) and Nasutitermes corniger (Nc)

106

  • 108
  • 4.4

4.5

109

  • 110
  • 4.6

5.1
Median nest temperature (a) and % RH (b) across nest architectures Diagram depicting the timing of hemolymph extractions one and two,

  • and the combinations of treatment groups
  • 131

134 135
5.2 5.3 5.4
Example hemolymph protein profiles from a NCE individual Median masses of new proteins across direct exposure treatments Median numbers of new proteins per individual across direct exposure

  • treatments
  • 136

137 138
5.5 5.6
Median masses of new proteins across naïve treatments Median numbers of new proteins per individual across naïve treatments
11

List of Tables

  • 1.1
  • A comparison of traits used as indicators of social complexity in

  • termites and their subsocial ancestors
  • 31

32 48 59 74
1.2 2.1 2.2 3.1 3.2
Selective forces implicated in the evolution of social insect nests Final regression model outputs for bacterial (A) and fungal (B) CFUs Climatological data for the U.S. Virgin Islands Identifications of fungi associated with Nasutitermes acajutlae Jaccard coefficients comparing the similarity of core, trail, and soil

  • fungal genera
  • 77

  • 79
  • 3.3

4.1
Taxonomic groups of fungi associated with Nasutitermes acajutlae Survival parameters for control and conidia exposed termites of the

  • three study species
  • 107

12

Chapter 1 – Introduction

Termites are eusocial roaches (Inward et al., 2007a) and are unique in that they are the only roaches to have evolved eusociality. Termites are of particular interest in studies of eusociality because 1) they are the only insects to have developed eusociality outside the order Hymenoptera, and 2) unlike Hymenopterans, which have haplo-diploid sex determination, termites are diplo-diploid and thus lack the genetic predispositions for the evolution of sterile castes (Thorne, 1997). Investigating the evolutionary history of termites and eusocial development is difficult due to the lack of extant solitary species within the order Isoptera. Instead, factors promoting eusociality must be inferred through studies of the fossil record, the presocial roach ancestors of termites, and different aspects of the biology of extant species, including the degree of complexity of their nests. Nest building is an innate behavior (Emerson, 1938; Theraulaz et al., 1998, 2003), often resulting in species-specific architectures. These nests are the result of the cooperative effort of a large number of individuals working together to build, organize, and expand the structure. In other words, nests represent an extended phenotype of a superorganism (Dawkins, 1999; Turner, 2004). Studies of extant termites have shown a correlation between increasing sociality and larger colonies with more complex architectures (Eggleton and Tayasu, 2001). When preserved as fossils, nest structures provide a historical record of the degree of social complexity of the builders (Hasiotis, 2003). As such, nests provide a useful tool for indirectly studying eusocial evolution in insects.
Although ubiquitous among social insects, the nests of termites in particular exhibit a wide diversity of structures. This diversity is not only related to differences in
13 social complexity, but also to adaptations evolved in response to the needs of the colony:

defense, food storage, and a homeostatic environment necessary for survival and growth of offspring (Emerson, 1938; Noirot, 1970; Noirot and Darlington, 2000). Evolution of social insect nest architectures across genera has been studied to a certain degree in bees (Eickwort and Sakagami, 1979; Packer, 1991; Franck et al., 2004; Roubik, 2006; Rasmussen and Camargo, 2008), and wasps (Jeanne, 1975; Wenzel, 1991) likely due to the ability to compare a wide range of solitary to eusocial extant species. In ants, evolution of diverse nesting strategies in the genus Polyrhachis has been examined (Robson, 2004; Robson and Kohout, 2005, 2007). Tschinkel (2003, 2004, 2005) and Mikheyev and Tschinkel (2004) have done extensive work on the nest architectures and ontogeny of subterranean ant nests that provide vital information for analysis of trace fossils. However, studies of phylogenetic relationships of termite nests are few, leaving the evolution of the most diverse nest structures largely unexplored (Schmidt 1955, 1958; Abe, 1987; Inward et al., 2007b). This work reviews what is currently known regarding social insect nesting and evolution, explores the interrelationships between pathogens and termites in nests and the role nests play in pathogen resistance, and the possible impact of pathogens as selection pressures in the evolution of termite nesting behavior.

