Phylogeny,diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae)

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on ～4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1‐α (EF1‐α), arginine kinase (ArgK) and RNA polymerase II (Pol‐II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil‐calibrated molecular clocks and the evolution of morphological traits using disparity‐through‐time plots. In addition, we inferred past biogeographical events by implementing model‐based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27–42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 552–572.     

Molecular systematics and evolution of the subgenus Mesocarabus Thomson, 1875 (Coleoptera: Carabidae: Carabus), based on mitochondrial and nuclear DNA

The subgenus Mesocarabus Thomson, 1875 is a western Palaearctic group that currently includes five species: four of them inhabiting western Europe (Carabus lusitanicus Fabricius, 1801, Carabus problematicus Herbst, 1786, Carabus dufourii Dejean & Boisduval, 1829, and Carabus macrocephalus Dejean, 1826) and one found in the Rif Mountains in northern Morocco (Carabus riffensis Fairmaire, 1872). Representatives of Mesocarabus have been included in previous molecular phylogenetic studies, but taxon‐ or gene‐sampling limitations yielded inconclusive results regarding its monophyly and sister relationship. Here we perform molecular phylogenetic analyses based on five mitochondrial (3625 nt) and eight nuclear (5970 nt) genes sequenced in many Mesocarabus populations, and in related western Palaearctic Carabus Linnaeus, 1758. We conducted parsimony, maximum‐likelihood, and Bayesian analyses and found a well‐supported sister relationship between a monophyletic Mesocarabus with Iberian species of the subgenus Oreocarabus Géhin, 1876. Within Mesocarabus, the European species form a monophyletic lineage sister to Moroccan C. riffensis. A time‐calibrated phylogeny suggests the split between Mesocarabus and Oreocarabus occurred at 11.8 Mya (95% highest posterior density, HPD, 8.7–15.3 Mya), and the divergence between C. riffensis and European Mesocarabus at 9.5 Mya (95% HPD 7.0–12.5 Mya). The early diversification of Mesocarabus and related subgenera during the Miocene, and alternative hypotheses concerning the origin of Mesocarabus in the Iberian Peninsula and the Betic‐Riffian plate are discussed using calibration data and dispersal–vicariance biogeographic analyses. Finally, we found instances of incongruence between mitochondrial DNA and nuclear‐based phylogenies of Mesocarabus, which are hypothesized to be the result of introgressive hybridization. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 787–804.     

Molecular phylogeny of the bee genus Hoplitis (Megachilidae: Osmiini) – how does nesting biology affect biogeography?

The genus Hoplitis (Megachilidae: Osmiini) comprises about 360 described species and occurs on all continents except Australia, South America, and Antarctica. Using five genes, we inferred the phylogeny of Hoplitis including 23 out of the 27 currently recognized subgenera, applying both Bayesian and maximum likelihood methods. Compared to the current morphology‐based classification, our phylogeny resulted in three classificatory changes: first, the subgenera Alcidamea, Cyrtosmia, Dasyosmia, Megalosmia, Monumetha, and Prionohoplitis are merged into one large subgenus Alcidamea Cresson, 1864, comb. nov. ; second, the subgenera Annosmia, Bytinskia, Coloplitis, and Hoplitis are merged into one large subgenus Hoplitis Klug, 1807, comb. nov. ; third, the subgenera Acrosmia, Hoplitina, Penteriades, and Proteriades are merged into one large subgenus Proteriades Titus, 1904, comb. nov. We provide evidence that the genus Hoplitis has a Palaearctic origin and that colonization events to southern Africa and to the Nearctic, as well as recolonization events from the Nearctic to the Palaearctic occurred. The species of the genus Hoplitis exhibit an extraordinary diversity in nesting behaviour, comprising both below and above ground nesting. Parsimony mapping revealed that ground nesting in excavated burrows is the ancestral state amongst Hoplitis bees. We hypothesize that nesting biology strongly affected both range expansion and long‐distance dispersal in Hoplitis. © 2012 The Linnean Society of London     

Specialized pollination by carpenter bees in Calanthe striata (Orchidaceae), with a review of carpenter bee pollination in orchids

