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The Taxonomy and General Biology of Some Southern Australian

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The Taxonomy and General Biology of Some Southern Australian 3ó .r-:5 BIOLOGY THE TAXONO¡4Y AND GENERAL OF SOME (OIPIERA: nn¡¡etocnn¡) SOUTHERN AUSTRALIAI{ CHIRONOMIDAE BY INGRID A. HERGSTROM B.Sc (ttons) Adel. UNIVERSITY OF ADELAIDE DEPARTMENT OF ZOOLOGY' L974 of Adelaide A thesis submitted to ttre University for the degree of Doctor of Philoscphy' J DECLARATION been accepted This thesis contains no material whictr- has at any university for the award of any other degree or diploma andtothebestofmyknowledgecontainsnomaterialprrblished reference is or written by anottrer Person' except where due made. (r. a. HERGSTRoM) CONTEI{TS Page (i) SUMMARY (ii) ACKNOW'LEDGEME}CTS 1 I. INIRODUCTION 3 II. GENERAL BIOLOGY 3 I INTRODUCTION 4 2. METHODS 2.I Collecti-on 4 2.2 Emergence Data 6 2.3 Rearing 6 2.4 Behaviour 8 I 2.5 Measurement of Humidity I 2.6 lvleasurernent of Acidity I 2-7 Measurement of OxYgen Levels 9 3 COI,LECTING SITES 3.1 Site descriPtions 9 29 4 BIOLOGY OF INDIVIDUAL SPECIES SubfamilY Podonominae 30 32 Subf amilY APhroteniinae SubfamilY Diamesinae' Tril¡e HePtagYini 32 32 Sr:bf anilY Orthocladiinae 51 5. DISCUSSTON 54 III. MORPHOLOGY 54 1. INTRODUCTION 56 2. METHODS 2.I Collection of sPecimens 56 2.2 Preservation of sPecimens 56 2.3 PreParation of sPecimens 56 Page 2.4 Drawings 57 2.5 Recommendations regarding the preservation and curating of Chironomid collections 57 57 3 STRUCTURES OF TAXONOMIC IMPORTANCE 3.1 Adult 58 3.2 Pupa 65 3.3 Larva 65 4 SPECIES DESCRIPTIONS 66 Sub-familY APHROTENIINAE 68 Genus Aphroteniinae 68 Sub-family DIAMESINAE, TRIBE HEPTAGYINI 69 Genus ParahePtagYia 69 Sr¡b-f amilY ORTHOCIJIDI INAE 72 Genus Austrobrillia 76 Genus Cardiocladius 79 Genus Stfcbcladlus 80 Genus Eukiefferiella 84 86 Genus Svnorthocladius 'ee Genus Paratrictrocladius Genus Cr 92 Genus Microcricotopus 98 Genus Nasuticladius 10r Genus Àustrocladius 117 Genus LimnoPhYes r29 Genus ParalimnoPhYes L32 Genus SParsicladius 136 143 Genus Nemoricladius Genus KiefferoPhYes L47 Genus Parametriocn ernus L54 156 Genus Gymnolnel-r iocnemus Page Genus Rhinocladius 160 Genus Smittia I63 Genus Parakief feriella 165 Genus Pseudosmittia 167 Genus Doloplastus r69 IV. PHYLOGENY 175 I INTRODUCTION 175 2. BRT]NDTN S USE OF HENNIGIS PIIYLOGENETTC SYSTE¡4ATTCS r78 3 PHYLOGENETIC SYSTEMATICS AND THE EVOLUTIONARY INDEX r84 4 A METHOD OF TND]CATING RELIABILITY OF PROPOSED PHYLOGENIES 186 5 PTIYLOGENETIC RE],ATTONSHIPS WITHTN TIIE SUB- FAMILY APHROTENITNAE 186 6. PHYLOGENETIC REI.ATIONSHTPS WITHIN THE SUB' FAMILY ORTHOCI,ADIINAE t87 6.1 Important Character Phylegsti.= 190 6.2 Genus Austrobrillia 193 6.3 cenus Cardiocladius ]-94 6.4 The Diplocladius-S tictocladius genus pa]-r 195 6.5 Genus Eukiefferiella 196 6.6. Genus SYnorÈhocladius l-97 6.7 Ttre Paratrichocladius groups of genera L97 6 .8 Genus Ml-crocricotopus 199 6.9 the Çb¿S.cla4ius group of genera 2OO 6. to The Nasutic lad.ius group 203 6. 11 Genus Parametriocnemus 206 6.L2 Genus jj¡g5l.!!ig 2c-7 6.13 Genus ParakfefferfellE 2U.7 Page 6.L4 The Pseudosmittia and Doloplastus pair 2o-7 6.15 Phylogenetic Tree of the Australian Orthocladiinae 208 209 V. NT]MER]CAL ANALYSIS 209 1. INTRODUCTION 2ro 2. 2IO 3. 4. 2LO vI. 2L4 References 2L7 SUMMARY After collecting widely in the wetter regions of South Australia and with the assistance of a small amount of material collecÈed in other parts of south eastern Australia, I have provided descriptions and biological details of 73 species in 3 subfamilies of Chironomidae (Diptera : Nema- tocera), Èhe Aphroteniinae, Diamesinae (tribe Heptagyini) and Ortho- cladiinae. Of these, the single member of the Aphroteniinae and both species of Heptagyini are new species, while two genera and forty-eight species of Orthocladiinae are new. I have examined the approach Èo taxonomy used by both the Mayr school and the Hennig school and also critically examined the methodology of Hennig's followers. Hennig's phylogenetic systematics has been adopted as the most suitable for the present work. A proposal has been made for indicating the level- of reliability of phylogenetic estimates of relationship, and its use illustrated by a consideraÈion of relationships within the Orthocladiinae, f.or which an incomplete phylogeny has been suggested.. \ A phenogram has been prepared using the program of Lance g Williams (L967) on a computer, relating all Australian species of Orthocladiinae. This has then been compared with the proposals for phylogeny- The implications of this work in relation to biogeographical problerns are briefly examined. ':i (i) ACKNOWLEDGEMENTS My supervisor, Dr. D. A. Duckhouse has greatly assisted me through discussion and criticism of this manuscript: for this and for his continued encouragement, I sincerely thank hím' I am grateful to Professor Andrewartha (recently retired) and to his successor, Mr. I. l'1. Thomas, for making available the facilities of the Zool-ogy DePartment. some collections of specimens were made on privately owned properties and for access to these I thank Dr. D. Hea1ey, Mrs' Fisher, Mr. I. Robertson and