Nothofagus Biogeography Revisited with Special Emphasis on the Enigmatic Distribution of Subgenus Brassospora in New Caledonia

Dispersals versus vicariance events and the presence of subgenus Brassospora in New Caledonia are two riddles of Nothofagus biogeography, a genus also distributed in New Guinea, New Zealand, South America, Southeast Australia, and Tasmania. Within a cladistic framework using the software COMPONENT 2.0, we demonstrate that most parsimonious area cladograms (areagrams) sensu cladistic biogeography need not always be the most plausible explanation nor reflect alternative geological hypotheses. The most parsimonious Nothofagus history sensu historical biogeography is reconstructed where a minimum of dispersed taxa is hypothesized and vicariance events are identified. A fully resolved well-established Nothofagus phylogeny was reconciled with three geological hypotheses (geograms) of East Gondwana break-up: (a) the conventional view, (b) an Australian—New Caledonian relationship, and (c) a biotic interchange between New Guinea and New Caledonia. Fossils determined to subgenus were optimized to the predicted lineages in the reconciled tree. Due to extensive extinctions, a maximum of three vicariance events are inferred, all being basal in the subgenera, an indication of subgeneric diversification prior to the break-up of Gondwana. Two taxa, N. gunnii and N. menziesii, are hypothesized as being long-distance dispersed. The most parsimonious solution suggests a close relationship between New Guinea and New Caledonia, supporting a Brassospora colonization route, but this hypothesis fails to predict numerous extinct lineages observed in the fossil record and thus must be rejected. The traditional break-up sequence of Gondwana is not the most parsimonious solution, indicating one incongruent node, but causes no overall incongruence with the fossil record. Considering all parameters, the occurrence of Brassospora in New Caledonia is most parsimoniously explained as a single colonization event from New Zealand where the subgenus subsequently went extinct in the Pliocene.     

Local or global? Biogeography of some primitive Lepidoptera in New Zealand

Abstract

Distribution maps are presented for 30 species of small forest moths representing two genera of Micropterigidae and one genus of Mnesarchaeidae. Evolutionary patterns are interpreted by means of the panbiogeographic method in order to seek a relationship between dispersal patterns within New Zealand and the overseas links of the taxa in question. It is established that different evolutionary lines indeed exhibit different geographic dispersal patterns within New Zealand. Moreover, certain characteristics of these patterns can be attributed to the overseas affinities of the groups, i.e., groups of organisms with New Caledonian or Australian affinities have a western disjunct pattern, whereas those with circum-Pacific affinities have a different pattern in which the taxa occupying a basal position in the phylogeny are to be found in the eastern South Island. However, both groups have overlapping centres of diversity, particularly in NW Nelson.     

First phylogenetic and biogeographical study of the southern bluebells (Wahlenbergia, Campanulaceae)

Wahlenbergia is a largely southern hemisphere genus of at least 260 species; within Campanulaceae only Campanula is larger. This first phylogeny of Wahlenbergia was reconstructed using about 20% of the 260 species in the genus based on the nuclear ribosomal ITS marker and the chloroplast trnL-F marker with samples from South Africa, Europe, Australia and New Zealand. Wahlenbergia was confirmed to be non-monophyletic, though most of the species form a clade. Our tree topology and date estimates indicate that Wahlenbergia diverged in South Africa about 29.6 mya, then dispersed to Australasia about 4.8 mya, thus indicating the radiation of Wahlenbergia occurred relatively recently. Radiations occurred in both of these main centres; there are currently about 170 species in South Africa and 45 species and subspecies in Australasia. New Zealand species comprise two clades, both rooted within the Australasian clade. We thus propose two dispersals from Australia to New Zealand, one leading to a radiation of species with the rhizomatous herbaceous growth form ca. 1.6 mya, and the other leading to a radiation of species with the radicate growth form 0.7 mya. Dispersals from Australia to New Zealand match the expected direction, following the west wind drift and ocean currents. The herbaceous growth form was shown to be ancestral for the genus as a whole, and polyploidy has been a mechanism of the evolution of the genus in Australasia.     