Calanthe striata has nectarless flowers that are self‐compatible, but pollinator dependent. Field observations showed that the flowers were pollinated exclusively by the carpenter bee Xylocopa appendiculata circumvolans, although the bees occasionally wasted pollen by delivering to the stigmatic surface pollinaria that retained their anther caps. Fruit set ratios at the population level varied spatiotemporally, but were generally low (8.3–17.3%). Calanthe striata blooms in spring when post‐overwintering carpenter bees have not yet started foraging for brood production. It can therefore exploit an abundance of opportunistic/naïve foragers. This timing may also increase the possibility of pollinator visits, because no rewarding co‐flowering plants are available in the orchid habitats. A literature review of Orchidaceae pollinated by carpenter bees revealed that at least 14 species of Orchidoideae and Epidendroideae have evolved flowers specialized for carpenter bee pollination. They typically have shallow pink/magenta flowers with a foothold for pollinators; pollinaria are attached to the head, ventral thorax or base of the middle legs of carpenter bees when they insert their heads and/or proboscises into flowers; pollination success is generally low, a probable consequence of the deceptive pollination systems. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 730–743.     

Comparison of social and solitary nesting carpenter bees in sympatry reveals no advantage to social nesting

Facultatively social bees allow for comparisons of social and solitary behaviour under similar environmental conditions. When such bees are polymorphic within the same population it provides a special and somewhat unusual opportunity to examine factors leading to cooperative (social) behaviours where many parameters such as environment are not variable. Some species of bees in the genus Xylocopa offers such opportunities. Studies of these bees often point to guarding and resource limitation as factors leading to social nesting. The large carpenter bee Xylocopa virginica L. (Hymenoptera: Apidae) is one species that exhibits both solitary and social nesting behaviour within the same populations. This paper first compares social and solitary nests in an Ontario population of X. virginica to determine if there is a reproductive advantage to social nesting. Following this, a series of possible explanations for social nesting, and the roles of females in social nests are examined. Social nests have similar brood sizes to solitary nests and productivity as a function of colony size is reduced with increasing number of foundresses. Additional foundresses are not effective guards, do not prevent parasitism, do not likely perform significant work, and do not assume foraging with the loss of a primary foundress. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 998–1010.     

Molecular phylogeny of the large carpenter bees, genus Xylocopa (Hymenoptera: apidae), based on mitochondrial DNA sequences

Carpenter bees, genus Xylocopa Latreille, a group of bees found on all continents, are of particular interest to behavioral ecologists because of their utility for studies of the evolution of mating strategies and sociality. This paper presents phylogenetic analyses based on sequences of two mitochondrial genes cytochrome oxidase 1 and cytochrome b for 22 subgenera of Xylocopa. Maximum-parsimony and maximum-likelihood methods were used to infer phylogenetic relationships. The analyses resulted in three resolved clades of subgenera: a South American group (including the subgenera Stenoxylocopa, Megaxylocopa, and Neoxylocopa), a group including the subgenera Xylocopa s.s. and Ctenoxylocopa, and an Ethiopean group (including the subgenera Afroxylocopa, Mesotrichia, Alloxylocopa, Platynopoda, Hoploxylocopa, and Koptortosoma). The relationships between the 11 other subgenera and the resolved clades are unclear. Within the Ethiopian group we found a clear separation of the African and the Oriental taxa and apparent polyphyly of the subgenus Koptortosoma. Using an evolutionary rate for ants, we investigated whether Gondwana vicariance or more recent dispersal events could best explain the present-day distribution of subgenera. Although some taxa show divergences that approach Gondwanan breakup times, most divergences between geographic groups are too recent to support a vicariance hypothesis.     

Biogeographic distribution of the Phytoseiidae (Acari: Mesostigmata)

More than 1982 species in 90 genera were included in an analysis of the biogeography of the Phytoseiidae, a family of predatory mites. Seven biogeographic regions were taken into account: Nearctic, Neotropical, Ethiopian, West Palaearctic, East Palaearctic, Oriental, and Australasian. The number of species was particularly high in the Neotropical, Oriental, and West Palaearctic regions. These regions also present the highest levels of species endemism. The number of genera was quite similar in all regions except for the Neotropics, which also had a high level of endemism. The possible Gondwanian (Neotropical, Ethiopian, Australasian, and Oriental regions) origin of the Phytoseiidae, most probably in the Neotropics, and their possible radiation to Laurasia (Nearctic, West Palaearctic, and East Palaearctic regions) are discussed. The comparison between genera and species in the different biogeographic regions indicate the importance of both dispersal and vicariance events in the evolution of the group. Dispersal is assumed to have been most important between Neotropical and Nearctic regions and between East Palaearctic and Oriental regions, whereas vicariance could have been the dominating process between Australasian, Ethiopian, and Oriental regions, as well as between West and East Palaearctic regions. A parsimony analysis of endemicity showed the Neotropical and the Nearctic regions to be isolated from the other regions. This is certainly due to a diversification after the continents drifted apart and then a high dispersal between Nearctic and Neotropical regions. Different phylogenetic hypotheses and scenarios are proposed for each subfamily based on the results obtained and further investigations are proposed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 845–856.     

