Best linear unbiased prediction of host-range of the facultative parasite Colletotrichum gloeosporioides f. sp. salsolae, a potential biological control agent of Russian thistle

Russian thistle or tumbleweed (Salsola tragus L.) is an introduced invasive weed in N. America. It is widely distributed in the US and is a target of biological control efforts.The fungus Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. in Penz. f. sp. salsolae (CGS) is a facultative parasite under evaluation for classical biological control of this weed. Host-range tests were conducted with CGS in quarantine to determine whether the fungus is safe to release in N. America. Ninetytwo accessions were analyzed from 19 families: Aizoaceae, Alliaceae, Amaranthaceae, Apiaceae, Asteraceae, Brassicaceae, Cactaceae, Campanulaceae, Chenopodiaceae, Cucurbitaceae, Cupressaceae, Fabaceae, Malvaceae, Nyctaginaceae, Phytolaccaceae, Poaceae, Polygonaceae, Sarcobataceae, and Solanaceae and 10 tribes within the Chenopodiaceae: Atripliceae, Beteae, Camphorosmeae, Chenopodieae, Corispermeae, Halopepideae, Polycnemeae, Salicornieae, Salsoleae, and Suaedeae. These included 62 genera and 120 species. To facilitate interpretation of results, disease reaction data were combined with a relationship matrix derived from internal transcribed spacer DNA sequences and analyzed with mixed model equations to produce Best Linear Unbiased Predictors (BLUPs) for each species. Twenty-nine species (30 accessions) from seven closely-related Chenopodiaceae tribes had significant levels of disease severity as indicated by BLUPs, compared to six species determined to be susceptible with least squares means estimates. The 29 susceptible species were: 1 from Atripliceae, 4 from Camphorosmeae, 1 from Halopepideae, 2 from Polycnemeae, 6 from Salicornieae, 8 from Salsolae, and 7 from Suaedeae. Most species in the genus Salsola, which are all introduced and weedy, were very susceptible and damaged by CGS. Statistical comparisons and contrasts of BLUPs indicated that these Salsola species were significantly more susceptible than non-target species, including 15 species from relatives in the closely-related genera Bassia (=Kochia), Nitrophila, Salicornia, Sarcocornia, and Suaeda. Of the 29 susceptible species, 10 native or commercially important species in N. America were identified as needing additional tests to determine the extent of any damage caused by infection.     

Towards a species level tree of the globally diverse genus Chenopodium (Chenopodiaceae)

Chenopodium is a large and morphologically variable genus of annual and perennial herbs with an almost global distribution. All subgenera and most sections of Chenopodium were sampled along with other genera of Chenopodieae, Atripliceae and Axyrideae across the subfamily Chenopodioideae (Chenopodiaceae), totalling to 140 taxa. Using Maximum parsimony and Bayesian analyses of the non-coding trnL-F (cpDNA) and nuclear ITS regions, we provide a comprehensive picture of relationships of Chenopodium sensu lato. The genus as broadly classified is highly paraphyletic within Chenopodioideae, consisting of five major clades. Compared to previous studies, the tribe Dysphanieae with three genera Dysphania, Teloxys and Suckleya (comprising the aromatic species of Chenopodium s.l.) is now shown to form one of the early branches in the tree of Chenopodioideae. We further recognize the tribe Spinacieae to include Spinacia, several species of Chenopodium, and the genera Monolepis and Scleroblitum. The Chenopodium rubrum and the Ch. murale-clades were newly discovered as distinct major lineages but their relationships within Chenopodioideae will need further evaluation. Based on our results, we suggest the delimitation of Chenopodium to include Einadia and Rhagodia because these are part of the crown group composed of species of subg. Chenopodium that appear sister to the Atripliceae. The tetraploid crops such as Ch. berlandieri subsp. nuttalliae and Ch. quinoa also belong to Chenopodium sensu stricto. Trees derived from trnL-F and ITS were incongruent within this shallow crown group clade. Possible biological causes are discussed, including allopolyploidization.     