The biogeography of the austral, subalpine genus Ourisia (Plantaginaceae) based on molecular phylogenetic evidence: South American origin and dispersal to New Zealand and Tasmania

Molecular phylogenetic analyses of 26 of the 28 species of Ourisia , including eight of ten subspecies and two purported natural hybrids, are presented and used to examine the biogeography of the genus, which is distributed in subalpine to alpine habitats of South America, New Zealand and Tasmania. Gondwanan vicariance, often cited as the cause of this classic austral biogeographical pattern, was rejected by parametric bootstrapping of our combined dataset. Alternatively, various lines of evidence are presented in favour of a South American origin of Ourisia and subsequent dispersal to Australasia. Specifically, the genus likely arose in the Andes of central Chile and spread to southern Chile and Argentina, to the north-central Andes, and finally to Tasmania and New Zealand. The ancestor of the New Zealand species probably first arrived on the South Island, where the New Zealand species of Ourisia are most diverse, and migrated to the North and Stewart Islands. Because the Tasmanian and New Zealand species are sister to one another, the direction of dispersal between these two areas is equivocal. These results agree with other molecular phylogenetic studies that show that past dispersal between southern hemisphere continents has played an important role in the evolutionary history of many high-elevation austral plants. Our data also show that within South America, many of the geographical barriers (with the exception of the Atacama Desert) that have played a role in the evolution of other plant groups have not affected Ourisia species. Within New Zealand, the phylogeny and biogeography of species of Ourisia coincide with the geological history of the country and patterns of other alpine plants. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 479–513.     

The biogeography of Gunnera L.: vicariance and dispersal

Aim The genus Gunnera is distributed in South America, Africa and the Australasian region, a few species reaching Hawaii and southern Mexico in the North. A cladogram was used to (1) discuss the biogeography of Gunnera and (2) subsequently compare this biogeographical pattern with the geological history of continents and the patterns reported for other Southern Hemisphere organisms. Location Africa, northern South America, southern South America, Tasmania, New Zealand, New Guinea/Malaya, Hawaii, North America, Antarctica. Methods A phylogenetic analysis of twenty‐six species of Gunnera combining morphological characters and new as well as published sequences of the ITS region, rbcL and the rps16 intron, was used to interpret the biogeographical patterns in Gunnera. Vicariance was applied in the first place and dispersal was only assumed as a second best explanation. Results The Uruguayan/Brazilian Gunnera herteri Osten (subgenus Ostenigunnera Mattfeld) is sister to the rest of the genus, followed sequentially upwards by the African G. perpensa L. (subgenus Gunnera), in turn sister to all other, American and Australasian, species. These are divided into two clades, one containing American/Hawaiian species, the other containing all Australasian species. Within the Australasian clade, G. macrophylla Blume (subgenus Pseudogunnera Schindler), occurring in New Guinea and Malaya, is sister to a clade including the species from New Zealand and Tasmania (subgenus Milligania Schindler). The southern South American subgenus Misandra Schindler is sister to a clade containing the remaining American, as well as the Hawaiian species (subgenus Panke Schindler). Within subgenus Panke, G. mexicana Brandegee, the only North American species in the genus, is sister to a clade wherein the Hawaiian species are basal to all south and central American taxa. Main conclusions According to the cladogram, South America appears in two places, suggesting an historical explanation for northern South America to be separate from southern South America. Following a well‐known biogeographical pattern of vicariance, Africa is the sister area to the combined southern South America/Australasian clade. Within the Australasian clade, New Zealand is more closely related to New Guinea/Malaya than to southern South America, a pattern found in other plant cladograms, contradictory to some of the patterns supported by animal clades and by the geological hypothesis, respectively. The position of the Tasmanian G. cordifolia, nested within the New Zealand clade indicates dispersal of this species to Tasmania. The position of G. mexicana, the only North American species, as sister to the remaining species of subgenus Panke together with the subsequent sister relation between Hawaii and southern South America, may reflect a North American origin of Panke and a recolonization of South America from the north. This is in agreement with the early North American fossil record of Gunnera and the apparent young age of the South American clade.     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