Phylogenetic relationships within Plantago (Plantaginaceae): evidence from nuclear ribosomal ITS and plastid trnL-F sequence data

A molecular phylogenetic study of Plantago L. (Plantaginaceae) analysed nucleotide variation in the internal transcribed spacers (ITS) of nuclear ribosomal and plastid trnL-F regions. Included are 57 Plantago species, with two Aragoa species as the ingroup and three Veronica species as the outgroup. Phylogenetic analysis using maximum parsimony identified five major clades, corresponding to the taxonomic groups Plantago subgenera Plantago, Coronopus, Psyllium, Littorella and Bougueria . Aragoa is sister to genus Plantago . Plantago subgenus Littorella is sister to the other subgenera of Plantago . The results are in general correlated with a morphological phylogenetic study and iridoid glucoside patterns, but Plantago subgenus Albicans is paraphyletic and should be included in Plantago subgenus Psyllium sensu lato to obtain a monophyletic clade with six sections. Plantago section Hymenopsyllium is more closely related to section Gnaphaloides than to section Albicans . Plantago subgenus Bougueria is sister to subgenus Psyllium s.l. section Coronopus in Plantago subgenus Coronopus is subdivided in two series. Only some of the sections can be resolved into series. DNA variation within genus Plantago is high, a result that would not have been predicted on the basis of morphology, which is relatively stereotyped. If we calibrate a molecular clock based on the divergence of P. stauntoni , endemic to New Amsterdam in the southern Indian Ocean, we calculate the time of the split between Plantago and Aragoa to be 7.1 million years ago, which is congruent with the fossil record.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 323–338.     

Brood development and nest parasitism of Xylocopa (Neoxylocopa) augusti Lepeletier (Hymenoptera: Apidae), a promising crop pollinator in Argentina

Pollination of passion fruit and other crops by species of carpenter bees of the genus Xylocopa Latreille significantly increases both the quality and the quantity of fruits. To enhance pollination services, bee nests are either introduced into the crop area or females are encouraged to nest using trap nests. Thus, knowledge of trap-nest preference, brood development and nest parasitism is essential for effective and sustainable nest management practices. Xylocopa (Neoxylocopa) augusti Lepeletier is a promising pollinator in some areas of Argentina because of its high abundance, ability to buzz-pollinate and easy acceptance of trap nests. However, limited information is available on the biology of this pollinator. Herein, we provide detailed information on its nesting cycle, brood development and parasitism from trap nests at an urban area in Buenos Aires province, Argentina. Our study indicates that X. augusti is a solitary and likely univoltine species that shares some nesting and developmental features with other species in the subgenus Neoxylocopa, which might facilitate the adoption of existing management techniques developed for other species and regions. Biological information on Physocephala wulpi Camras (Diptera: Conopidae), a parasitoid of X. augusti, is also given for the first time.     

An alien Sennertia mite (Acari: Chaetodactylidae) associated with an introduced Oriental bamboo‐nesting large carpenter bee (Hymenoptera: Apidae: Xylocopa) invading the central Honshu Island,Japan

Since 2006, an introduced Oriental bamboo‐nesting large carpenter bee, Xylocopa tranquebarorum, has been recorded from the central Honshu Island, Japan, which is inhabited only by the endemic subspecies, Xylocopa appendiculata circumvolans. Carpenter bees (tribes Xylocopini and Ceratinini) have ecological associations with specific Sennertia spp. in all geographic regions of their distribution, thus it is worried that the introduced carpenter bee has brought non‐indigenous mites into Japan. In their native ranges, X. a. circumvolans and X. tranquebarorum each has specific Sennertia mite faunas: the four Japanese Alloxylocopa bees including X. a. circumvolans have associations with S. alfkeni, while X. tranquebarorum has association with S. potanini in China (except Taiwan) and with S. horrida in South to East Asia including Taiwan. In the present study, we examined phoretic mite fauna on the introduced X. tranquebarorum, and determined whether the mites are indigenous or not based on morphological character and two gene sequences (mitochondrial cytochrome oxidase subunit I gene and nuclear ribosomal internal transcribed spacer). It was found from the result of this study that the non‐indigenous Sennertia mite has invaded Japan with the introduced X. tranquebarorum. We discuss geographic origin of the introduced X. tranquebarorum based on associated mite fauna and potential ecological risk caused by the introduced Xylocopa‐Sennertia association.     

