SPECIES DIVERSITY AND BIOGEOGRAPHY OF ANTS (HYMENOPTERA: FORMICIDAE) IN LOUISIANA, WITH NOTES ON THEIR ECOLOGY A Thesis Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College In partial fulfillment of the Requirements for the degree of Master of Science In The Department of Entomology by Shawn T. Dash B.S., University of Delaware, 2002 December 2004 Dedication This work is dedicated to my mother, who got up before dawn to take me bird watching when I was young, who “allowed” me to use the bathtub for my turtles, who never minded hundreds of bottles and cans of insects in the freezer. Two other people share responsibility for who I am today, and I dedicate this work to them as well. Charles Bartlett who is a teacher, a mentor, and a life-long friend and Jake Bowman, without his advice on science and life I may never have made it. ii Acknowledgements Many persons contributed to my research and this letter is in the spirit of gratitude and acknowledgment. I would like to thank my major professor Dr. Linda Hooper-Bui, who was there to lend support and offer an ear to hear my ideas. Dr. Chris Carlton who always had useful advice and suggestions to improve as a scientist and a writer. Dr. Dorothy Prowell who listened to my thoughts, and was kind enough to allow me to use specimens from her ongoing research . A number of people lent their time and energy to this project, Andrew Cline provided hours of conversation and methodology (especially the skill of sifting). He also generously gave me specimens for my analysis and often joined me in the field. A heart-felt thank you to Michael Seymour who photographed the majority of species and aided me in the creation of the distributional maps. A very special acknowledgment must be give to Dr. John Moser who always encouraged me to strive for more and was always willing to talk science; he was also kind enough to donate a number of specimens. Other persons who have made major contributions of specimens included, M. Seymour, D. Colby, A. Tishechkin, J. Rosson, K. Landry, L. Womack, N. Nguyen, E. Watson, A T. Pranschke. I would also like to thank those who helped me in the field and in the laboratory: K. O’Brien, R. Baillif, J. Fleming, S. Scoby, N. Saucier, K. Prejean and A. Williamson. Stacy Clayton in the creation of web page supplements to my thesis. I would also like to thank Brian Fisher and Stefan Cover for organizing “The Ant Course” which allowed me to improve and tune my taxonomic skills. I would also like to thank Kye Hudland, Lloyd Davis, Bill MacKay, James Trager, and Mark Deyrup who were always available with friendly advice and were kind enough to review my identifications. iii Table of Contents ACKNOWLEDGEMENTS………………………………………………………..……………..iii ABSTRACT………………………. …………………………………….……………..…………v CHAPTER 1. INTRODUCTION……………………...………………………….….. …...…………1 2. METHODS………………………………….…………………………..……………..7 Study Area……………………………………………………………………...…7 Sampling Protocol………………………………………………………………....7 Processing, Identification, and Analysis………………………………………....10 3. RESULTS……………………………………………………...……………...….….14 Summary of Results...……………………………………………...…………….14 Foreward to Species Accounts….....………………………………..…………....17 Key to the Ants of Louisiana…………...…………………………..……….…...20 Ectioniae……………………………………………………………..………..…21 Ponerinae……………………………………………………………………...…38 Proceratiinae……………………………………………………...………...……50 Ectatomminae……………………………………………………………………55 Pseudomyrmecinae…………………………………………………….………...63 Myrmecinae…………………………………………………………………...…71 Dolichoderinae………………………………………………………… ………185 Formicinae……………………………………………………………………...203 4. DISCUSSION...……..…………………………………………………………...…249 LITERATURE CITED………………………………………………………………………....263 APPENDIX A. ECOREGIONS IN LOUISIANA……………………………………………………283 B. PARISHES IN LOUISIANA………………………………………………………...284 C. LIST OF ANTS IN LOUISIANA……………………………………………………285 D. RECORDS OF SPECIES CONSIDERED TO BE PREVIOULSY UNREPORTED FROM LOUISIANA …………………………………………………………..........288 E. SPECIES OF ANTS THAT ARE LIKELY TO OCCUR IN LOUISIANA ………..289 VITA……………………………………………………………………………………………290 iv Abstract Over three decades have passed since the completion of Moser and Blum’s (1960) treatment of the ants of Louisiana. This research represents a significant contribution to and update of their work. I report the first-ever comprehensive assessment of the Louisiana Formicid fauna. More than 100 species from 38 genera and eight subfamilies are covered. This faunistic survey gives valuable data of biodiversity and macro/microhabitat selection of ant species. Included are keys to all subfamilies, genera, and species with illustrations photographs, and a regional map for each species. v Chapter 1 Introduction This research examines the biodiversity of ants in Louisiana. This project also expands the knowledge base of ants found in the state by providing details on biogeography and observations on natural history. This research