Molecular phylogeny ofNothofagus (Nothofagaceae) based on theatpB-rbcL intergenic spacer of the chloroplast DNA

The genusNothofagus is distributed in the Southern Hemisphere from South America to Oceania, and its distribution has been assumed to be formed by continental drift by means of Gondwana break-up during the Mesozoic era. The phylogeny of the genus was elucidated by the sequences ofatpB-rbcL intergenic spacer of cpDNA for the better understanding of its evolution and biogeography. The phylogeny ofNothofagus corresponded completely to the pollen morphology which recognizes four pollen types in extant species, and agrees well with the taxonomic system of Hill and Read (1991) although there, the subgenusNothofagus showed in unresolved polytomy. The topology of the phylogenetic tree reveals that subgenusLophozonia was derived first, and thenFuscospora, Nothofagus andBrassospora. Species from South America and New Zealand were assigned to each cluster according to their pollen morphology. Therefore, diversification ofNothofagus should have already proceeded at the subgenus level before the completion of Gondwana break-up Tropical species distributed in New Guinea and New Caledonia whose evolutionary history has been controversial were revealed to be a derived group. All five New Caledonian species formed a monophyletic group with very few sequence divergences in the intergenic spacer of cpDNA, thus showing rapid adaptive radiation in the island. Evolutionary trends of several morphological traits ofNothofagus are discussed. The evolution of valve number of cupules, number of nuts per cupule, and habit of leaf-fall (evergreen or deciduous) which are diversified in the genus, were revealed as having occurred several times as the result of convergence.     

Phylogenetics of the australasian mudfishes: evolution of an eel-like body plan

The mudfish genus Neochanna (Osmeriformes: Galaxiidae) contains six species that exhibit varying degrees of morphological and ecological adaptation to life in swampy conditions. Here, we present the first molecular phylogenetic analysis (16S rRNA+cytochrome b; 1681bp) of the entire genus to (1) test for monophyly of Australian and New Zealand taxa and (2) elucidate morphological character evolution. In addition, we analyse a matrix of 21 morphological characters to test for congruence between mitochondrial DNA and morphology, and to examine total evidence under a Bayesian framework. Molecular data indicate that the diadromous Tasmanian mudfish, N. cleaveri, is sister to a clade of five non-diadromous New Zealand mudfishes. Mapping of morphological characters onto the molecular phylogeny suggests an evolutionary transition from a plesiomorphic "stream" galaxiid morphotype to a more specialised "anguilliform" galaxiid morphotype. Pelvic fins have become increasingly reduced and dorsal, anal, and caudal fins are increasingly confluent. Associated with these changes are elongated nostrils, reduced eyes, and increased anterior cranial ossification. Morphological and total evidence analyses yield similar or identical topologies, respectively. The phylogenetic distribution of diadromy in Neochanna is consistent with a single loss of this character state in New Zealand. However, the strong sister relationship (3.6% divergent; 100% bootstrap support) detected between non-diadromous N. burrowsius (South Island, NZ) and N. rekohua (Chatham Islands) indicates geologically recent dispersal across 850km of ocean. Diadromy may therefore have been retained in the common ancestor of these two mudfish species, and subsequently lost from both lineages.     

Uncommon zoogeographical connections in the subgenus <Emphasis Type="Italic">Exopalpiger</Emphasis> Schultze of the genus <Emphasis Type="Italic">Ixodes</Emphasis> Latreille (Acari,Ixodidae)

The species composition of the subgenus Exopalpiger Schulze, 1935 (genus Ixodes Latreille, 1795) and species ranges are considered. Altitude and biotopic preferences and host-parasite relations of species are analyzed. A hypothesis explaining the palaeogenesis of the disjunctive distribution (Europe; South Africa; north and east of South America; New Guinea, southern Australia, New Zealand, and Tasmania) of the subgenus Exopalpiger is proposed.     