The historical biogeography of Apsilochorema (Trichoptera,Hydrobiosidae) revised,following molecular studies

The genus Apsilochorema Ulmer, 1907 is unique in the family Hydrobiosidae Ulmer, being widely distributed in the Palaearctic, Oriental and Australian Regions. All other 49 genera in the family, except the New World Atopsyche Banks, 1905, are confined to a single biogeographical Region. This unique distribution has independently stimulated researchers to formulate competing hypotheses about the biogeographical history of the genus. Molecular sequence data from mitochondrial cytochrome oxidase I (COI) and nuclear cadherin (CAD) genes of Apsilochorema species from the Oriental and Australian areas were analysed phylogenetically. The results retain a monophyletic Apsilochorema, which forms the sistergroup to the other genera in the subfamily Apsilochorematinae. The results from the biogeographical analyses dispute the earlier assumptions of an Oriental or northern Gondwana origin for the genus, revealing unambiguously an initial Australian radiation of the ancestral Apsilochorema with a subsequent dispersal into the Oriental Region. All but one of the Apsilochorema species occurring on the Pacific islands had an Oriental ancestor. The exception is the sistergroup to the New Caledonian species, which is found in both Australia and Oriental Regions. The molecular dating analysis, using a relaxed clock model, indicates that the genus Apsilochorema is about 36.4 MY old and that it dispersed from Australia into the Oriental Region about 28.3 Ma. It also gives an estimate of the approximate ages of the dispersals into New Caledonia to about 15.3 Ma; to the Solomon Islands at about 16.2 Ma; to the Fiji Islands at about 16.1 Ma; and to the Vanuatu Islands at about 5.4 Ma.     

A tale of North and South America: time and mode of dispersal of the amphitropical genus Munroa (Poaceae,Chloridoideae)

Plant disjunctions have provided some of the most intriguing distribution patterns historically addressed by biogeographers. We evaluated the three hypotheses that have been postulated to explain these patterns [vicariance, stepping‐stone dispersal and long‐distance dispersal (LDD)] using Munroa, an American genus of grasses with six species and a disjunct distribution between the desert regions of North and South America. The ages of clades, cytology, ancestral characters and areas of distribution were investigated in order to establish relationships among species, to determine the time of divergence of the genus and its main lineages, and to understand further the biogeographical and evolutionary history of this genus. Bayesian inference recovered the North American M. pulchella as sister species to the rest. Molecular dating and ancestral area analyses suggest that Munroa originated in North America in the late Miocene–Pliocene (7.2 Mya; 8.2–6.5 Mya). Based on these results, we postulate that two dispersal events modelled the current distribution patterns of Munroa: the first from North to South America (7.2 Mya; 8.2–6.5 Mya) and the second (1.8 Mya; 2–0.8 Mya) from South to North America. Arid conditions of the late Miocene–Pliocene in the Neogene and Quaternary climatic oscillations in North America and South America were probably advantageous for the establishment of populations of Munroa. We did not find any relationship between ploidy and dispersal events, and our ancestral character analyses suggest that shifts associated with dispersal and seedling establishment, such as habit, reproductive system, disarticulation of rachilla, and shape and texture of the glume, have been important in these species reaching new areas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 110–125.     

Testing the role of the Himalaya Mountains as a dispersal barrier in freshwater gastropods (Gyraulus spp.)

