ANT DISPERSAL OF CALATHEA (MARANTACEAE) SEEDS BY CARNIVOROUS PONERINES (FORMICIDAE) IN A TROPICAL RAIN FOREST

Two tropical herbaceous plant species, Calathea microcephala and Calathea ovandensis, exhibit morphological adaptations for seed dispersal by ants. In the field, 21 ant species are attracted to the arils of the seeds. Previously known as predatory carnivores, the ponerine ants Odontomachus laticeps, O. minutus, Pachycondyla harpax, and P. apicalis carry the seeds toward their nests, behaving as though they were carrying prey. These ants remove the aril in the nest, and seeds without arils germinate more readily than seeds with arils. Ant foraging distances can account for seedling distributions. Myrmecochory may be far more common in tropical rain forests than has been previously suspected.     

BIOSYSTEMATICS OF RUELLIA TUBEROSA L. (ACANTHACEAE)

Ruellia tuberosa is a wide-ranging tropical, perennial herb that has been designated the type species for the large genus Ruellia. To clarify interspecific relationships, the R. tuberosa species complex, including R. nudiflora, R. malacosperma, and R. lorentziana, and R. tuberosa, has been proposed as the redefined and restricted genus Ruellia. All other Ruellia species would be transferred to segregate genera. Biosystematic investigations that included artificial hybridization experiments, herbarium comparisons, chromatographic analyses, and cytological studies demonstrated that R. tuberosa is genetically related to a number of North American temperate taxa and to certain other tropical species. The narrow definition of Ruellia cannot be supported from the biosystematic evidence at hand. Morphological comparisons suggest that R. tuberosa may be a primitive component in a broadened generic concept.     

Tropical Asian species show that the Old World clade of ‘spiny solanums’ (Solanum subgenus Leptostemonum pro parte: Solanaceae) is not monophyletic

The tropical Asian taxa of the species‐rich genus Solanum (Solanaceae) have been less well studied than their highly diverse New World relatives. Most of these tropical Asian species, including the cultivated brinjal eggplant/aubergine and its wild progenitor, are part of the largest monophyletic Solanum lineage, the ‘spiny solanums’ (subgenus Leptostemonum or the Leptostemonum clade). Here we present the first phylogenetic analysis of spiny solanums that includes broad sampling of the tropical Asian species, with 42 of the 56 currently recognized species represented. Two nuclear and three plastid regions [internal transcribed spacer (ITS), waxy, ndhF‐rpL32, trnS‐trnG and trnT‐trnF] were amplified and used to reconstruct phylogenetic relationships using maximum likelihood and Bayesian methods. Our analyses show that Old World spiny solanums do not resolve in a single clade, but are part of three unrelated lineages, suggesting at least three independent introductions from the New World. We identify and describe several monophyletic groups in Old World solanums that have not been previously recognized. Some of these lineages are coherent in terms of morphology and geography, whereas others show considerable morphological variation and enigmatic distribution patterns. Tropical Asia occupies a key position in the biogeography of Old World spiny solanums, with tropical Asian taxa resolved as the closest relatives of diverse groups of species from Australia and Africa.     

Historical biogeographical patterns of the species of Bursera (Burseraceae) and their taxonomic implications

Aim The plant genus Bursera, with 104 species of trees and shrubs, has been used as a model for biogeographical analyses because of its high species richness and large number of endemic species. The biogeographical patterns of Bursera and their implications for its phylogenetic classification are reviewed in order that some hypotheses on the historical biogeography of tropical Mexico can be proposed. Location Bursera is found in the south‐western USA, most of Mexico, mainly below 1700 m elevation in tropical forests, with some species in xeric shrublands, diversifying along the Pacific slope, Central America, and north‐western South America. A few species occur on the Galapagos and Revillagigedo archipelagos, some of which are endemics, whereas in the Antilles species are distributed extensively, with several endemics in the Bahamas, Cuba, Jamaica, and Hispaniola. Methods Data from specimens in herbaria and the literature were used to construct a matrix of 104 species in 160 areas. Distributional patterns of the species of Bursera were inferred applying track analysis, parsimony analysis of endemicity (PAE), and Brooks parsimony analysis (BPA). Results Track analysis revealed four individual tracks: (1) a circum‐Caribbean track, comprising species of the Bursera simaruba species group; (2) an Antillean track, including species that have been transferred to Commiphora based on their pollen traits; (3) a Mexican Pacific track, including species of the B. fragilis, B. microphylla, and B. fagaroides species groups, called ‘cuajiotes’; and (4) a Neotropical Pacific track, including the two species groups assigned to section Bullockia, in which the individual track of the Bursera copallifera species group is nested within the track of the B. glabrifolia species group. The four tracks overlap in a node in the Mexican Pacific slope, where they are highly diversified. PAE allowed us to identify 22 areas of endemism: 12 in Mexico (11 along the Mexican Pacific slope), six in the Antilles, two in Central America, one in South America, and one in the Galapagos. The general area cladogram obtained by BPA has two main clades: one includes the greater Antilles; and the other, 12 Mexican areas of endemism. Main conclusions Bursera fragilis, B. microphylla, and B. fagaroides species groups can be treated together as a new section within Bursera, sect. Quaxiotea, because they are segregated from the other groups of sect. Bursera based on morphological, anatomical, molecular and geographical evidence.     

