Peter S. Cranston

COULD A CLADOGRAM THIS SHORT HAVE ARISEN BY CHANCE ALONE?: ON PERMUTATION TESTS FOR CLADISTIC STRUCTURE

Abstract Absolute criteria for evaluating cladistic analyses are useful, not only because cladistic algorithms impose structure, but also because applications of cladistic results demand some assessment of the degree of corroboration of the cladogram. Here, a means of quantitative evaluation is presented based on tree length. The length of the most‐parsimonious tree reflects the degree to which the observed characters co‐vary such that a single tree topology can explain shared character states among the taxa. This “cladistic covariation” can be quantified by comparing the length of the most parsimonious tree for the observed data set to that found for data sets with random covariation of characters. A random data set is defined as one in which the original number of characters and their character states are maintained, but for each character, the states are randomly reassigned to the taxa. The cladistic permutation tail probability, PTP, is defined as the estimate of the proportion of times that a tree can be found as short or shorter than the original tree. Significant cladistic covariation exists if the PTP is less than a prescribed value, for example, 0.05. In case studies based on molecular and morphological data sets, application of the PTP shows that:

Botryocladius gen.n: a new transantarctic genus of orthocladiine midge (Diptera: Chironomidae)

A new genus of orthocladiine Chironomidae, Botryocladius (type species B. grapeth sp.n. from eastern Australia) is described and illustrated in all life history stages. All thirteen included species are described as new, six from eastern Australia (B. grapeth, B. brindabella, B. mdfrc, B. collessi, B. tasmania, B. australoalpinus), two from Western Australia (B. bibulmun and B. freemani), one from ephemeral streams in Australia (B. petrophilus) and four from Patagonian Argentina and Chile (B. edwardsi, B. glacialis, B. mapuche and B. tronador). All Australian species are known from at least pupal exuviae, most from adult males and several from larvae. In contrast, only B. edwardsi amongst Neotropical species is known from the adult male; all others are described from pupal exuviae. The immature stages are lotic in Australian permanent and temporary streams and Patagonian glacial streams and rivers, and lentic in Neotropical glacial‐fed and Australian subalpine lakes. Botryocladius appears to belong with a grouping centred on two formally undescribed taxa from Australia. The genus evidently demonstrates a vicariant distribution with at least two sister‐group relationships between South American and Australian taxa, providing a minimum dating for the clade of 38u2003Ma., with apparent absence from New Zealand indicating a maximum date of 80u2003Ma.

A systematic reappraisal of the Australian Aphroteniinae (Diptera: Chironomidae) with dating from vicariance biogeography

Abstract. The chironomid subfamily Aphroteniinae is revised for the Australian fauna. The larval‐based genus Anaphrotenia Brundin is synonymized with Aphroteniella Brundin, and Anaphrotenia lacustris Brundin with Aphroteniella filicomis Brundin, and all stages redescribed and illustrated. A second Australian species, Aphroteniella tenuicornis Brundin, has been reared and the larva is described and the pupa and male redescribed here.

Parapsectrocladius: a new genus of orthocladiine Chironomidae (Diptera) from Patagonia, the southern Andes

Parapsectrocladius is described in all life history stages as a new genus with type-species Cardiocladius acuminatus Edwards, for four species from Andean South America. The female, pupa and larva of P. acuminatus are described for the first time, from Chile; all stages of P. escondido Cranston & Anon Suarez, sp. n., are described from lentic patagonian Argentina, all stages of Parapsectrocladius reissi sp. n. from a pond in Valdivia, Chile, and P. longistylus sp. n. from males from several locations in south Chile and Argentina and the putatively associated female and pupa from southern Chile. The clade is Patagonian, with larvae occurring in Andean pools, ponds and lakes, including glacial-fed ones, and also in one lowland river in Chiloe. Previous suggested relationships to Cardiocladius Kieffer or Paratrissocladius Zavrel are rejected following phylogenetic analysis, with these placements suggested as due to adult male -based convergent character states. The immature stages, which show resemblance to Psectrocladius Kieffer and Rheocricotopus Thienemann & Harnisch, and the female contribute evidence that Parapsectrocladius forms the sister group to Psectrocladius, which appears monophyletic despite its subgeneric heterogeneity. Immediate next relations to Psectrocladius + Parapsectrocladius are postulated as (Rheocricotopus (both subgenera) + Paracricotopus) then (Nanocladius + Doncricotopus) at next remove.

The current distribution and geographical origin of the scale insect pest Ceroplastes sinensis (Hemiptera: Coccidae)

Ceroplastes sinensis Del Guercio is a pest of commercial citrus in Australia and has been recorded from New Zealand, southern Europe, northern Africa, North America and Mexico. Its distribution is mainly temperate and it is more restricted climatically than previously believed. Uncertainty concerning the place of origin of C. sinensis has thwarted previous attempts at biological control. In the present study, a novel and cost-effective approach using cladistic estimates is demonstrated to determine the place of its origin. Cladistically derived phylogenies are commonplace for many organisms, but the method has not been used before to predict the area of origin of a cosmopolitan pest. Cladistic analysis of the wax scales predicts a native range for C. sinensis within Central or South America. This prediction appears to have been confirmed by the recent discovery of heavily parasitized individuals of C. sinensis in Argentina. The cladistic analysis also allows prediction of the geographical origin of three other widespread wax scale speries: Ceroplastes ceriferus (Fabricius) may be of Neotropical origin, C. rubens Maskell and C. rusci (Linnaeus) are probably native to the Afrotropical region. The approach used has wide implications because the native areas of many other economically important pests are still unknown or uncertain.