The Himalayas are considered to constitute a biogeographical boundary between the Palaearctic and Oriental regions. However, this mountain range does not form a uniform barrier because several large river valleys deeply cut into the Himalayas, crossing drainage divides and potentially forming dispersal corridors, particularly for freshwater organisms. In the present study, the effectiveness of these corridors is tested for the first time based on molecular data, using the amphi‐Himalayan freshwater gastropod genus Gyraulus as a model group. Based on a broad spatial sampling from the Himalayas and the Tibetan Plateau, phylogenetic analyses were performed using mitochondrial DNA sequence data. The resulting phylogeny shows that northern and southern Himalayan clades are geographically distinct (i.e. they exclusively consist of either northern or southern specimens). This pattern suggests a lack of gene flow across the Himalayas probably at least since the late Pliocene/early Pleistocene. Successful dispersal and/or establishment of gastropods might have been impeded by geographical and ecological features. Instead of direct dispersal across the range, multiple colonizations of regions north and south of the Himalayas from extralimital areas have to be assumed. The Himalayas thus represent a very strong dispersal barrier for freshwater snails, and probably for other taxa as well. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 526–534.     

Phylogeny and divergence time of island tiger beetles of the genus Cylindera (Coleoptera: Cicindelidae) in East Asia

To examine the diverse colonization histories in eight tiger beetle species of the genus Cylindera (Coleoptera: Cicindelidae) on the East Asian islands, we conducted phylogenetic analyses and divergence time estimation using mitochondrial cytochome oxidase subunit I (COI) and nuclear 28S rDNA sequences. The island fauna consisted of four subgenera: Apterodela, Cicindina, Ifasina, and Cylindera. Apterodela is a flightless group with large bodies, whereas the others are fliers with small bodies. In Apterodela, the divergence among endemic species in Taiwan, Japan, and the mainland was ancient (2.1–4.7 Mya), as expected from their flightlessness. Their dispersal might have occurred across the extended landmass in East Asia during the Pliocene. In the subgenus Cicindina, Cylindera elisae has spread throughout East Asia, from which an endemic species, Cylindera bonina, was derived on the oceanic Bonin Islands during the early Pleistocene (0.9 Mya). This indicates the significance of Cylindera bonina, which is currently confined to a single island, for conservation. In the subgenus Ifasina, Cylindera kaleea is widely distributed in East Asia, and its sister species Cylindera humerula, endemic to Okinawa Island, diverged 1.0 Mya, whereas Cylindera psilica on Taiwan and the Yaeyama Islands diverged approximately 0.8 Mya. In the subgenus Cylindera, the colonization of Cylindera gracilis in Japan from the mainland occurred during the last glacial period. With the exception of C. bonina, which likely colonized new territories by flight or drifting, other dispersal events might have used land connections that occurred repeatedly during the Pliocene and Pleistocene. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 715–727.     

Geometric morphometric analysis of a new Miocene bumble bee from the Randeck Maar of southwestern Germany (Hymenoptera: Apidae)

The first fossil bumble bee (Apinae: Bombini) from the Miocene Randeck Maar of southwestern Germany is described and illustrated. The specimen is subjected to a geometric morphometric analysis along with a diversity of other bumble bee species representing most major extant lineages, and particularly the subgenus Bombus s.s. The morphometric analysis supports the placement of the Randeck Maar species within Bombus s.s., as a species distinct from all others in the subgenus. It shows that extant subgenera of bumblebees were already derived in the early/middle Miocene. The Randeck Maar fossil is formally described as Bombus (Bombus) randeckensis Wappler & Engel sp. n. .     

Revision and phylogeny of the bee genus Paratetrapedia Moure,with description of a new genus from the Andean Cordillera (Hymenoptera,Apidae, Tapinotaspidini)

The bee genus Paratetrapedia represents a commonly collected group of bees and is especially diverse in forested areas of the Neotropics. Its taxonomy has remained poorly understood because of a lack of modern revisionary work and numerous species described as Tetrapedia whose type specimens have not been re‐examined in recent times. Here, a comprehensive study was carried out to review the taxonomy of the genus Paratetrapedia and to investigate cladistically the relationships amongst its species. Eighteen new species of Paratetrapedia are described, giving a total of 32 species in the genus. A phylogenetic analysis of the species of Paratetrapedia was carried out using 61 morphological characters for 41 terminal taxa. The phylogenetic results confirmed the monophyly of Paratetrapedia and allowed the recognition of five species groups: the lugubris, moesta, bicolor, lineata, and flavipennis groups. Nasutopedia gen. nov. , recognized as the sister group of Paratetrapedia and with its distribution restricted to the western forested portions of the Andean highlands, is proposed based on distinct morphology, its placement in the phylogenetic tree, and biogeographical patterns. Species of Paratetrapedia are especially diverse in the Amazon Forest; the eastern Brazilian Atlantic Forest contains four endemic species, and one species is endemic to the Cerrado of central Brazil. Paratetrapedia shows a biogeographical pattern similar to other Neotropical groups of bees and birds, with wide distribution and high diversity in lowland forests and whose sister taxon occurs on highlands of north‐western portions of the Andean cordillera. Identification keys for males and females of all species are provided, as well as distribution maps and illustrations of general external morphology and genitalia. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 351–442.     