MOLECULAR SYSTEMATICS OF THE GRACILARIACEAE (GRACILARIALES,RHODOPHYTA) WITH EMPHASIS ON SOUTHERN AFRICA1

Southern Africa has economically exploited populations of terete gracilarioids on the cool temperate west coast and numerous species of endemic and Indo‐Pacific tropical Gracilariaceae on the south and east coasts. Gross morphological characters have been the main means of identification, and incorrect applications have led to a number of misidentifications. In this study, small subunit rDNA and RUBISCO spacer sequences were used to determine phylogenetic relationships. Whereas rDNA sequences successfully differentiate major groups within the family as well as species belonging to the Gracilariopsis and the Curdiea/Melanthalia clade, RUBISCO spacer sequencing was required to distinguish between species of Gracilaria. The southern African gracilarioid complex (stringy, terete, elongate members of the Gracilariaceae) was resolved into three species: Gracilaria gracilis, Gracilariopsis longissima, and Gracilariopsis funicularis. South African Gracilaria protea was shown to be conspecific with tropical Indian Ocean G. corticata. Apart from G. gracilis and G. corticata, South African Gracilaria species were differentiated into a temperate‐tropical terete grouping and a temperate‐tropical flattened grouping.     

Taxonomy ofNeurolaena (Asteraceae-Heliantheae)

A taxonomic treatment of the tropical genusNeurolaena (tribeHeliantheae, subtribeGalinsoginae) is rendered. Ten species are recognized: the widespread, annual, weedy,N. lobata and nine relatively rare, relictual, shrubby or small tree species, mostly restricted to southern Mexico and Central America. Distributional maps, key to species, and complete synonymy are presented. A chromosomal and chemical review is also provided.     

The subgenus Scaptodrosophila (Diptera: Drosophilidae)

Four major subgenera, Drosophila, Sophophora, Hirtodrosophila and Scaptodrosophila, and several minor subgenera, have been recognized in the genus Drosophila. The hitherto little studied subgenus Scaptodrosophila, possessing a complicated synonymy, contains 144 named species, or about 11% of the total described in the genus; it is likely that many more Scaptodrosophila species await discovery. Scaptodrosophila probably originated in tropical Asia, and the greatest Scaptodrosophila faunas occur in Asia, south-east Asia, New Guinea, Australia and Africa, with very few species in north and south America and Europe. Scaptodrosophila species usually possess a pair of prescutellar bristles, a propleural bristle, and three large subequal sternopleural bristles; other morphological features are more variable. Ecological information concerning the majority of Scaprodrosophila species is scanty or absent, but known feeding or breeding sites include tree sap, fungi, fruit, flowers and 1eaf litter; larvae of at least one species are gall forming in plant tissues. The establishment of species groups is complicated by lack of detailed knowledge of many species, but six groups containing four or more species are recognized.     

Biogeographical contingency and the evolution of tropical anchovies (genus Cetengraulis) from temperate anchovies (genus Engraulis)