1.1 Ancestry and eusocial evolution

Recent phylogenetic analyses of extinct and extant species have confirmed that roaches of genus Cryptocercus are the living sister group to termites (Lo et al., 2000; Inward et al., 2007a; Engel et al., 2009). Morphologically speaking, Mastotermes darwiniensis Froggatt most closely resembles cryptocercid roaches, and is considered to
14 be the most primitive extant termite. However, in terms of social behavior, the extant

Termopsidae (recently renamed Archotermopsidae by Engel et al., 2009) exhibit the most primitive condition. Similar to members of Archotermopsidae, colonies of C. punctulatus live in networks of mostly horizontal galleries in dead wood of varying decay (Nalepa, 1984). Members of these two taxonomic groups sometimes occupy the same piece of wood, and share similar nesting habits: there is no evidence they leave the nest to forage for food. Instead, they consume wood they excavate from their nest, and digest it by means of symbiotic cellulolytic gut microorganisms (Cleveland et al., 1934; Bell et al., 2007). Cryptocercus punctulatus exhibits the presocial condition by nesting in family groups with overlapping generations that cooperate in brood care (Seelinger and Seelinger, 1983; Nalepa, 1984). A factor that separates subsocial cryptocercids from eusocial termites is that the former lack sterile castes, while termite workers not only remain in the parental nest, but also forego their own direct fitness to help raise the next generations of siblings (Wilson, 1971). It is not clear why extant cryptocercids have not evolved eusociality, but it is apparent that wood-feeding and colonial nesting preceded termite sociality. Nests under bark provide protection from the external environment, promote inbreeding through restricted movement, and also are costly to construct, all of which may have encouraged persistence of social behavior (Hamilton, 1978; Hansell, 1987). As a result, the nests of their presocial ancestors may have been a preadaptation for social evolution in termites.
15

1.2 Termites in the fossil record

Evidence of insect eusocial evolution is preserved in the fossil record. Termites were the first animals to evolve eusociality, predating ants by roughly 35 million years. However, there are gaps in the fossil history because not all species lived in environments conducive to fossil preservation. Moreover, there are surely specimens yet to be found. Only recently have scientists begun to analyze relationships between fossil species of termites and the 2,958 extant species (Engel et al., 2009). The oldest specimens of eusocial termites come from the Early Cretaceous, between 137-130 mya (MartínezDelclòs and Martinell, 1995; Rasnitsyn, 2008), but termites are believed to have diverged from roaches in the Late Jurassic, ~156-145 mya (Engel et al., 2009). Most fossil termites found are winged reproductives preserved in amber, or as depressions in limestone (reviewed by Engel et al., 2009). One worker was found from the Early Cretaceous (Martínez-Delclòs and Martinell, 1995) and soldiers have been found in Miocene (~23-5 mya) Dominican amber (Krishna and Grimaldi, 2009). These individual termite specimens demonstrate the existence of eusociality by the presence of differentiated castes and winged reproductives, which are absent in presocial roaches. They can also be compared to extant species to gauge what other common characteristics these species share. However, individual specimens cannot provide evidence for how large or complex their colonies were. Only fossil nests can show the dynamics of extinct superorganisms.
Compared to fossil insects, diagnosing structures within the fossil record as termite nests is open to interpretation. Bordy et al. (2004) reported having found an advanced fossil termite nest from the Early Jurassic (~200-175 mya) with evidence of fossil fungus gardens. Despite termite origins possibly dating back to the Jurassic, termite
16 body fossil evidence and estimates of divergence times have calculated mound-building

and fungus cultivation to have evolved much more recently (Genise et al., 2005; Engel et al., 2009). Currently, the oldest confirmed termite structure dates back to the Early Cretaceous, about the same age as the oldest termite body fossils (Francis and Harland, 2006). This find was a permineralized bored chamber within conifer wood. Although only a simple chamber, possibly part of a larger structure, it was identified as a termite excavation by the cylindrical, subhexagonal fecal pellets within the boring. Although cryptocercids also excavate galleries in wood (Nalepa, 1984), only extant termites are known to excrete fecal pellets of this shape (Genise, 1995). Fossil nests similar to those currently constructed by members of Hodotermitidae and Mastotermitidae have been found dating back to the Late Triassic, ~228-200 mya (Hasiotis and Dubiel, 1995). Other, more complex fossil nests have been found in the Chad basin, Africa dating ~7-3 mya. These included not only excavated galleries, but also structures obviously constructed and having a complexity akin to today’s Macrotermitinae, part of the most derived termite family Termitidae (Duringer et al., 2007). The range of ichnofossils, from simple to complex, has proven to be a useful interpretive tool for tracking social evolution through the fossil record (reviewed by Hasiotis, 2003).