Cryptic dioecy and leaky dioecy in endemic species of Dombeya (Sterculiaceae) on La Re´union

The high frequency of dioecy on oceanic islands such as Hawaii and New Zealand has attracted a great deal of attention from plant evolutionary biologists. One reason suggested for the high prevalence of dioecy on oceanic islands is that taxa considered truly dioecious may have occasional hermaphrodite flowers, i.e., show leaky dioecy. In this study, we quantified the presence and distribution of leaky dioecy in a group of congeneric endemic species of the genus Dombeya (Sterculiaceae) on La Re´union island (Indian Ocean). All eight species show cryptic dioecy. Five species show strict dioecy and three species show leaky dioecy due to the presence of male trees that set fruit. Species with strict dioecy and large populations tend to occur in mid- to high-altitude moist tropical cloud forest, whereas species in smaller populations at lower altitude and in semidry tropical forest tend to show leaky dioecy. Two reasons for this differential distribution of strict dioecy and leaky dioecy are discussed. First, environmental variation along the altitudinal gradient, biotic and/or abiotic, may influence the breeding system. Second, leaky dioecy may be favored in lowland populations due to the small size and disturbed nature of such populations.     

中国-喜玛拉雅特有属——蓝钟花属的分类修订

Cyananthus Wallich ex Bentham, the only genus of Campanulaceae with superior ovary, is revised to clarify infrageneric relationships and phylogeny of the genus. Evidence obtained from the comparative gross morphology, anatomy, palynology, and karyomorpho-logy recommends a new infrageneric classification of the genus, recognizing 23 species, belonging to two subgenera, four sections and four subsections. One subgenus(Subgen. Mi-cranthus), one section(Sect. Suffruticulosi) and two subsections(Subsect. Flavi and Sub-sect. Lichiangenses)are described as new taxa. New combinations at sectional (Sect. Annui) and subsectional(Subsect. Stenolobi) ranks are also proposed. The genus Cyananthus is strictly distributed in the high mountains of China(Xizang, Yunnan and Sichuan), extending to Bhutan, Nepal and India (Kumaon-Garhwal, Assam and Sikkim), with altitudinal ranges from 2500 ~ 5300 m. It is observed that 13 species are endemic to SW China and only three species are endemic to the Himalayas( two species in Ne     

The evolution of dioecy, heterodichogamy, and labile sex expression in Acer

The northern hemisphere tree genus Acer comprises 124 species, most of them monoecious, but 13 dioecious. The monoecious species flower dichogamously, duodichogamously (male, female, male), or in some species heterodichogamously (two morphs that each produce male and female flowers but at reciprocal times). Dioecious species cannot engage in these temporal strategies. Using a phylogeny for 66 species and subspecies obtained from 6600 nucleotides of chloroplast introns, spacers, and a protein-coding gene, we address the hypothesis (Pannell and Verdú, Evolution 60: 660-673. 2006) that dioecy evolved from heterodichogamy. This hypothesis was based on phylogenetic analyses (Gleiser and Verdú, New Phytol. 165: 633-640. 2005) that included 29-39 species of Acer coded for five sexual strategies (duodichogamous monoecy, heterodichogamous androdioecy, heterodichogamous trioecy, dichogamous subdioecy, and dioecy) treated as ordered states or as a single continuous variable. When reviewing the basis for these scorings, we found errors that together with the small taxon sample, cast doubt on the earlier inferences. Based on published studies, we coded 56 species of Acer for four sexual strategies, dioecy, monoecy with dichogamous or duodichogamous flowering, monoecy with heterodichogamous flowering, or labile sex expression, in which individuals reverse their sex allocation depending on environment-phenotype interactions. Using Bayesian character mapping, we infer an average of 15 transformations, a third of them involving changes from monoecy-cum-duodichogamy to dioecy; less frequent were changes from this strategy to heterodichogamy; dioecy rarely reverts to other sexual systems. Contra the earlier inferences, we found no switches between heterodichogamy and dioecy. Unexpectedly, most of the species with labile sex expression are grouped together, suggesting that phenotypic plasticity in Acer may be a heritable sexual strategy. Because of the complex flowering phenologies, however, a concern remains that monoecy in Acer might not always be distinguishable from labile sex expression, which needs to be addressed by long-term monitoring of monoecious trees. The 13 dioecious species occur in phylogenetically disparate clades that date back to the Late Eocene and Oligocene, judging from a fossil-calibrated relaxed molecular clock.     