Origin,Differentiation, and Geographic Distribution of the Chenopodiaceae

Numbers of species and genera,endemic genera,extant primitive genera,relationship and distribution patterns of presently living Chenopodiaceae(two subfamilies,12 tribes,and 118 genera)are analyzed and compared for eight distributional areas,namely central Asia,Europe,the Mediterranean region,Africa,North America,South America, Australia and East Asia． The Central Asia,where the number of genera and diversity of taxa are greater than in other areas,appears to be the center of distribution of extant Chenopodiaceae．North America and Australia are two secondary centers of distribution． Eurasia has 11 tribes out of the 12,a total of 70 genera of extant chenopodiaceous plants,and it contains the most primitive genera of every tribe． Archiatriplex of Atripliceae,Hablitzia of Hablitzeae,Corispermum of Corispermeae,Camphorosma of Camphorosmaea,Kalidium of Salicornieae,Polecnemum of Polycnemeae,Alexandra of Suaedeae,and Nanophyton of Salsoleae,are all found in Eurasia,The Beteae is an Eurasian endemic tribe,demonstrating the antiquity of the Chenopodiaceae flora of Eurasia．Hence,Eurasia is likely the place of origin of chenopodiaceous plants． The presence of chenopodiaceous plants is correlated with an arid climate．During the Cretaceous Period,most places of the continent of Eurasia were occupied by the ancient precursor to the Mediterranean,the Tethys Sea．At that time the area of the Tethys Sea had a dry and warm climate．Therefore,primitive Chenopodiaceae were likely present on the beaches of this ancient land．This arid climatic condition resulted in differentiation of the tribes Chenopodieae,Atripliceae,Comphorosmeae,Salicornieae,etc．,the main primitive tribes of the subfamily Cyclolobeae. Then following continental drift and the Laurasian and Gondwanan disintegration, the Chenopodiaceae were brought to every continent to propagate and develop, and experience the vicissitudes of climates, forming the main characteristics and distribution patterns of recent continental floras. The tribes Atripliceae, Chenopodieae, Camphorosmeae, and Salicornieae of recent Chenopodiaceae in Eurasia, North America, South America, southern Africa, and Australia all became strongly differentiated. However, Australia and South America, have no genera of Spirolobeae except for a few maritime Suaeda species. The Salsoleae and Suaedeae have not arrived in Australia and South America, which indicates that the subfamily Spirolobeae developed in Eurasia after Australia separated from the ancient South America-Africa continent, and South America had left Africa. The endemic tribe of North America, the tribe Sarcobateae, has a origin different from the tribes Salsoleae and Suaedeae of the subfamily Spirolobeae. Sarcobateae flowers diverged into unisexuality and absence of bractlets. Clearly they originated in North America after North America had left the Eurasian continent. North America and southern Africa have a few species of Salsola, but none of them have become very much differentiated or developed, so they must have arrived through overland migration across ancient continental connections. India has no southern African Chenopodiaceae floristic components except for a few maritime taxa, which shows that when the Indian subcontinent left Africa in the Triassic period, the Chenopodiaceae had not yet developed in Africa. Therefore, the early Cretaceous Period about 120 million years ago, when the ancient Gondwanan and Laurasian continents disintegrated, could have been the time of origin of Chenopodiaceae plants.The Chinese flora of Chenopodiaceae is a part of Chenopodiaceae flora of central Asia. Cornulaca alaschnica was discovered from Gansu, China, showing that the Chinese Chenopodiaceae flora certainly has contact with the Mediterranean Chenopodiaceae flora. The contact of southeastern China with the Australia Chenopodiaceae flora, however, is very weak.     

藜科植物的起源、分化和地理分布

全球藜科植物共约130属1500余种,广泛分布于欧亚大陆、南北美洲、非洲和大洋洲的半干旱及盐碱地区。它基本上是一个温带科,对亚热带和寒温带也有一定的适应性。本文分析了该科包含的1l族的系统位置和分布式样,以及各个属的分布区,提出中亚区是现存藜科植物的分布中心,原始的藜科植物在古地中海的东岸即华夏陆台(或中国的西南部)发生,然后向干旱的古地中海沿岸迁移、分化,产生了环胚亚科主要族的原始类群;起源的时间可能在白垩纪初,冈瓦纳古陆和劳亚古陆进一步解体的时期。文章对其迁移途径及现代分布式样形成的原因进行了讨论。     

Endemic or introduced? Phylogeography of Asparagopsis (Florideophyceae) in Australia reveals multiple introductions and a new mitochondrial lineage