1.3 Nest types, classifications, and associated levels of sociality

Abe (1984) identified six nesting trends based on different nest systems, and the feeding habits of the nest inhabitants: a) Drywood termites, which nest in and consume only the dry, hard wood in which they live (mainly the Kalotermitidae); b) Dampwood termites, which feed exclusively on the wet, decaying wood in which they live (the
17
Archotermopsidae, and some Rhinotermitidae); c) Intermediate termites, which nest in

wood and construct galleries in the soil or on the ground, and consume other wood sources in addition to the nest itself (Mastotermitidae, some Kalotermitidae, most Rhinotermitidae, and some Termitidae); d) Arboreal termites, which nest on living tree trunks and construct covered galleries leading to food sources completely separate from the nest (many Termitidae); e) Subterranean termites, which make epigeous and /or subterranean nests, also constructing galleries to food sources separate from the nest (Hodotermitidae, some Rhinotermitidae, Serritermitidae, and many Termitidae); and f) humus feeding termites, which make epigeous or subterranean nests and construct galleries to access humus food sources (many Termitidae). These trends were dubbed life types, and can be further grouped into three main categories of nests: one-piece, intermediate, and separate types of nests (Abe, 1987).
Similar to their cryptocercid ancestors, termites from the basal Archotermopsidae live in and consume the same piece of decaying wood, excavating galleries that result in a relatively simple nest architecture (Abe, 1987). These more primitive termites, also part of the lower termites, are distinguished by having flagellated gut protozoa for cellulose digestion (Krishna, 1969), and having “functional” workers that are not completely sterile. Instead these pseudergates (also known as false workers) have a highly flexible development, and retain the ability to achieve reproductive status when the primary queen and king die (Thorne, 1997). In the higher termites of the Termitidae, a true worker caste is present. These workers have lower postembryonic developmental flexibility, no external sexual characteristics present, and in most species have no reproductive potential (Noirot, 1969). The nests in this family vary considerably in size, shape, materials used,
18 as well as methods of construction. For the termite species used in this research, and their

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  • Blattodea: Hodotermitidae) and Its Role As a Bioindicator of Heavy Metal Accumulation Risks in Saudi Arabia

    Blattodea: Hodotermitidae) and Its Role As a Bioindicator of Heavy Metal Accumulation Risks in Saudi Arabia

    Article Characterization of the 12S rRNA Gene Sequences of the Harvester Termite Anacanthotermes ochraceus (Blattodea: Hodotermitidae) and Its Role as A Bioindicator of Heavy Metal Accumulation Risks in Saudi Arabia Reem Alajmi 1,*, Rewaida Abdel-Gaber 1,2,* and Noura AlOtaibi 3 1 Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia 2 Zoology Department, Faculty of Science, Cairo University, Cairo 12613, Egypt 3 Department of Biology, Faculty of Science, Taif University, Taif 21974, Saudi Arabia; [email protected] * Correspondence: [email protected] (R.A.), [email protected] (R.A.-G.) Received: 28 December 2018; Accepted: 3 February 2019; Published: 8 February 2019 Abstract: Termites are social insects of economic importance that have a worldwide distribution. Identifying termite species has traditionally relied on morphometric characters. Recently, several mitochondrial genes have been used as genetic markers to determine the correlation between different species. Heavy metal accumulation causes serious health problems in humans and animals. Being involved in the food chain, insects are used as bioindicators of heavy metals. In the present study, 100 termite individuals of Anacanthotermes ochraceus were collected from two Saudi Arabian localities with different geoclimatic conditions (Riyadh and Taif). These individuals were subjected to morphological identification followed by molecular analysis using mitochondrial 12S rRNA gene sequence, thus confirming the morphological identification of A. ochraceus. Furthermore, a phylogenetic analysis was conducted to determine the genetic relationship between the acquired species and other termite species with sequences previously submitted in the GenBank database. Several heavy metals including Ca, Al, Mg, Zn, Fe, Cu, Mn, Ba, Cr, Co, Be, Ni, V, Pb, Cd, and Mo were measured in both collected termites and soil samples from both study sites.
  • Complementary Symbiont Contributions to Plant Decomposition in a Fungus-Farming Termite

    Complementary Symbiont Contributions to Plant Decomposition in a Fungus-Farming Termite