Aim Similar regimes of selection in different geographical settings can deterministically produce similar adaptive morphologies. We tested the hypothesis that the evolutionary trajectories of fish in upwelling zones can be altered by biogeographic contingencies in the biological and physical environment. Location Eastern Pacific and western Atlantic oceans. Methods We estimated phylogenetic relationships among eastern Pacific temperate anchovies (genus Engraulis) and tropical anchovies (genus Cetengraulis) with neighbour‐joining and Bayesian tree analysis of a 521‐bp segment of mitochondrial DNA cytochrome b. Available sequences for five additional engraulid taxa were included to establish polarity of the tree. Bayesian estimates (BEAST) of time to most recent common ancestor (TMRCA) for the nodes in the phylogeny were calibrated with divergence between Cetengraulis edentulus and Cetengraulis mysticetus precipitated by the rise of the Panama Isthmus 2.8–3.2 Ma. Results Neighbour‐joining and Bayesian trees indicate that South American Engraulis anchoita (Argentina) and Engraulis ringens (Chile) together are basal sister taxa to the California anchovy (Engraulis mordax) and Old World anchovies (Engraulis japonicas, Engraulis australis, Engraulis capensis and Engraulis encrasicolus). The two tropical species of Cetengraulis are sister‐taxa to Californian E. mordax, even though their phenotypes and ecologies differ markedly. A relaxed molecular clock indicates a TMRCA between Californian E. mordax and Cetengraulis at about 4.2 Ma (3.0–6.3 Ma 95% highest probability density). Main conclusions The TMRCA between the California anchovy, E. mordax, and tropical Cetengraulis coincides with the formation of the Gulf of California, which provided opportunities for allopatric isolation during climate oscillations. Mid‐Pliocene warming (3.1–2.9 Ma) may have trapped ancestors of Cetengraulis in the Gulf of California, where they evolved digestive tract morphologies to exploit inshore tropical habitats with low plankton productivities. While populations of several other temperate fishes have become isolated in the Gulf of California, few of these derived species show strong adaptive shifts from temperate sister taxa or range expansions into the tropical provinces of the western Atlantic and eastern Pacific.     

Drosophila parasitoids (Hymenoptera) of Japan

The Drosophila parasitoid system is an excellent model for the study of biological and ecological interactions. In Japan, extensive studies have been carried out on Drosophila parasitoids since the early 2000s. Here we provide up-to-date information on their diversity, distributions, host use and reproductive mode. Until now, 45 wasp species have been recorded. In addition, four genetically distinct groups (tentatively assigned as “races”) have been found in Ganaspis brasiliensis (Figitidae). Most of them are larvo-pupal parasitoids belonging to the families Figitidae and Braconidae, and some are pupal parasitoids of the families Pteromalidae and Diapriidae. A species of Encyrtidae was also recorded. Their major hosts are indicated with information on the host's breeding sites. Except Ganaspis xanthopoda and two races of G. brasiliensis that are distributed both in tropical and temperate regions, they are divided into tropical and temperate species. The tropical species are mainly distributed in Southeast Asia, and their distributions in Japan are restricted to the Ryukyu islands, particularly Iriomote-jima. The temperate species have been recorded from the main islands of Japan and surrounding islands but not from the Ryukyu islands, except two braconid species, Asobara japonica and Phaenocarpa sp. TK1. Some of the temperate species have also been recorded from Korea, China, Europe and/or North America. Among the 45 Japanese parasitoid species, at least seven are assumed to be thelytokous.     

Systematic study of the genusDelilia (Asteraceae,Heliantheae)

A taxonomic study of the genusDelilia is presented. Two species are recognized:D. biflora, a widespread weed of tropical and subtropical regions of the New World, extending from Mexico to northern Argentina and recently introduced in the Cape Verde Islands; andD. repens, endemic to the Galapagos islands. A key to the species, illustrations, distribution maps, and complete synonymy are presented.     

The Hiletini, an ancient and enigmatic tribe of Carabidae with a pantropical distribution (Coleoptera)

Abstract. Species of the tribe Hiletini are rarely represented in collections. Of the twenty known species, only three have been adequately collected. These twenty species are here arrayed in two genera, Hiletus (six species in two spe'cies groups), Eucamaragnathus (fourteen species in five species groups). Hiletus species occur in tropical Africa. Eucamaragnathus species of the alluaudi group occur in tropical and southeastern Africa, those of the suberbiei group are found only in Madagascar, those of the bocandei group are found in tropical western Africa, species of the sumatrensis group occur in southeastern Asia (Burma, Vietnam, Sumatra and Borneo), and those of the batesi group are found in the western Amazon Basin, northeastern Amazon Basin, and along the Paraquay River drainage system. Four new species of Hiletini are described from the following type localities: Hiletus nimba, GUINEA, Nimba Mountains; Eucamaragnathus borneensis, BRUNEI, (BORNEO); E.jaws, BRAZIL, Parana; E.amapa, BRAZIL, Territory of Amapa. Most species of Hiletini live in tropical climates with a mean annual temperature above 21°C and mean annual rainfall above 200 cm (exceptions are noted in text). All species apparently frequent latosolic soils in broadleaf evergreen or deciduous forests or in grassland savannahs with scattered or gallery broadleaf evergreen trees. Records available indicate that pupation occurs during the dry season and that adults emerge with the onset of rains. The structure of the mandibles, other mouthparts, crop and proventriculus suggests that only liquid food is normally taken in, and it is probable that at least some preoral digestion occurs as in other carabids. A suite of newly discovered character states associated with the tarsal claws unites the Hiletini with Cnemacanthini, Elaphrini, Migadopini, Promecognathini, Pseudomor-phini, Scaritini and Siagonini. These stocks together form a sister lineage to the ozaenine-brachinine lineage, all having distinct epimera, brushy non-styliform parameres, long empodial unguitractor plates, but not having conjunct mesocoxae (type I). Hiletini is the sister group of the combined Scaritini-Cnemacanthini-Pseudomorphini stocks. We suggest that the origin of the Hiletini occurred at least as early as the Jurassic Period from an equatorial position near the centre of the combined Africa/South America landmass. By mid-Cretaceous, radiation of taxa occurred across southern Laurasia into the southeastern part of that landmass, but never leaving equatorial climates. Later in the Cretaceous or early Tertiary, when continents began their rapid break-up, hiletines were stranded in tropical parts of South America, Africa/Madagascar, and southeastern Asia where they still occur today.     