Chloroplast DNA diversity of Hieracium Pilosella (Asteraceae) introduced to New Zealand: reticulation, hybridization, and invasion

The European hawkweed Hieracium pilosella is a successful invader and a troublesome weed in New Zealand. The systematics of the genus Hieracium is extremely complex and contentious, probably due to recent speciation, hybridization, polyploidy, and diverse reproductive strategies. In the first chloroplast DNA survey of the group, we sequenced 285 plants (including H. pilosella and 12 other species of subgenus Pilosella) from New Zealand and Europe for 900 bp of trnL-trnF. Eleven haplotypes were identified with much sharing among species. Three haplotypes (A, D, G) were found in seven, three, and four species, respectively, but two species (H. lactucella and H. auricula) had single, private haplotypes. Our cpDNA data for subgenus Pilosella are consistent with the group's having incomplete lineage sorting and/or recent reticulate evolution. Six haplotypes were identified in H. pilosella, four of these unique to this taxon in our sample. In New Zealand, haplotype A was common and occurred in plants of different ploidy (i.e., 4×, 5×, 6×), whereas haplotypes C, B, and M were restricted to 4×, 5×, and 6× plants, respectively. The distribution of haplotype variation suggests that some or all of the H. pilosella seeds accidentally introduced into New Zealand probably came from east Europe rather than the United Kingdom and that a minimum of four lineages were introduced. Within New Zealand, hybridization of H. pilosella with a related taxon (probably H. praealtum) has occurred at least three times, involving both obligate sexual tetraploids and facultative apomictic pentaploids of H. pilosella.     

Patterns of long-distance dispersal in Tiquilia subg. Tiquilia (Boraginaceae): implications for the origins of amphitropical disjuncts and Galapagos Islands endemics

Plant biogeographers have long argued whether plant disjunctions result from vicariance or dispersal. One of the classic patterns of plant disjunction involves New World amphitropical disjuncts, as exemplified by Tiquilia subg. Tiquilia (Boraginaceae). Subgenus Tiquilia forms a heterogeneous group of ~20 species that is amphitropically distributed in the deserts of North and South America, with four taxa endemic to the Galápagos Islands. The current study reconstructs the biogeographic history of subg. Tiquilia in order to explore the origins of New World amphitropical disjunction and of Galápagos endemism. A strongly supported phylogeny of the subgenus is estimated using sequence data from matK, ndhF, rps16, ITS, and waxy. Biogeographic analyses using combined and individual marker data sets reveal a complex history of long-distance dispersal in subg. Tiquilia. Biogeographic reconstructions imply a North American origin of the subgenus and its three major lineages and require at least four long-distance dispersal events to explain its current distribution. The South American taxa of subg. Tiquilia result from three independent and nonsimultaneous colonization events, while the monophyly and continental origins of the Galápagos endemics are unresolved. This study contributes to a growing body of evidence that intercontinental dispersal has been more common than previously realized.     

South Pacific biogeography,tectonic calibration,and pre‐drift tectonics: cladogenesis in Abrotanella (Asteraceae)

Abrotanella is the basal genus in the large tribe Senecioneae (Asteraceae) and has a disjunct distribution in Australasia and South America. A recent molecular phylogeny of the genus was used to investigate whether the main biogeographical patterns in the group could be related to the region's tectonic history in a coherent way. The phylogenetic/biogeographical breaks and overlaps in the genus imply a series of vicariance and range expansion events. Each of these can be related to one of the main tectonic events in the region, including assembly of the New Zealand terranes, crustal extension, and magmatism in Gondwana that preceded seafloor spreading, opening of the Tasman and Pacific basins, and transcurrent movement on the New Zealand Alpine fault. The coincident sequence indicates that pre‐drift tectonics and magmatism have been more important for the origin of trans‐Tasman and trans‐Pacific groups than the final rifting of Gondwana that led to their disjunction. For example, during the pre‐drift phase of break‐up, the Whitsunday volcanic province of Australia and the Median Batholith of New Zealand formed a large, active igneous belt. Its distribution is aligned with the break between New Zealand–south‐eastern Australia clades, and New Zealand–New Guinea clades. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