    Complementary symbiont contributions to plant decomposition in a fungus-farming termite Michael Poulsena,1,2, Haofu Hub,1, Cai Lib,c, Zhensheng Chenb, Luohao Xub, Saria Otania, Sanne Nygaarda, Tania Nobred,3, Sylvia Klaubaufe, Philipp M. Schindlerf, Frank Hauserg, Hailin Panb, Zhikai Yangb, Anton S. M. Sonnenbergh, Z. Wilhelm de Beeri, Yong Zhangb, Michael J. Wingfieldi, Cornelis J. P. Grimmelikhuijzeng, Ronald P. de Vriese, Judith Korbf,4, Duur K. Aanend, Jun Wangb,j, Jacobus J. Boomsmaa, and Guojie Zhanga,b,2 aCentre for Social Evolution, Department of Biology, University of Copenhagen, DK-2100 Copenhagen, Denmark; bChina National Genebank, BGI-Shenzen, Shenzhen 518083, China; cCentre for GeoGenetics, Natural History Museum of Denmark, University of Copenhagen, DK-1350 Copenhagen, Denmark; dLaboratory of Genetics, Wageningen University, 6708 PB, Wageningen, The Netherlands; eFungal Biodiversity Centre, Centraalbureau voor Schimmelcultures, Royal Netherlands Academy of Arts and Sciences, NL-3584 CT, Utrecht, The Netherlands; fBehavioral Biology, Fachbereich Biology/Chemistry, University of Osnabrück, D-49076 Osnabrück, Germany; gCenter for Functional and Comparative Insect Genomics, Department of Biology, University of Copenhagen, DK-2100 Copenhagen, Denmark; hDepartment of Plant Breeding, Wageningen University and Research Centre, NL-6708 PB, Wageningen, The Netherlands; iDepartment of Microbiology, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria SA-0083, South Africa; and jDepartment of Biology, University of Copenhagen, DK-2100 Copenhagen, Denmark Edited by Ian T. Baldwin, Max Planck Institute for Chemical Ecology, Jena, Germany, and approved August 15, 2014 (received for review October 24, 2013) Termites normally rely on gut symbionts to decompose organic levels-of-selection conflicts that need to be regulated (12).
  • Morphology and Gonad Development of Normal Soldiers and Reproductive Soldiers of the Termite Zootermopsis Nevadensis Nevadensis (Isoptera, Archotermopsidae)

    A peer-reviewed open-access journal ZooKeysMorphology 148: 15–30 and(2011) gonad development of normal soldiers and reproductive soldiers of the termite... 15 doi: 10.3897/zookeys.148.1672 RESEARCH ARTICLE www.zookeys.org Launched to accelerate biodiversity research Morphology and gonad development of normal soldiers and reproductive soldiers of the termite Zootermopsis nevadensis nevadensis (Isoptera, Archotermopsidae) Susan E. Johnson1, Nancy L. Breisch1, Bahram Momen2, Barbara L. Thorne1 1 Department of Entomology, University of Maryland, College Park, MD 20742, USA 2 Department of En- vironmental Science & Technology, University of Maryland, College Park, MD 20742, USA Corresponding authors: Barbara L. Thorne ([email protected]), Nancy L. Breisch ([email protected]) Academic editor: Michael Engel | Received 7 June 2011 | Accepted 9 August 2011 | Published 21 November 2011 Citation: Johnson SE, Breisch NL, Momen B, Thorne BL (2011) Morphology and gonad development of normal soldiers and reproductive soldiers of the termite Zootermopsis nevadensis nevadensis (Isoptera, Archotermopsidae). In: Engel MS (Ed) Contributions Celebrating Kumar Krishna. ZooKeys 148: 15–30. doi: 10.3897/zookeys.148.1672 Abstract Reproductive or neotenic soldiers of the Archotermopsid Zootermopsis nevadensis nevadensis (Hagen) are compared to sterile soldiers and primary male reproductives. Several head capsule morphometrics cor- relate significantly with gonad size across all forms and both sexes of soldiers. The easily observed field character of ratio of mandible length to labrum length is a consistent and reliable feature of head capsule external morphology for predicting gonad development and reproductive potential of soldier forms re- gardless of age, sex, or live weight. Keywords evolution of soldier caste, reproductive soldier, neotenic soldier, Zootermopsis, morphometrics Introduction Soldiers are a non-reproductive defensive caste in termites (though they may sometimes have other roles (Traniello 1981) and are not always the exclusive defensive caste in a colony (e.g.
  • The Biology of Nine Termite Species (Isoptera: Termitidae) from the Cerrado of Central Brazil

    The Biology of Nine Termite Species (Isoptera: Termitidae) from the Cerrado of Central Brazil