The saline nuisance chironomid Carteronica longilobus (Diptera: Chironomidae): a systematic reappraisal

Abstract. Carteronica Kieffer and Nilodorum Kieffer sensu auctt. are synonymized with Kiefferulus Goetghebuer on the basis of cladistic analysis. An expanded diagnosis for all stages of Kiefferulus and a checklist of included species is given. K.longilobus comb.n. from Carteronica, a saline tolerant nuisance pest, is redescribed in all its life history stages. The following additional new combinations are proposed: K.brevibucca (Kieffer), K.brevipalpis (Kieffer), K.rugosum (Freeman), K.tainanus (Kieffer), K.umbraticola (Yamamoto). Many new characters from larval ventromental plates are illustrated and their use in phylo‐genetic analysis is discussed.

Conochironomus Freeman: an Afro-Australian Chironomini genus revised (Diptera: Chironomidae)

The chironomid genus Conochironomus is revised, with diagnoses provided for both sexes of adults and, for the first time, the immature stages. The Afrotropical genotype C.acutistilus Freeman and the newly described Australian species C.australiensis (for misidentified acutistilus in Australia), C.cygnus and C.kakadu are described in all stages. C.avicula Freeman is redescribed from the male, and C.deemingi is described as new from the male alone, both from the Afrotropical Region. The Australian species C.cervus is described as new, based on the pupa alone. Comments are made on the ecology and distribution. A previously little‐recognized ‘tropical’ Gondwanan biogeographic explanation for Conochironomus distribution is postulated.

The immature stages and phylogeny of imparipecten freeman, an australian endemic genus of wood‐mining chironomid (diptera)

The Australian endemic genus Imparipecten Freeman 1961 is revised, with descriptions newly provided for the immature stages. The single species, Imparipecten pictipes Freeman 1961, which mines immersed wood as a larva, is found in lotic waters in eastern Australia from northern New South Wales to Tasmania. A postulated phylogenetic relationship to Endochironomus Kieffer and Tribelos Townes (Freeman, 1961) is refuted by the female genitalia, the morphology of the immature stages and by parsimony analysis of combined life history data. The best substantiated phylogenetic hypothesis has Imparipecten lying within a grouping that includes Paratendipes Kieffer, Conochironomus Freeman and Skusel‐la Freeman ‐ a monophyletic grouping defined by the possession of a larval six‐segmented antenna with Lauterborn organs alternate on the second and third segments.

THE UNDERSIDE OF CLADISTICS-IN THE LAND OF OZ

The southern temperate fauna and flora, not least that of Australia, is a stimulating testing ground for phylogenetic and biogeographic ideas. Had Hooker (1860) stuck to his earlier ( 1853) explanation of a once widespread austral flora on a more contiguous landmass there might have been little controversy. However, Hooker’s second thoughts of a colonial invasion prevailed, culminating in Darlington’s (1965) dispersalist opus. When Hennig (1960) and Brundin (1966) came to the rescue, the austral biota was again central in demonstrations of how to elucidate phylogenetic and biogeographic relationships. The IXth meeting of the Willi Hennig Society, held in Australia’s capital, Canberra, allowed us, amongst other matters, to inspect recent progress in these areas. The inaugural southern hemisphere meeting of our Society, held between 23 and 27 August 1990, was organized by Ebbe Nielsen, Judy West, Pauline Ladiges, Robert Raven and this year’s president, Chris Humphries. The venue allowed many Australians to participate, although disappointingly few New Zealanders availed themselves of the opportunity to proselytize. Other gondwanan areas were also represented (Jorge Crisci and Pablo Goloboff-South America; Peter Linder South Africa). Some 200 in total attended, with the northern antipodean component enhanced by many trans-Pacific migrants, compensating for a paucity of oceanic islanders and continental Europeans, Danes excepted. Two of the themes reflected the location: Pauline Ladiges and Chris Humphries organized a full day on “Austral biogeography”, with the following day being devoted to “Austral flora and fauna: systematics and evolution” under the supervision of Mike Crisp and Robert Raven. The demarcation between these two sessions was indistinct. since most contributors in the Flora and Fauna session acknowledged contemporary biogeographic thinking. This contrasted somewhat with several papers presented in the biogeography session, where heresies such as biogeography without areas competed with biogeography without cladograms. Joel Cracraft (“Deciphering historical signals in the distribution of organisms”) may have rediscovered site classification and Dick VaneWright (whose paper was presented in absentia by Ian Kitching) transcended Wallace’s line (and cladograms) in assessing the relationships of Sulawesian butterflies. New Zealander Ian Henderson gave a “head-in-the-lion’s_mouth” panbiogeographic virw of “Biogeography without areas”, raising the important issue of the fuzzy relationship between point distribution records and areas, and then delving into metaphysics with “nodal ctntres of form-making”. More conventional case-history studies came from Peter Weston and Mike Crisp (“Trans-Pacific cladistic patterns in the Proteaceae”), Allan Rodrigo and Robert