The red seaweed Asparagopsis taxiformis embodies five cryptic mitochondrial lineages (lineage 1–5) introduced worldwide as a consequence of human mediated transport and climate change. We compared globally collected mitochondrial cox2‐3 intergenic spacer sequences with sequences produced from multiple Australian locations and South Korea to identify Asparagopsis lineages and to reveal cryptic introductions. We report A. taxiformis lineage 4 from Cocos (Keeling) Islands, Australia, and the highly invasive Indo‐Pacific Mediterranean lineage 2 from South Korea and Lord Howe Island, Australia. Phylogeographic analysis showed a clear haplotype and geographic separation between western Australian and Great Barrier Reef (GBR) isolates belonging to the recently described lineage 5. The same lineage, however, was characterized by a substantial genetic and geographic break between the majority of Australian specimens and Asparagopsis collections from South Solitary Island, Southern GBR, Lord Howe Island, Kermadec Islands, Norfolk Island, New Caledonia and French Polynesia. The disjunct geographic distribution and sequence divergence between these two groups supports the recognition of a sixth cryptic A. taxiformis mitochondrial lineage. As climatic changes accelerate the relocation of biota and offer novel niches for colonization, periodic surveys for early detection of cryptic invasive seaweeds will be critical in determining whether eradication or effective containment of the aliens are feasible.     

Fruit and Seed Anatomy of Chenopodium and Related Genera (Chenopodioideae,Chenopodiaceae/Amaranthaceae): Implications for Evolution and Taxonomy

A comparative carpological study of 96 species of all clades formerly considered as the tribe Chenopodieae has been conducted for the first time. The results show important differences in the anatomical structure of the pericarp and seed coat between representatives of terminal clades including Chenopodium s.str.+Chenopodiastrum and the recently recognized genera Blitum, Oxybasis and Dysphania. Within Chenopodium the most significant changes in fruit and seed structure are found in members of C. sect. Skottsbergia. The genera Rhagodia and Einadia differ insignificantly from Chenopodium. The evolution of heterospermy in Chenopodium is discussed. Almost all representatives of the tribe Dysphanieae are clearly separated from other Chenopodioideae on the basis of a diverse set of characteristics, including the small dimensions of the fruits (especially in Australian taxa), their subglobose shape (excl. Teloxys and Suckleya), and peculiarities of the pericarp indumentum. The set of fruit and seed characters evolved within the subfamily Chenopodioideae is described. A recent phylogenetic hypothesis is employed to examine the evolution of three (out of a total of 21) characters, namely seed color, testa-cell protoplast characteristics and embryo orientation.     

Examining the phylogeny of the Australasian Lymnaeidae (Heterobranchia: Pulmonata: Gastropoda) using mitochondrial, nuclear and morphological markers

We examined the species groups relationships of the freshwater snail genus Austropeplea using mitochondrial, nuclear and morphological markers in addition to traditional methods of shell shape analysis. Based primarily on the results of a combined molecular and morphological analysis, samples of the nominal species A. tomentosa form distinct lineages. The New Zealand populations of A. tomentosa are a very distinct lineage from any of the Australian populations attributed to A. tomentosa. Furthermore, within the Australian group, three lineages, south Australia, Tasmania and eastern Australia, appear to have undergone recent and/or rapid speciation events. Samples assigned to A. lessoni were resolved as two distinct lineages, representing the eastern and northern Australian populations. Kutikina hispida was resolved within the Australian A. tomentosa clade. Molecular results for A. viridis suggests that it is also composed of at least two distinct lineages that could be treated as species. Incongruence observed between the single mitochondrial, nuclear and morphological topologies highlight the importance of using a number of different datasets in the delimitation of species-group taxa.     

Radiation following long‐distance dispersal: the contributions of time,opportunity and diaspore morphology in Sicyos (Cucurbitaceae)