The evolution of reproductive systems and sex-determining mechanisms within rumex (polygonaceae) inferred from nuclear and chloroplastidial sequence data

The genus Rumex includes hermaphroditic, polygamous, gynodioecious, monoecious, and dioecious species, with the dioecious species being represented by different sex-determining mechanisms and sex-chromosome systems. Therefore, this genus represents an exceptional case study to test several hypotheses concerning the evolution of both mating systems and the genetic control of sex determination in plants. Here, we compare nuclear intergenic transcribed spacers and chloroplast intergenic sequences of 31 species of Rumex. Our phylogenetic analysis supports a systematic classification of the genus, which differs from that currently accepted. In contrast to the current view, this new phylogeny suggests a common origin for all Eurasian and American dioecious species of Rumex, with gynodioecy as an intermediate state on the way to dioecy. Our results support the contention that sex determination based on the balance between the number of X chromosomes and the number of autosomes (X/A balance) has evolved secondarily from male-determining Y mechanisms and that multiple sex-chromosome systems, XX/XY1Y2, were derived twice from an XX/XY system. The resulting phylogeny is consistent with a classification of Rumex species according to their basic chromosome number, implying that the evolution of Rumex species might have followed a process of chromosomal reduction from x = 10 toward x = 7 through intermediate stages (x = 9 and x = 8).     

A revision of the South American taxa Elnoretta Alexander, Euvaldiviana Alexander and Valdiviana Alexander (Diptera: Tipulidae), with a discussion on their phylogeny

Abstract The South American tipulid taxa Elnoretta, Euvaldiviana and Valdiviana are revised. Type material of all described species was examined. Euvaldiviana is raised from subgeneric to generic rank. The synonymy of Valdiviana synempora and V. neuquenensis is established. Details of the genitalia of the five recognized species as well as external characters are illustrated. A phylogeny is presented of a monophyletic group containing Elnoretta, Euvaldiviana and Valdiviana together with the genera Acracantha (Australia) and Austrotipula (New Zealand).     

Welcome back New Zealand: regional biogeography and Gondwanan origin of three endemic genera of mite harvestmen (Arachnida, Opiliones, Cyphophthalmi)

Aim Biogeographers have long been intrigued by New Zealand’s biota due to its unique combination of typical ‘continental’ and ‘island’ characteristics. The New Zealand plateau rifted from the former supercontinent Gondwana c. 80 Ma, and has been isolated from other land masses ever since. Therefore, the flora and fauna of New Zealand include lineages that are Gondwanan in origin, but also include a very large number of endemics. In this study, we analyse the evolutionary relationships of three genera of mite harvestmen (Arachnida, Opiliones, Cyphophthalmi) endemic to New Zealand, both to each other and to their temperate Gondwanan relatives found in Australia, Chile, Sri Lanka and South Africa. Location New Zealand (North Island, South Island and Stewart Island). Methods A total of 94 specimens of the family Pettalidae in the suborder Cyphophthalmi were studied, representing 31 species and subspecies belonging to three endemic genera from New Zealand (Aoraki, Neopurcellia and Rakaia) plus six other members of the family from Chile, South Africa, Sri Lanka and Australia. The phylogeny of these taxa was constructed using morphological and molecular data from five nuclear and mitochondrial genes (18S rRNA, 28S rRNA, 16S rRNA, cytochrome c oxidase subunit I and histone H3, totalling c. 5 kb), which were analysed using dynamic as well as static homology under a variety of optimality criteria. Results The results showed that each of the three New Zealand cyphophthalmid genera is monophyletic, and occupies a distinct geographical region within the archipelago, grossly corresponding to palaeogeographical regions. All three genera of New Zealand mite harvestmen fall within the family Pettalidae with a classic temperate Gondwanan distribution, but they do not render any other genera paraphyletic. Main conclusions Our study shows that New Zealand’s three genera of mite harvestmen are unequivocally related to other members of the temperate Gondwanan family Pettalidae. Monophyly of each genus contradicts the idea of recent dispersal to New Zealand. Within New Zealand, striking biogeographical patterns are apparent in this group of short‐range endemics, particularly in the South Island. These patterns are interpreted in the light of New Zealand’s turbulent geological history and present‐day patterns of forest cover.     

