PHYLOGENY OF THE AMMOBATINI AND REVISION OF THE AFROTROPICAL GENERA (HYMENOPTERA: ANTHOPHORIDAE: NOMADINAE) by CONNAL DESMOND EARDLEY Submitted in fulfilment of the academic requirements for the degree of Doctor of Philosophy in the Department of Zoology and Entomology University of Natal Pietermaritzburg 1994 ABSTRACT The phylogeny of the Ammobatini was studied, with regard to the principles of cladistics using parsimony, and the classification is revised. It is concluded that the tribe fonns a monophyletic group that comprises six distinct monophyletic genera: Pasite Jurine, Sphecodopsis Bischoff, Ammobates Latreille, Me/anempis Saussure, Spilwpasites Warncke and Oreopasites Cockerell, of which Pasites, Sphecodopsis, Ammobates and MeLanempis occur in the Afrotropical Region. The Afrotropical species of these four genera are revised. Pseudopasites Bischoff and Pseudodichroa Bischoff are synonymized with Sphecodopsis. Pasites includes 17 Afrotropical species, Sphecodopsis 10 species, and Ammobates and MeLanempis are each known from a single Afrotropical species. Ten new species are described: Pa..~ites nilssoni, P. paulyi, P. humecta, P. glwma, P. namibiensis, P. somaLica, Sphecodopsis vespericena, S. longipygidium, S. namaquensis and Ammobates auster. Thirty-three names are synonymized: they are P. nigerrima (Friese), P. argentata (Baker) (= P. barkeri (Cockereil»; P. chubbi Cockerell, P. nigritula Bischoff, P. peratra Cockerell (= P. atra Friese); P. nigripes (Friese), P. fortis Cockerell, P. subfortis Cockerell, P. stordyi Cockerell, P. voiensis Cockerell, P. aitior Cockerell (= P. carnifex (Gerstaecker»; P. Ilataiensis (Cockerell), P. aiboguttatus (Friese), P. ogiiviei (Cockerell) (= P. jenseni (Friese»; P. alivalensis (Cockerell), P. rufitarsis (Cockerell) (= P. histrio (Gerstaecker»; P. marshaUi (Cockerell) (= P. jonesi (Cockerell»; P. abessinica (Friese), P. fulviventris (Bischoff), P. rhodesialla (Bischoff), P. apicalis (Bischoff), P. turneri (Cockerell), P.polituLa (Cockerell), P. indecisa (Cockerell), P. nudicauda (Cockerell), P. bechuanica (Cockerell), P. breviceps (Cockerell) (= P. appLetoni (Cockerell); S. rufula (Cockerell) (= S. minutissima (Cockerell»; S. pygmaea (Friese), S. rufescens Bischoff, S. aLgoensis Bischoff, S. perpunctata Cockerell (= S. acuLeata (Friese»; S. leonis (Cockerell) (= S. semirufa (Cockerell». Keys to the genera and species are provided. 11 PREFACE The work described in this thesis was carried out at the Biosystematics Division, Plant Protection Research Institute, Pretoria, from February 1988 to December 1994, the last two years under the supervision of Professor Denis J. Brothers, of the University of Natal, Pietermari tzburg. These studies represent original work of the author and have not otherwise been submitted in any form for any degree or diploma to any University. Where use has been made of the work of others it is duly acknowledged in the text. iii CONTENTS 1 INTRODUCTION. .. 1 2 HISTORICAL REVIEW OF THE AMMOBATINI . .. 3 3 MATERIALS AND METIIODS .. .. 5 3.1 INTRODUCTION......................................... 5 3.2 MORPHOLOGICAL TERMS At'ID DESCRIPTIONS ............... 5 3.3 CLADISTIC METIIODS . .. 6 3.4 MATERIAL EXAMINED..... .............. ... ...... ...... 9 4 PHYLOGENY OF THE A.\1M:OBATINI .................. _. .. 11 4.1 INTRODUCTION ......................................... 11 4.2 ANALYSIS OF ADULT CHARACTERS ........................ 11 4.2.1 SPECIES Al'ID SPECIES GROUPS . .. 11 4.2.2 GENERIC GROUND PLANS ............. .. 13 4.2.3 GENERIC RELATIONSHIPS ........... .... ............... 13 4.2.4 TRIBAL RELATIONSHIPS ................................ 15 4.3 LARVAL CHARACTERS ................................... 15 4.4 ZOOGEOGRAPHY.... .. .... .. 15 4.5 CONCLUSION........................................... 17 5 REVISION OF THE AFROTROPICAL AMMOBATINI . .. 18 5.1 INTRODUCTION....... ................................. 18 5.2 GENUS PASITES JURINE .................................. 19 5.3 GENUS SPHECODOPSIS BISCHOFF .......................... 68 5.4 GENUS AMMOBATES LATREILLE ........................... 88 5.5 GENUS MELANEMPIS SAUSSURE ........................... 90 6 ACKNOWLEDGEMENTS . .. 91 7 REFERENCES ......... ................................ 92 8 INDEX ................. ................................ 