Aim  To infer the most plausible explanations for the presence of 14 species of the Neotropical cucurbit genus Sicyos on the Hawaiian Islands, two on the Galápagos Islands, two in Australia, and one in New Zealand. Location  Neotropics, the Hawaiian and Galápagos archipelagos, Australia and New Zealand. Methods  We tested long‐problematic generic boundaries in the tribe Sicyoeae and reconstructed the history of Sicyos using plastid and nuclear DNA sequences from 87 species (many with multiple accessions) representing the group’s generic and geographic diversity. Maximum likelihood and Bayesian approaches were used to infer relationships, divergence times, biogeographic history and ancestral traits. Results  Thirteen smaller genera, including Sechium, are embedded in Sicyos, which when re‐circumscribed as a monophyletic group comprises 75 species. The 14 Hawaiian species of Sicyos descended from a single ancestor that arrived c. 3 million years ago (Ma), Galápagos was reached twice at c. 4.5 and 1 Ma, the species in Australia descended from a Neotropical ancestor (c. 2 Ma), and New Zealand was reached from Australia. Time since arrival thus does not correlate with Sicyos species numbers on the two archipelagos. Main conclusions  A plausible mechanism for the four trans‐Pacific dispersal events is adherence to birds of the tiny hard fruit with retrorsely barbed spines found in those lineages that underwent long‐distance migrations. The Hawaiian clade has lost these spines, resulting in a lower dispersal ability compared with the Galápagos and Australian lineages, and perhaps favouring allopatric speciation.     

EVOLUTION OF ASEXUALITY IN THE COSMOPOLITAN MARINE CLAM LASAEA

The marine clam genus Lasaea is unique among marine bivalves in that it contains both sexual and asexual lineages. We employed molecular tools to infer intrageneric relationships of geographically restricted sexual versus cosmopolitan asexual forms. Polymerase chain reaction primers were used to amplify and sequence homologous 624 nucleotide fragments of COIII from polyploid, asexual, direct-developing individuals representing northeastern Pacific, northeastern Atlantic, Mediterranean, southern Indian Ocean, and Australian populations. DNA sequences also were obtained from the two known diploid congeners, the Australian sexual, indirect developer, Lasaea australis, and an undescribed meiotic Australian direct developer. Estimated tree topologies did not support monophyly for polyploid asexual Lasaea lineages. A robust dichotomy was evident in all phylogenetic trees and each of the two main branches included one of the diploid meitoic Australian congeners. Lasaea australis clustered with two of the direct-developing, polyploid asexual haplotypes, one from Australia, the other from the northeastern Atlantic. Monophyly is supported for the diploid Australian direct-developing lineage together with the remaining polyploid asexual lineages from the northeastern Pacific, northeastern Atlantic, Mediterranean, and southern Indian Ocean. These results indicate that asexual Lasaea lineages are polyphyletic and may have resulted from multiple hybridization events. The high degree of genetic divergence of asexual lineages from co-clustering meiotic congeners (16%–22%) and among geographically restricted monophyletic clones (9%–11%) suggests that asexual Lasaea lineages may be exceptionally long lived.     

Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis

? Premise of the study: Atripliceae (Chenopodiaceae), including Atriplex (300 spp.) as the largest genus of the family, are an ecologically important group of steppes and semideserts worldwide. Relationships in Atripliceae are poorly understood due to obscure and potentially convergent morphological characters. ? Methods: Using sequence variation of two chloroplast markers (rbcL gene, atpB-rbcL spacer) and one nrDNA marker (ITS) analyzed with BEAST, we investigated the systematics and biogeography of Atripliceae. We surveyed flower morphology and fruit anatomy to study the evolution of flowers and fruits in the tribe. ? Key results: Female flowers with persistent foliar cover (the diagnostic character of traditional Atripliceae) evolved three times in Chenopodioideae, in Atripliceae s.s., Axyrideae, and Spinacia. Atripliceae s.s. started to diversify during the Early Miocene in Eurasia, separating into the Archiatriplex and the Atriplex clades. The former consists of eight species-poor, disjunct, and morphologically heterogeneous genera and is likely a relictual lineage. The Atriplex clade comprises the majority of species and evolved one C(4) lineage 14.1-10.5 Ma, which diversified rapidly worldwide. The C(4) Atriplex entered North America during the Middle/Late Miocene and spread to South America subsequently. Australia was colonized by two C(4) lineages both arriving during the Late Miocene. One of them diversified rapidly, giving rise to most Australian Atriplex species. ? Conclusions: Atripliceae s.s. comprise Archiatriplex, Atriplex, Exomis, Extriplex, Grayia, Halimione, Holmbergia, Manochlamys, Proatriplex, and Stutzia. Microgynoecium is included based on morphology but only weak molecular support. Axyris, Krascheninnikovia, and Ceratocarpus (here described as Axyrideae) and Spinacia are excluded from Atripliceae.     