A molecular phylogeny of the Littorininae (Gastropoda: Littorinidae): unequal evolutionary rates,morphological parallelism,and biogeography of the Southern Ocean

A molecular phylogeny is presented for the subfamily Littorininae (including representatives of all subgeneric taxa and all members of a group of southern-temperate species formerly classified as 'Nodilittorina'), based on sequence data from two nuclear (18S rRNA, 28S rRNA) and two mitochondrial (12S rRNA, CO1) genes. The phylogeny shows considerable disagreement with earlier hypotheses derived from morphological data. In particular, 'Nodilittorina' is polyphyletic and is here divided into four genera (Echinolittorina, Austrolittorina, Afrolittorina new genus, and the monotypic Nodilittorina s.s.). The phylogenetic relationships of 'Littorina' striata have been controversial and it is here transferred to the genus Tectarius, a surprising relationship for which there is little morphological support. The relationships of the enigmatic Mainwaringia remain poorly resolved, but it is not a basal member of the subfamily. The two living species of Mainwaringia are remarkable for a greatly elevated rate of evolution in all four genes examined; it is suggested that this may be connected with their protandrous hermaphroditism, which is unique in the family. The molecular phylogeny provides a new framework for the adaptive radiation of the Littorininae, showing more frequent shifts between habitats and climatic regimes than previously suspected, and striking parallelism of morphological characters. The fossil record of littorinids is poor, but ages of clades are estimated using a calibration based on a Lower Eocene age of the genus Littoraria. Using these estimates, the antitropical distribution of Littorina and Afrolittorina is an ancient pattern of possibly Cretaceous age. The five members of Austrolittorina show a Gondwanan distribution in Australia, New Zealand, and South America. Based on the morphological uniformity within this clade, relatively recent (Plio-Pleistocene) trans-Pacific dispersal events seemed a likely explanation, as proposed for numerous other congeneric marine taxa. However, molecular estimation of ages of divergence suggest an initial vicariance between Australian and South American lineages at 40-73Ma, contemporary with the later stages of fragmentation of the Gondwanan supercontinent, followed by more recent (but still mid-Cenozoic) dispersal events across the Tasman Sea and the Pacific Ocean. Afrolittorina is another Cretaceous clade, now restricted to southern Africa and southern Australia, but divergence between these lineages (29-55Ma) post-dates Gondwanan fragmentation. Within both Austrolittorina and Afrolittorina all sister-species divergences are estimated to fall in the range 10-47Ma, so that there is no evidence for speciation events in the Plio-Pleistocene.     

Revision of Dennyus (Collodennyus) lice (Phthiraptera: Menoponidae) from swiftlets, with descriptions of new taxa and a comparison of host–parasite relationships

Lice of the subgenus Dennyus ( Collodennyus ) are host specific, permanent parasites of swiftlets (Aves: Apodidae). As a prelude to a test of the hypothesis that these lice have cospeciated with their hosts, we revise the taxonomy of the subgenus, redescribing the seven previously recognized species, and adding thirteen new species and three new subspecies. All twenty-three of these louse taxa are found on swiftlets (Apodiformes: Apodidae), with four from hosts of the genus Collocalia , eighteen from Aerodramus , and one from Hydrochous . Successful identification is associated in most cases with females; males are only tenuously separable. A complete host–parasite list for the subgenus Collodennyus is provided, as well as a key for the identification of these taxa. Limited morphological variation within the subgenus has prevented ready extraction of discrete characters for cladistic analysis. In the absence of such characters, a cluster analysis of female and male lice is presented. Comparison of a dendrogram for Dennyus ( Collodennyus ) with a molecular phylogeny for the swiftlet hosts suggests that the history of the swiftlet–louse association has been complex, including episodes of host switching and independent speciation by the lice.