Molecular phylogeny of the subgenus Holothuria (Selenkothuria) Deichmann, 1958 (Holothuroidea: Aspidochirotida)

The subgenus Selenkothuria comprises 12 species of tropical shallow water sea cucumbers that share morphological features, such as rods in the body wall and tube feet, modified tentacles for suspension feeding, and cryptic colours. The taxonomic status of this taxon has been controversial, but currently it is accepted as a subgenus of the genus Holothuria. Phylogenetic analyses of mitochondrial genes [cytochrome c oxidase subunit 1 (COI), 16S RNA] of ten species of Selenkothuria and related subgenera showed the polyphyly of this subgenus; monophyly was rejected by a likelihood ratio test. A geographical split divides the species of this subgenus into three different groups: one Indo‐West‐Pacific (IWP) group and two American groups. The IWP group is more closely related to Holothuria (Semperothuria) cinerascens and to other subgenera such as Roweothuria, Holothuria, and Vaneyothuria, whereas the two American groups are more closely related to each other and to some species of the subgenus Halodeima. These results suggest multiple parallel originations and diversification of ossicle morphology within the subgenus Selenkothuria. The current scheme of subgenera for the genus Holothuria is not supported, suggesting the need for a new classification. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 109–120.     

A synopsis of <Emphasis Type="Italic">Asclepias</Emphasis> (Apocynaceae: Asclepiadoideae) in tropical Africa

In preparation for two major regional Flora accounts, Asclepias has been reassessed in tropical Africa. 38 species not assigned to the allied genera Aspidoglossum, Glossostelma, Gomphocarpus, Margaretta, Pachycarpus, Stathmostelma, Stenostelma or Xysmalobium are recognised in the region. Nine species and one subspecies are described for the first time. The inclusion of Trachycalymma again within Asclepias requires four new combinations. A further three new names or combinations are required for the transfer of Odontostelma welwitschii, Schizoglossum alpestre and Stathmostelma verdickii to Asclepias. Five names are lectotypified, and a neotype proposed for Gomphocarpus meliodorus. The conservation status of all species is assessed.     

The Miocene Architectonicidae (Gastropoda) of Chile

Three new species of Architectonicidae Heliacinae,Heliacus (Torinista) chonos, Heliacus (Grandeliacus) antu andSolatisonax bieleri, are described. These include the first reports ofGrandeliacus andSolatisonax from Chile.Solarium australe Philippi, 1887 is an earlier name forHeliacus (Torinista) bahamondei Frassinetti & Covacevich, 1981, but is a junior homonym ofS. australe Philippi, 1849. Type material and new specimens of the five previously described architectonicid species from Chile are described and figured for comparison. Together with other gastropod taxa from the same deposits, Architectonicidae provide evidence for tropical to subtropical water temperatures in central Chile during the early Miocene.     