97 1 1 INTRODUCTION The purpose of this study was to provide a comprehensive revision of the systematics of the Afrotropical Ammobatini (Anthophoridae; Nomadinae). To attain this Objective three main aspects were investigated. The first was to establish whether the Ammobatini is a monophyletic taxon and, in so doing, establish its validity. The second was to review the generic classification of the Ammobatini and gain an understanding of the relationships between the genera . The third was to acquire a sound knowledge of the identity of the Afrotropical species and the relationships between species. The Ammobatini are cJeptoparasitic bees, also known as cuckoo-bees. They lay their eggs in the nests of pollen-collecting bees that provision their larval cells with sufficient food [or the larva to develop to maturity. The cleptoparasite kills the host larva and utilizes its provisions. Pollinating bees are one of the most important groups of beneficial insects. It is therefore important to study the systematics and biology of their cJeptoparasites because they affect the population dynamics of pollen-collecting bees. Bees of the ammobatine genus Sphecodopsis Bischoff, for example, are known to parasitise species of Scrapter Lepeletier & Serville (Rozen & Michener 1968). Both of these genera are endemic to southern Africa. Scrapter pollinates indigenous plants and is potentially important in the pollination of agricultural crops. It is prevalent in the semi-arid regions of southern Africa where insect-pollinated indigenous plants form an important component of the ground cover and pasture. The cuckoo-bee subfamily Nomadinae, to which the Ammobatini belongs, is the largest and most diverse lineage of cleptoparasitic bees. The Nomadinae comprise ten distinct tribes (Roig-Alsina 1991) and about 1200 described species (Alexander 1990). This subfamily has its greatest diversity in the Nearctic and Neotropical Regions, is fairly well represented in the Afrotropical and Palaearctic Regions and is poorly known from the Oriental and Australian Regions. In the Afrotropical Region, the subfamily comprises four tribes: Nomadini, Epeolini, Ammobatoidini and Ammobatini. The Nomadini, Epeolini and Ammobatoidini are 2 represented there only by their nominate genera. The Nomadini and the Epeolini were revised by Eardley & Schwarz (1991) and Eardley (1991), respectively. The Ammobatoidini is known from the Afrotropical Region from a single female specimen described as Ammobatoides braunsi Bischoff. The Arnmobatini is the largest and most diverse tribe of Afrotropical nomadine bees. The Ammobatini occur mainly in the Nearctic, Palaearctic and Afrotropical Regions. Although they have not been recorded from the Oriental Region, several Palaearctic species. are known from areas that border the Oriental Region and these species possibly extend into that region. They do not occur in the Neotropical and Australian Regions. The Nearctic and Palaearctic faunas have been well studied and the Afrotropical fauna is revised here. The tribal classification of the subfamily has been extensively reviewed (Alexander 1990; Roig-Alsina 1991). Current knowledge of the systematics of the Nomadinae makes the Arnmobatini well suited for phylogenetic research and such a study is needed before the systematics of the subfamily can be properly understood. Most of the previous work on the systematics of the Afrotropical Ammobatini consists of descriptions of new species and distribution records. The original descriptions are generally vague, without illustrations and inadequate for the recognition of the species. Bischoff (1923) provided a comprehensive revision of the Afrotropical Arnmobatini, but his work has several shortcomings, such as that he did not study much of the type material, and it has subsequently become outdated. The present study is the first treatment of these bees that included an examination of nearly all the type material and a study of male and female terminalia. The research was based mainly on adult morphology as only sufficient material of the adult stage was available for such a study. In spite of the excellent work on ammobatine larvae by Rozen (1954, 1966), Rozen & McGinley (1974) and Rozen & Roig-Alsina (1991), Rozen & McGinley (1974) clearly stated that insufficient data were available on ammobatine larvae for a phylogenetic analysis of the tribe, and little additional information has subsequently accrued. Nevertheless, a preliminary investigation into the phylogeny of ammobatine larvae 3 was undertaken, but proved to be of little value in the interpretation of the relationships between the genera. 2 mSTORICAL REVIEW OF THEAMMOBATINI The history of the classification of this group of closely-related bees may be outlined as follows. Recent literature, e.g. Alexander (1990) and Roig-Alsina (1991), considers them to belong to the Anthophoridae. The current familial placement of these bees, a topic that is beyond the scope of the study, has been accepted. The first significant attempt to arrange the bees into a system of higher classification was by Dalla Torre (1896). He placed all bees in the family Apidae and placed the genera that are currently considered to belong to the Ammobatini, namely