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.     

Phylogeography of the whelk genus Cominella (Gastropoda: Buccinidae) suggests long‐distance counter‐current dispersal of a direct developer

The gastropod genus Cominella Gray, 1850 consists of approximately 20 species that inhabit a wide range of marine environments in New Zealand and Australia, including its external territory, the geographically isolated Norfolk Island. This distribution is puzzling, however, with apparently closely‐related species occurring either side of the Tasman Sea, even though all species are considered to have limited dispersal abilities. To determine how Cominella attained its current distribution, we derived a dated molecular phylogeny, which revealed a clade comprising all the Australian and Norfolk Island species nested within four clades of solely New Zealand species. This Australian clade diverged well after the vicariant separation of New Zealand from Australia, and implies two long‐distance dispersal events: a counter‐current movement across the Tasman Sea from New Zealand to Australia, occurring at the origination of the clade, followed by the colonization of Norfolk Island. The biology of Cominella suggests that the most likely method of long‐distance dispersal is rafting as egg capsules. Our robust phylogeny also means that the current Cominella classification requires revision. We propose that our clades be recognized as subgenera: Cominella (s.s.), Cominista, Josepha, Cominula, and Eucominia, with each subgenus comprising only of New Zealand or Australian species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 315–332.     

FACTORS INFLUENCING THE REPRODUCTIVE SUCCESS OF IPOMOEA PES-CAPRAE (CONVOLVULACEAE) AROUND THE GULF OF MEXICO

The reproductive biology of Ipomoea pes-caprae, a pantropical beach morning glory, was studied at five sites around the Gulf of Mexico. The primary pollinators were Xylocopa species (carpenter bees) which dispersed pollen up to 90 m. Exclusion experiments demonstrated that ants feeding on extrafloral nectaries increased seed set but did not protect seeds from predation by the bruchid beetle Megacerus. The water-dispersed seed and long-range dispersal of pollen may function to counter the sporophytic incompatability mechanism of the plant; populations exhibit a large neighborhood size. Key reproductive factors in the life cycle of I. pes-caprae are the long-range pollen flow and mass germination of water-dispersed seeds.     

Evidence for mutualism between a flower-piercing carpenter bee and ocotillo: use of pollen and nectar by nesting bees

Abstract.

• 1 Carpenter bees (Xylocopa californica arizonensis) in west Texas, U.S.A., gather pollen and ‘rob’ nectar from flowers of ocotillo (Fouquieria splendens). When common, carpenter bees are an effective pollen vector for ocotillo. We examined ocotillo's importance as a food source for carpenter bees.
• 2 The visitation rate of carpenter bees to ocotillo flowers in 1988 averaged 0.51 visits/flower/h and was 4 times greater than that of queen bumble bees (Bombus pennsylvanicus sonorus), the next most common visitor. Nectar was harvested thoroughly and pollen was removed from the majority of flowers. Hummingbird visits were rare.
• 3 Pollen grains from larval food provisions were identified from sixteen carpenter bee nests. On average, 53% of pollen grains sampled were ocotillo, 39% were mesquite (Prosopis glandulosa), and 8% were Zygophyllaceae (Larrea tridentata or Guaiacum angustifolium). Carpenter bee brood size averaged 5.8 per nest.
• 4 We measured the number of flowers, and production of pollen and nectar per flower by mature ocotillo plants, as well as the quantity of pollen and sugar in larval provisions. An average plant produced enough pollen and nectar sugar to support the growth of eight to thirteen bee larvae. Ocotillo thus has the potential to contribute significantly to population growth of one of its key pollinators.
• 5 Although this carpenter bee species, like others, is a nectar parasite of many plant species, it appears to be engaged in a strong mutualism with a plant that serves as both a pollen and as a nectar source during carpenter bee breeding periods.