Biogeography and conservation of the genus Ficus (Moraceae) in Mexico

Aim The main objective of this study is to document the biogeographical patterns, endemism and degree of conservation of the species of Ficus (Moraceae) in Mexico. There are over 750 species of the genus Ficus distributed worldwide, and Mexico practically represents its northernmost limit in the American continent. Detailed studies at regional scales may help to understand the biogeography of large genera such as Ficus. Location Mexico. Methods The biogeographical patterns of Mexican Ficus were obtained from information of fig specimens available in two of the main herbaria of Mexico (2140 vouchers), collecting figs throughout this country, and revising the specialized literature. The presence of each species of Ficus was recorded for every one of Mexico's states and several tropical countries of America. Besides, the Mexican territory was divided into cells of 1° × 1° and the presence or absence of all species of the genus was recorded. Rarity of species was classified based on the width of geographic distribution, habitat specificity and population size. Results A total of 21 species of Ficus occur in Mexico, including six species (28.6%) that are endemic to this country. Five species are included in subgenus Pharmacosycea and 16 species are documented under subgenus Urostigma. Affinities of Ficus flora with other tropical countries in America generally decreased as geographical distances from Mexico increased. Mexican states and cells with highest values of Ficus species richness (both total and endemic species) were located. Ten species, including three endemics, presented a wide distribution. Five species, including two endemics, possess the three attributes of rarity (narrow geographical distribution, high habitat specificity and scarce local populations). Three species of Ficus, including the endemic and very rare Ficuslapathifolia (Liebm.) Miq., are not recorded in any protected area existing in Mexico. Main conclusions Most of the Mexican Ficus show a great morphological variation and occupy different habitats along their geographic distribution. The biogeographical patterns described here establish a fundamental scenario for ongoing studies on Ficus–pollinator interactions. However, many local populations are considered to be at risk, as there have been significant reductions in the number and size of local populations. Further studies are needed to understand the process of colonization, maintenance and persistence of fig–pollinator mutualism in species with different patterns of geographic distribution. Mexican Ficus require special policies for conservation due to their complex degree of rarity, particularly their geographic distribution in different types of vegetation, ranging from dry scrublands to tropical rain forests.     

UPPER ORDOVICIAN BRYOZOANS OF THE PIN FORMATION (SPITI VALLEY, NORTHERN INDIA)

Abstract: Twenty‐nine species of bryozoans from the Upper Ordovician–Lower Silurian Pin Formation (Spiti, India) have been identified. Eight of these are new: Trematopora minima, Ulrichostylus bhargavai, Ptilodictya exiliformis, Phaenopora ordinarius, Oanduellina himalayaica, Pesnastylus? vesiculosum, Ralfina? originalis and Pinocladia triangulata. The fossil record and facies analyses of the area investigated indicate shallow‐water conditions within the subtropical–tropical realm. The distribution pattern of fossils among the Ordovician/Silurian succession on the Northern Gondwana shelf and the influence of the Late Ordovician cooling phases on marine organisms are distinctive owing to a dramatic reduction in diversity globally. As far as the bryozoan taxa of Spiti are concerned, only one (Helopora fragilis) of the 29 species was recorded above the Ordovician/Silurian boundary. Observed bryozoan communities are very similar to faunas of Laurentia, the Baltic, Siberia and southern China of early–late Ordovician age.     

A monograph of the betel nut palms (Areca: Arecaceae) of East Malesia

The palm genus Areca is widespread in tropical Asia and includes the economically important betel nut palm, A. catechu. The genus has three centres of high species diversity: the Sunda Region, the Philippines and East Malesia (to the east of Wallace's line). The taxonomy of the genus in East Malesia has been neglected. Prior to this study, 19 species were accepted for this area, all but one endemic, but their limits and differences were not understood. Here, we provide a taxonomic monograph of East Malesian Areca spp., based on an extensive study of the genus in herbaria and in the field. We recognize six species of Areca in East Malesia, including the widespread cultivated A. catechu. Five wild species are accepted, namely A. macrocalyx, A. mandacanii, A. novohibernica, A. oxycarpa and A. vestiaria. We place 12 of the previously accepted species into synonymy, and provide additional new synonymy in A. catechu. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 147–173.     

Dormancy cycles in buried seeds of three perennial Xyris (Xyridaceae) species from the Brazilian campo rupestre

• Dormancy cycles are an important mechanism for avoiding seed germination under unfavourable periods for seedling establishment. This mechanism has been scarcely studied in tropical species. Here, we studied three tropical and perennial species of Xyris, X. asperula, X. subsetigera and X. trachyphylla, to investigate in situ longevity and the existence of seasonal seed dormancy cycles.
• Seeds of three species of Xyris were buried in their natural habitat, with samples exhumed bimonthly for 18 months. Germination of exhumed seeds was assessed under a 12‐h photoperiod over a broad range of temperatures. Seeds of X. trachyphylla were also subjected to treatments to overcome secondary dormancy.
• Seeds of all species are able to form a persistent seed bank and exhibit seasonal changes in germinability. Secondary dormancy was acquired during the rainy summer and was overcome during the subsequent dry season (autumn/winter). Desiccation partially overcomes secondary dormancy in X. trachyphylla seeds.
• Soil seed bank persistence and synchronisation of seed germination under favourable conditions for seedling establishment contribute to the persistence and regeneration of X. asperula, X. subsetigera and X. trachyphylla in their natural environment.