is not manipulative but rather observational or measureative. It presents contributions to biology, based on scientific investigations of a diverse family of insects. One of the most fundamental areas in biology is the study of a species; once one knows what something is the door to understanding it opens. Investigations into taxonomy, phylogenics, behavior, and natural history have been afforded to relatively few taxa. Even with many scientist studying invertebrates, limited research coverage has been accomplished. The limited treatment may be an artifact of the diversity of these groups. For example the class Mammalia has about 4,000 species, and the phylum such as Ctenophora/Cnidaria has about 9,000 species (Wilson 1992), but the family Formicidae (ants) has nearly triple as many species as mammals (LaSalle and Gauld 1993). The limited amount of research on insects is understandable with such a hyper- diverse group, further study is needed. To understand the broader themes in biological research such as ecology, evolution, as well as medical research (pharmaceuticals), the establishment of what an organism is, what it does, and how it is related to other biota is required. As Sharkey (2001) observed, contrary to popular belief, the oldest profession of humans has been taxonomy. He cites the Bible as stating in Genesis 4:14 God presented to man the duty of naming all the “beasts of the land, all the fish of the sea, and the birds in the sky.” This naming tradition has continued for thousands of years and, in the past 246 years (Linnaeus 1758), has become a well-defined science. However, once a popular part of natural history study, 1 the field of taxonomy does not have the same number of scientist it once had. For a number of years, the number of up-and-coming taxonomists has diminished rapidly. Nevertheless, a push has been made to rekindle the most basic unit of natural history studies, and in recent years, classical taxonomy and systematic research has begun to reemerge (Wilson 2000) however, the resources to study biodiversity are deteriorating (Edwards 1984). The importance of knowing what species exist and how many occur is one of the most basic questions in biology and leads to many more. Thoroughly understanding ecosystems has become extremely important as humankind’s practices have led to a decline in environmental conditions (Wilson 1998). There is a widely- held idea that mass decline of species diversity causes ecological instability, which directly affects all life on earth (Agosti et al. 2000). Luckily, considerable research is underway to investigate patterns of biodiversity in the context of the ongoing mass extinction (Wilson 1992, Agosti et al. 2000). For some an innate biophilic desire to know what is out there and what is being lost exist (Wilson 1985). One emerging method is that of rapid assessments; these studies take a picture of local diversity. Such studies often produce regional lists that not only aid in further research but also are beneficial to local communities by raising local awareness. Once a common practice in biology, the creation of regional lists for species is not a common practice today yet the need is apparent. One group receiving attention is the insect order Hymenoptera (ants, bees, wasps, and their kin, LaSalle and Gauld 1993, Agosti et al. 2000). As Gauld is persistent in pointing out, this order is hyper-diverse and this biodiversity is indicated by the lack of a “unifying common name” (LaSalle and Gauld 1993, Gauld and Bolton 1998). In terms of numbers, dominance, and ecological importance, Hymenoptera are fundamentally significant (LaSalle and Gauld 1993). 2 One family alone within this order can be used to exemplify the importance of insects and biodiversity, the ants (formicidae) (Hölldobler and Wilson 1990, Agosti et al. 2000, LaSalle and Gauld 1993). LaSalle and Gauld (1993) note that ants strongly influence the functionality of ecosystems. Ants dominate by their massive biomass, manipulate species composition, influence trophic interactions, possess numerous mutalistic and symbiotic relationships, and shape both the abiotic (eg: moving soil) and biotic (eg: plant-ant mosaics) matrix of community interactions. To accurately assess biodiversity and community dynamics, focal taxa require certain criteria. These criteria include: ecological significance, diversity, and ease of collection. In the past, these efforts were focused mostly on birds, vascular plants, and mammals (Johnson 1986, Wilson 2000). A number of recent studies have shifted the focus to include invertebrates such as beetles, butterflies, ants and aquatic macroinvertbrates (Anderson 1997, McCafferty 1981, Olembo 1991, Wilson 2001). Ants have been recognized as excellent candidate insects for such studies (Agosti 2003). Formicidae contains 21 subfamilies, 358 genera, and more than 11,000 species worldwide (Agosti 2003, Bolton 