    THE BIOLOGY OF NINE TERMITE SPECIES (ISOPTERA: TERMITIDAE) FROM THE CERRADO OF CENTRAL BRAZIL BY HELEN R. COLES DE NEGRET AND KENT H. REDFORD INTRODUCTION The Neotropical region is second to the Ethiopian in numbers of described termite species (Araujo 1970). However, little is known of their biology. The literature on Brazilian termites is largely re- stricted to isolated taxonomic descriptions of species from the Amazon Basin and southern states of Brazil (Araujo 1961, 1969, 1977 and Fontes 1979). Exceptions to this include information re- lating termite species and their distribution to vegetation types in Mato Grosso State (Mathews 1977), the effect of deforestation on termites in the Amazon (Bandeira 1979) and data on the ecology and defense of termites in the cerrado vegetation of the Distrito Federal (Coles 1980). The present study was done in conjunction with a study on mammalian termite predators, in particular the giant anteater, Myrmecophaga tridactyla (Coles 1980 and Redford in prep.). Six aspects of termite biology of importance in defense by termites against mammalian predators were studied for nine of the most common mound-building termite species in the Distrito Federal, Brazil. Reported here are individual weights, morphology of soldier castes, worker-soldier ratios, mound sizes and forms, mound hard- nesses and nest materials, distributions and abundances of nests and feeding habits for these nine species. All species studied were from the family Termitidae (see Fig. for comparison of soldier heads), subfamily Apicotermitinae, Grigioter- mes rnetoecus (Matthews); subfamily Nasutitermitinae, Armitermes Laboratoria de Zoologia e Ecologia Animal, Universidade de Brasilia, Brasilia D. F. 80910, Brazil.
  • 14128 JGI CR 07:2007 JGI Progress Report

    14128 JGI CR 07:2007 JGI Progress Report

    U.S. JOINT DEPARTMENT GENOME OF ENERGY INSTITUTE PROGRESS REPORT 2007 On the cover: The eucalyptus tree was selected in 2007 for se- quencing by the JGI. The microbial community in the termite hindgut of Nasutitermes corniger was the subject of a study published in the November 22, 2007 edition of the journal, Nature. JGI Mission The U.S. Department of Energy Joint Genome Institute, supported by the DOE Office of Science, unites the expertise of five national laboratories—Lawrence Berkeley, Lawrence Livermore, Los Alamos, Oak Ridge, and Pacific Northwest — along with the Stanford Human Genome Center to advance genomics in support of the DOE mis- sions related to clean energy generation and environmental char- acterization and cleanup. JGI’s Walnut Creek, CA, Production Genomics Facility provides integrated high-throughput sequencing and computational analysis that enable systems-based scientific approaches to these challenges. U.S. DEPARTMENT OF ENERGY JOINT GENOME INSTITUTE PROGRESS REPORT 2007 JGI PROGRESS REPORT 2007 Director’s Perspective . 4 JGI History. 7 Partner Laboratories . 9 JGI Departments and Programs . 13 JGI User Community . 19 Genomics Approaches to Advancing Next Generation Biofuels . 21 JGI’s Plant Biomass Portfolio . 24 JGI’s Microbial Portfolio . 30 Symbiotic Organisms . 30 Microbes That Break Down Biomass . 32 Microbes That Ferment Sugars Into Ethanol . 34 Carbon Cycling . 39 Understanding Algae’s Role in Photosynthesis and Carbon Capture . 39 Microbial Bioremediation . 43 Microbial Managers of the Nitrogen Cycle . 43 Microbial Management of Wastewater . 44 Exploratory Sequence-Based Science . 47 Genomic Encyclopedia for Bacteria and Archaea (GEBA) . 47 Functional Analysis of Horizontal Gene Transfer . 47 Anemone Genome Gives Glimpse of Multicelled Ancestors .
  • Termite Prevention and Control by Design in Florida

    Termite Prevention and Control by Design in Florida

    Termite Prevention and Control by Design in the Southeast Presented by Faith Oi, Ph.D., University of Florida, Entomology and Nematology Dept., Gainesville, FL 32611. email: [email protected] Disclaimer: This presentation was developed by a third party and is not funded by WoodWorks or the Softwood Lumber Board. “The Wood Products Council” is a This course is registered with AIA CES Registered Provider with The American for continuing professional education. Institute of Architects Continuing As such, it does not include content Education Systems (AIA/CES), Provider that may be deemed or construed to #G516. be an approval or endorsement by the AIA of any material of construction or Credit(s) earned on completion of this any method or manner of handling, course will be reported to AIA CES for using, distributing, or dealing in any AIA members. Certificates of Completion material or product. for both AIA members and non-AIA ______________________________ members are available upon request. Questions related to specific materials, methods, and services will be addressed at the conclusion of this presentation. Course Description Building enclosures are responsible for controlling heat flow, air flow, vapor flow and a number of other elements. In the southern US, they are also essential for termite prevention. This presentation will explore design considerations associated with wood- frame building enclosures and the role of control layers in addressing items such as durability, termite prevention and control, and thermal continuity. Beginning with a review of building enclosure design fundamentals and considerations, it will then cover best practices for a variety of wood-frame building enclosure assemblies and details.