Diversification across biomes in a continental lizard radiation

Ecological opportunity is a powerful driver of evolutionary diversification, and predicts rapid lineage and phenotypic diversification following colonization of competitor‐free habitats. Alternatively, topographic or environmental heterogeneity could be key to generating and sustaining diversity. We explore these hypotheses in a widespread lineage of Australian lizards: the Gehyra variegata group. This clade occurs across two biomes: the Australian monsoonal tropics (AMT), where it overlaps a separate, larger bodied clade of Gehyra and is largely restricted to rocks; and in the larger Australian arid zone (AAZ) where it has no congeners and occupies trees and rocks. New phylogenomic data and coalescent analyses of AAZ taxa resolve lineages and their relationships and reveal high diversity in the western AAZ (Pilbara region). The AMT and AAZ radiations represent separate radiations with no difference in speciation rates. Most taxa occur on rocks, with small geographic ranges relative to widespread generalist taxa across the vast central AAZ. Rock‐dwelling and generalist taxa differ morphologically, but only the lineage‐poor central AAZ taxa have accelerated evolution. This accords with increasing evidence that lineage and morphological diversity are poorly correlated, and suggests environmental heterogeneity and refugial dynamics have been more important than ecological release in elevating lineage diversity.     

Molecular evidence for Gondwanan origins of multiple lineages within a diverse Australasian gecko radiation

Aim Gondwanan lineages are a prominent component of the Australian terrestrial biota. However, most squamate (lizard and snake) lineages in Australia appear to be derived from relatively recent dispersal from Asia (< 30 Ma) and in situ diversification, subsequent to the isolation of Australia from other Gondwanan landmasses. We test the hypothesis that the Australian radiation of diplodactyloid geckos (families Carphodactylidae, Diplodactylidae and Pygopodidae), in contrast to other endemic squamate groups, has a Gondwanan origin and comprises multiple lineages that originated before the separation of Australia from Antarctica. Location Australasia. Methods Bayesian (beast ) and penalized likelihood rate smoothing (PLRS) (r 8s ) molecular dating methods and two long nuclear DNA sequences (RAG‐1 and c‐mos) were used to estimate a timeframe for divergence events among 18 genera and 30 species of Australian diplodactyloids. Results At least five lineages of Australian diplodactyloid geckos are estimated to have originated > 34 Ma (pre‐Oligocene) and basal splits among the Australian diplodactyloids occurred c. 70 Ma. However, most extant generic and intergeneric diversity within diplodactyloid lineages appears to post‐date the late Oligocene (< 30 Ma). Main conclusions Basal divergences within the diplodactyloids significantly pre‐date the final break‐up of East Gondwana, indicating that the group is one of the most ancient extant endemic vertebrate radiations east of Wallace’s Line. At least five Australian lineages of diplodactyloid gecko are each as old or older than other well‐dated Australian squamate radiations (e.g. elapid snakes and agamids). The limbless Pygopodidae (morphologically the most aberrant living geckos) appears to have radiated before Australia was occupied by potential ecological analogues. However, in spite of the great age of the diplodactyloid radiation, most extant diversity appears to be of relatively recent origin, a pattern that is shared with other Australian squamate lineages.     

Contrasting patterns of radiation in African and Australian Restionaceae

The floras of the Mediterranean-climate areas of southern Africa and southwestern Australia are remarkably species rich. Because the two areas are at similar latitudes and in similar positions on their respective continents, they have probably had similar Cenozoic climatic histories. Here we test the prediction that the evolution of the species richness in the two areas followed a similar temporal progression by comparing the rates of lineage accumulation for African and Australian Restionaceae. Restionaceae (Poales) are typical and often dominant elements in the fynbos vegetation of the Cape Floristic Region of southern Africa and the kwongan vegetation of the Southwestern Floristic Province of Western Australia. The phylogeny of the family was estimated from combined datasets for rbcL and trnL-F sequences and a large morphological dataset; these datasets are largely congruent. The monophyly of Restionaceae is supported and a basal division into an African clade (approximately 350 species) and an Australian clade (146 species) corroborated. There is also support for a futher subdivision of these two large sister-clades, but the terminal resolution within the African clade is very weak. Fossil pollen records provided a minimum age of the common ancestor of Australian and African Restionaceae as 64-71 million years ago, and this date was used to calibrate a molecular clock. A molecular clock was rejected by a likelihood ratio test; therefore, rate changes between the lineages were smoothed using nonparametric rate smoothing. The rate-corrected ages were used to construct a plot of lineages through time. During the Palaeogene the Australian lineage diversity increased consistent with the predictions of the constant birthrate model, while the African lineage diversity showed a dramatic increase in diversification rate in the Miocene. Incomplete sampling obscures the patterns in the Neogene, but extending the trends to the modern extant diversity suggests that this acceleration in the speciation rate continued in the African clade, whereas the Australian clade retained a constant diversification rate. The substantial morphological and anatomical similarity between the African and Australian Restionaceae appear to preclude morphological innovations as possible explanations for the intercontinental differences. Most likely these differences are due to the greater geographical extent and ecological variation in temperate Australia than temperate Africa, which might have provided refugia for basal Restionaceae lineages, whereas the more mountainous terrain of southern Africa might have provided the selective regimes for a more rapid, recent speciation.     