2003). In North America, there are 10 subfamilies, 70 genera and 983 species. Ants are well suited for studies in taxonomy, biodiversity, community composition, and biogeography (Hölldobler and Wilson 1990, Alonso and Agosti 2000). For example, ants are noted for their proportionally large biomass in ecosystems, making them an important and dominant component of all terrestrial habitats. One area in the Amazon rain forest, ants and termites together weigh more than all land vertebrates combined (Hölldobler and Wilson 1990). Erwin (1989) reported in his canopy studies that 69% of specimens were ants. Ants also fill numerous niches and often act as “keystone species.” Modifying Solbrig’s (1991) categories of keystone species: predator, mutualist, and resource species, LaSalle and 3 Gauld (1993) generate support for the contributions in each context for ants. Ants can act as a keystone predator, such as weaver ants (Oecophylla). In the category of keystone mutualist, ants are needed by some plants for protection, nutrient recycling, and dispersal of pollen and seeds (myrmecochory, seed dispersal by ants) (LaSalle and Gauld 1993 and references therein). Ants are keystone resource species as well, they influence soil structure by turning earth and transporting organic material into their nest, acting to fertilize soil (Hölldobler and Wilson 1990, LaSalle and Gauld 1993). In addition to being a food resource for other organisms, from other arthropods to mammals (Xenarthra) that specialize in eating ants, ants are associated with numerous organisms, including vertebrates that require them for their survival. Such interaction can be seen in army ants and ant birds which follow the raiding trails of army ants. One other example is the nest location of some neotropical birds that “nest in ant-occupied trees” (LaSalle and Gauld 1993). In addition to high diversity, ants possess a quasi-stable taxonomic and systematic status (Bolton 2003) which creates a situation where identification is assessable and widely understood (Kaspari 2000). A significant study would be the assessment of biodiversity, community composition, biogeography, and other basic investigation of ecology of a regional biota. Such a study would grant researchers a background from which to begin further more detailed studies. The current research will examine regional ant fauna in Louisiana in a taxonomic and ecological context. Louisiana is distinctive in the number of diverse communities and unique habitats. The Nature Conservancy lists eight main ecoregions. Some more distinctive habitats are the Tunica Hills ecoregions and southern prairies in the coastal prairie ecoregions. More studies investigating the unique fauna and flora of Louisiana are badly needed (The Nature Conservancy of Louisiana 2004). Louisiana is in a current state of flux with wide-spread logging and 4 urbanization. Consequently research on biodiversity is paramount and required in a time frame when resident species diversity can still be assessed in Louisiana. Moser and Blum (1960) generated a list of ants in Louisiana that included a total of 128 potential species for the state. This number requires some amending to reflect taxonomic status and actual occurrence in the state. Sixty four of the 123(after considering synonyms) species were presumed species. They were recorded in adjacent states and were thought to occur in Louisiana. Because of the heterogeneity of habitats in Louisiana and surrounding states, the assumption of species overlap requires reevaluation of those species. In addition, their study only included species and no distributional data. Louisiana’s ant diversity not only needs research to clarify previous work, but could yield new species as well. When the Wheelers traveled through St. Tammany Parish, they discovered a species of Brachymyrmex not known to occur in the US (Wheeler and Wheeler 1978). They found a small colony of Brachymyrmex musculus Forel at a wayside park and cited the likelihood of many introduced species to Louisiana because of the busy port in New Orleans (Wheeler and Wheeler 1978). Moser (pers. com.) noted that at the time the Louisiana list was published, the red imported fire ant (Solenopsis invicta Buren) had only infested a few parishes west of East Baton Rouge Parish (about 23 parishes). Callcott and Collins (1996) noted that every parish in the state had been infested by 1975. With the introduction of S. invicta and Linepithema humile (Mayr), the assumption of community disruption and species displacement would occur is logical. Additionally other eradication methods for these pest ants may have affected species diversity. With this information the need for an updated list becomes clear. A study that examines the biodiversity of ants in Louisiana is overdue. This research investigates three major research questions: 1) has the species richness changed over time 2) has 5