Mï. L. E. M. Salter. Special thanks are due to Mr. and Mrs. salter for their hospitality and to Mr. salter for regularly tending my collecting apparatus over a period of a year and a ha1f. I examined. many type specimens at c.s.I.R.O. Division of Entomolog!, Canberra and I especially thank Dr. D. H. Colless and his staff for their assistance in this work and also for the loan of material from the Australian National Insect Collection. I examined other type specimens aÈ the laboratories of the school of public Health and Tropical Medicíne, sydney university, and I am grateful to Professor D. J. Lee for the use of these facilities. For typing the drafts and the larger part of this manuscript I am indebted to my mother, M-rs. J. B. Hergstrom' I SECTION I INTRODUCTION The ísolated fauna of Australia and New Zealand poses many questions for systematists and biogeographers. Hennig (1960) outlined the problems posed by the then known Dipteran fauna of New Zealand and illustrated by example how phylogenetic systematics, his particular approach to the analysis of retationships, can provide a means of clarifying these questions and assist in answering some of them. Hennig r,vas prevented from anal-ysíng relationships more extensively by a lack of detailed information about the groups involved. However, he exhorted specialist workers to be aware of the type of question to be answered and to approach the analysis of relationships in a way that may provide the answers- Brundin (1966) in his investigations of southern representatives of the subfamilies Podonominae, Aphroteniinae and Diamesinae of the fainily chironomidae responded to Hennig's plea and used the methods of phylogenetic systematics to analyse the relationships between species of these sub- fanilies in depth. on Èhe basis of this analysis he came to far reactring biogeographical and geological conclusions. Brundin uses Hennig's principles with great flexibility, but if his extrapolaÈions were valid then this approach would be a useful and powerful tool for solving biogeographical Problems - If, however, his extrapolations are ulìlvarranted, care must also be taken in evaluating other works written in this style (e.g. Hirvenoja, Lg73). Considerable criticism has already been made both of Hennigrs phylogenetic systematics and of Brundin's application of these principles' I have approached the problem througlr- a consideration of some of these criticisms and by closely ana]-ysing some of Bru¡rdinrs proposals' In addition I have attempted to use Hennig's methods to construct a partial phylogeny of a second group, the Orthocladiinae' 2 A survey of the literaÈure (Brundi¡r, 1956; Edwards, 1931; Freeman' 1956, Ig5g, 1961) showed that of the remaining sr:bfanilies of Chironomid.ae, the Orthoclacliinae include genera with a distrilcution in the southern la¡d masses similar to that of tJ.e groups studied by Brundin. An analysis of the relationships within Orttrocladiinae could thus provide an j¡dication of the plausibility of Brundin's conclusions. For a nuniber of reasons it seemed wiser to choose Orthocladiinae for this analysis, rather than an entirely different familY. l. The characters of Orthoctadiinae would be somewhat si:nil-ar to those of the groups analysed by Brundin and thus migtr-t provide some clues about the accuracy of his evaluations of synapomorphies and synplesiomorphies- 2. In many cases Orthocladiinae are found in places where Podonomi¡rae, Aphroteniinae and Heptagryini would also be expected and thus while collecting Orthocladiinae, ne\¡¡ species of the latter might be taken which would provide a test of relationships proposed by Brundin. 3. A preliminary survey showed Orthocladiinae to be widespread and abundant in areas of South Australia not completely altered by human activity. So that proposals of relationships could be broadl-y based, I have co1lecÈed extensively and made investigations into tlre bioloqy (SECTION II) of as many species as possible. In proposing relationships I have also nade use of phenetic methods of analysis (SECTION V). 3 SBCTION II GENERAL BIOLOGY 1. INTRODUCTION Australian Chironomidae outside Chironominae have only rarely been collected on a large scale (Australian National Insect Collections,' Brundin 1966) and biological studies have been restrícted in scope (D. H. Edward, 1961; D. H. Edward and D. H. colfess, 1968; R. Jones, L973). At the outset of the present work almost nothing was known of the South Australian faqna and my objective has been to obtain a preliminary but comprehensive picture of ttre diversity of species associated wíth lotic habitats in this state. By collecting widely and using a variety of methods for trapping, extracting and rearing, I have attempted, to discover the immatures as well as the adults of as many species as possiJcle. In addition' observations have been made of such habitat preferences, tolerances and behaviour as might be useful in evaluating characters for phylogenetic purposes, Most of the collecting sites are, or were until recent clearing, surrounded by native woodland or forest comprising a dominant eucalypt and an associated understorey in which species diversity increases in the vatleys.
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