Repeated trans-Atlantic dispersal catalysed a global songbird radiation

Aim Turdus thrushes are one of the most speciose and widespread songbird genera, comprising nearly 70 species that combined have a near‐global distribution. Herein, we use molecular phylogenetic, molecular clock and behavioural evidence to examine the historical biogeography of the genus. Ancestral area reconstructions in conjunction with divergence estimates and palaeoclimatogical data are used to test whether the long‐standing paradigm of Beringian colonization or trans‐Atlantic dispersal best explains modern distributions in the New and Old Worlds. Location Worldwide, with emphasis on New World–Old World biotic interchange. Methods Using a molecular phylogenetic hypothesis of Turdus thrushes, we reconstructed ancestral area relationships utilizing the five major continental or regional areas occupied by species in the genus. We also examined the evolution of behaviours on the phylogeny, and estimated the timing of major lineage divergences via a molecular clock. Results Turdus originated in Eurasia, and following the colonization of Africa underwent a series of five trans‐Atlantic sweepstake dispersals. The data reject the alternative hypothesis that connections between Old and New World Turdus species can be attributed to movement through Beringia with subsequent extinction. Divergence estimates indicate that these dispersals all occurred near the Miocene–Pliocene boundary, 5 Ma. A significant phylogenetic correlation between migratory and flocking behaviour is evident in the genus. Main conclusions The initial divergence of Turdus in the Old World was followed by a series of trans‐Atlantic sweepstake dispersal events. These dispersals are temporally correlated with a specific palaeoclimatic system, which would have facilitated transport of Turdus from the Caribbean to the Old World across the Atlantic. Uplift of the Central American Seaway 4.7 Ma effectively shut down the palaeoclimatic system, and no additional trans‐Atlantic dispersals are evident in Turdus after this time. Migratory movements by ancestral lineages in flocks, rather than as single individuals, suggest an increased likelihood of successfully colonizing new areas, post‐dispersal.     

Phylogeny of kangaroo apples (<Emphasis Type="Italic">Solanum</Emphasis> subg. <Emphasis Type="Italic">Archaesolanum</Emphasis>, Solanaceae)

Kangaroo apples, subgenus Archaesolanum, are a unique and still poorly known group within the genus Solanum. Here we aimed to reveal phylogeny, historical biogeography and age of diversification of Archaesolanum. We sampled all recognized species of the group and sequenced three chloroplast regions, the trnT-trnL spacer, trnL intron and trnL-trnF spacer to calibrate a molecular clock to estimate the age of the group. Distributional data were combined with the results of phylogenetic analysis to track the historical processes responsible for the current range of the group. Our analysis supported the monophyly of the kangaroo apples and the biogeographical disjunction between the two subclades within the group. Based on the divergence time estimates the most recent common ancestor of kangaroo apples is from the late Miocene age (~9 MYA). Based on the age estimate the common ancestors of the kangaroo apples are presumed to have arrived in Australia by long-distance dispersal. The two distinct lineages within the group have separated during the aridification of the continent and further speciated in the brief resurgence of rainforests during the Pliocene.     

Long-distance colonization of the Western Mediterranean by Cistus ladanifer (Cistaceae) despite the absence of special dispersal mechanisms

Aim There are few biogeographical and evolutionary studies that address plant colonization and lineage origins in the Mediterranean. Cistus serves as an excellent model with which to study diaspore dispersal and distribution patterns of plants exhibiting no special long‐distance dispersal mechanisms. Here we analyse the pattern of genetic variation and divergence times to infer whether the African–European disjunction of C. ladanifer L. is the result of long‐distance dispersal or of vicariance events. Location Principally the Western Mediterranean region, with a focus on the Strait of Gibraltar. Methods We used DNA sequence phylogenetic approaches, based on plastid (rbcL/trnK‐matK) and nuclear (ITS) sequence data sets, and the penalized likelihood method, to date the diversification of the 21 species of Cistus. Phylogenetic relationships and phylogeographical patterns in 47 populations of C. ladanifer were also analysed using two plastid DNA regions (trnS‐trnG, trnK‐matK). These sequence data were analysed using maximum parsimony, Bayesian inference and statistical parsimony. Results Dating estimates indicated divergence dates of the C. ladanifer lineage in the Pleistocene. Eight nucleotide‐substitution haplotypes distributed on the European (four haplotypes) and African (five haplotypes) sides of the Strait of Gibraltar were revealed from C. ladanifer sequences. Both the haplotype network and the phylogenetic analyses depicted two main Cistus lineages distributed in both Europe and North Africa. An Iberian haplotype forms part of the North African lineage, and another haplotype distributed on both continents is related to the European lineage. Haplotype relationships with respect to outgroup sequences supported the hypothesis that the centre of genetic diversity is in northern Africa. Main conclusions Based on lineage divergence‐time estimates and disassociation between geographical and lineage haplotype distributions, we inferred at least two intercontinental colonization events of C. ladanifer post‐dating the opening of the Strait of Gibraltar (c. 5 Ma). This result supports a hypothesis of long‐distance dispersal rather than a hypothesis of vicariance. We argue that, despite limited dispersal abilities, preference for disturbed habitats was integral to historical colonization after the advent of the Mediterranean climate (c. 3.2 Ma), when Cistus species diverged and became established as a dominant element in the Mediterranean scrub.     

Charting the course of reed‐warblers across the Pacific islands

Aim Deciphering the complex colonization history of island archipelagos is greatly facilitated by comprehensive phylogenies. In this study we investigate the phylogeny and biogeography of the insular reed‐warblers (genus Acrocephalus) of the tropical Pacific Ocean, from Australia to eastern Polynesia. Location Oceania. Methods We used sequences of mitochondrial DNA (cytochrome b, ND2 and ATP8 genes) to infer the colonization patterns of reed‐warblers endemic to Pacific islands and Australia. We sampled all known taxa of Acrocephalus in the Pacific except A. luscinius nijoi, for which no sample was available. Most taxa were represented by toe‐pad samples from museum specimens collected in the 19th and 20th centuries. With a few exceptions, several specimens per taxon were sequenced independently in two institutions (Smithsonian Institution and Natural History Museum of Geneva). Results Our data indicate that Pacific reed‐warblers do not form a monophyletic group, because A. luscinius luscinius from Guam falls outside the main Pacific radiation. The remaining Pacific taxa are divided into two clades: one clade includes all the reed‐warblers from Micronesia (except Guam) and Australia, and two Polynesian taxa from the Line Islands and the southern Marquesas; the other clade includes all remaining Polynesian taxa. The taxa endemic to three archipelagos (Mariana, Marquesas and Society islands) are polyphyletic, suggesting several independent colonizations. Main conclusions Our results provide evidence for a complex pattern of colonization of the Pacific by reed‐warblers. Calibration analyses suggest that reed‐warbler lineages are much younger than the ages of the islands they occupy. Several remote archipelagos were colonized independently more than once. Consequently, we infer that the colonization of reed‐warblers in the Pacific did not follow a regular, stepping‐stone‐like pattern. The phylogeny also suggests a previously undetected case of reverse colonization (from island to continent) for the Australian lineage and indicates that A. luscinius, as currently defined, is not monophyletic. We discuss the supertramp strategy of reed‐warblers in the Pacific and show that, although Pacific reed‐warblers meet some of the supertramp criteria in their aptitude for colonizing remote archipelagos, their life history characteristics do not fit the model.     

Reproductive isolation between phylogeographic lineages scales with divergence

Phylogeographic studies frequently reveal multiple morphologically cryptic lineages within species. What is not yet clear is whether such lineages represent nascent species or evolutionary ephemera. To address this question, we compare five contact zones, each of which occurs between ecomorphologically cryptic lineages of skinks from the rainforests of the Australian Wet Tropics. Although the contacts probably formed concurrently in response to Holocene expansion from glacial refugia, we estimate that the divergence times (τ) of the lineage pairs range from 3.1 to 11.5 Ma. Multi-locus analyses of the contact zones yielded estimates of reproductive isolation that are tightly correlated with divergence time and, for lineages with older divergence times (τ > 5 Myr), substantial. These results show that phylogeographic splits of increasing depth represent stages along the speciation continuum, even in the absence of overt change in ecologically relevant morphology.     

Phylogeography of the parasitic fly Batrachomyia in the Wet Tropics of north-east Australia, and susceptibility of host frog lineages in a mosaic contact zone

Parasites could differentially impact intraspecific host lineages due to genetic, phenotypic, ecological, or behavioural differences between the lineages, or the development of reproductive isolation between them. Batrachomyia (Diptera: Chloropidae) are flies that exclusively parasitize Australian frogs, and in the Wet Tropics rainforest of north‐east Australia larvae are largely restricted to the green‐eyed tree frog Litoria genimaculata (Anura: Hylidae). This frog species consists of two highly divergent genetic lineages that overlap in two nearby, but independent, contact zones. At one contact zone there has been extensive phenotypic divergence and speciation between the lineages whereas, at the other contact relatively lower levels of phenotypic divergence and reproductive isolation suggest that speciation has not occurred. In the present study we tested: (1) whether the deep phylogeographic divergence between northern and southern host populations is mirrored by congruent genetic structuring in the parasite populations and (2) whether the host lineages are differentially impacted by parasitism. We found that the two divergent frog lineages are parasitized by a single lineage of Batrachomyia, which exhibits strikingly little phylogeographic structuring. We found a significant difference in Batrachomyia prevalence between the host lineages at mixed lineage sites in both contact zones, with the magnitude and direction of this effect being consistent in both. The pattern did not differ between the two contacts even though recent phenotypic divergence and speciation has occurred between the lineages at one contact but not the other. Taken together, this suggests a fundamental difference in susceptibility between the genetically divergent host lineages. Using weight relative to body length as a measure of body condition, we found no differential impact of parasitism on the body condition of each host lineage, and no evidence that parasitism impacts the body condition of the host in general. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 593–603.     

Navigating the mtDNA road map out of the morphological maze: interpreting morphological variation in the diverse Monomorium rothsteini (Forel) complex (Hymenoptera: Formicidae)

Monomorium is a large and diverse ant genus with speciose radiations in both the Afrotropical and Australian regions. According to the most recent taxonomic revision, many Australian species are characterised by very broad distributions and variable morphology, which suggests that some species may be unrecognised species complexes. With a continent‐wide distribution and diverse yet overlapping morphology, M. rothsteini (Forel) is representative of the greater challenge that exists in Australian Monomorium systematics. Here we investigate species boundaries in M. rothsteini using a molecular phylogenetic framework to interpret the complex overlap of nine morphological characters (with 31 states) and examine biogeographic relationships among the lineages. Bayesian inference resolved 38 mtDNA lineages that were morphologically separable, at least from their sister lineage. Although the morphological characters were intermixed across the phylogeny, instances of inseparable morphology among sister clades was rare. Seventeen lineages exhibited complete morphological overlap with one or more other lineages and could not be separated by Principal Component Analysis based on 12 morphometric variables. Two‐thirds of all lineages occurred sympatrically with one or more both genetically and morphologically divergent lineages. The two nuclear markers EF1αF2 and wingless were used to generate haplotype networks which were characterised by a star‐like pattern indicative of a rapid and recent radiation. Several haplotypes for both nuclear gene regions were shared among individuals occurring in separate mtDNA clades which we were also unable to distinguish morphologically or that were occurring in sympatry, indicating possible introgression in both the mtDNA and nuclear genomes. Clear biogeographic affinities among samples within a lineage were detected but there was no overall pattern in the biogeographic relationships among the lineages. We conclude that M. rothsteini is a large species complex that has undergone a complex evolutionary history following aridification of the Australian continent, and discuss the implications of this conclusion for the systematics of Australian Monomorium more generally.