A seasonal evergreen forest in Belize: unusually high tree species richness for northern Central America

Most of the undisturbed forests of northern Central America remain undescribed. Some studies predict that tree species richness in lowland forests of northern Central America should be much lower than in similar forest types in southern Central America. This paper describes the physical and biological structure of two permanent, one-hectare plots on a valley floor in the Bladen Nature Reserve, Belize and compares these plots with other forests in the Neotropics. The plots have 91 and 89 species of trees ≥ 10 cm diameter (327 and 358 individuals, respectively) and comparable or higher measures of alpha-diversity than other such forests in Central America. The recorded tree species richness from other forests in northern Central America may be lower than those in southern Belize. The plots are dominated by Mortoniella pittieri Woodson (Apocynaceae), a disjunct from Costa Rica and Nicaragua. More than 50% of the tree species have distributions ranging into Bolivia or Peru, and >75% are wide-ranging species. The plots have a lower tree density and a higher proportion of large trees (>70 cm dbh) than other wet Neotropical forests. Like other Neotropical forests, tree species in the plots are generally aggregated and have a high proportion of vertebrate-dispersed fruits. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 275–296.     

Diversity, flower visitation frequency and generalism of pollinators in temperate rain forests of Chiloé Island, Chile

Species richness and taxonomic composition of pollinator assemblages are documented for 26 plant species from temperate rain forests of northern Chiloé Island, southern Chile (42°30'S). We investigated the patterns of generalism and specialization among plants and animal pollinators by comparing the flower visit frequency by different pollen vectors during the spring and summer months of three consecutive years (2000–2002). Species studied exhibited a range of floral morphologies (radial vs. zygomorphic, open vs. tubular) and rewards (nectar and/or pollen). Overall, we recorded 172 pollinator species, with an average of 6.6 species of pollen vectors/plant species. Pollinators visited an average of 15.2 plant species/pollen vector. Pollinator assemblages were dominated by Coleoptera (75 species), Diptera (56 species) and Hymenoptera (30 species), but passerine birds and hummingbirds were also important. The most specialized plants were vines, including the bee-pollinated genus Luzuriaga (Philesiaceae) and two endemic species of hummingbird-pollinated Gesneriaceae. Hymenoptera contributed 41.2% of all visits, with the bumblebee Bombus dalhbomii accounting for 22.5% of these. Plants with unspecialized flower morphology supported a higher species richness of pollinators, but visiting rates did not differ from specialized flowers.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 399–416.     

A multidisciplinary approach to discriminating different taxa in the species complex Pachycondyla villosa (Formicidae)

A multidisciplinary approach provides new evidence that strongly supports the differentiation of three taxa previously confounded in a single species, Pachycondyla villosa . All specimens studied were collected in the same location at Ilhéus, Bahia, Brazil. Bivariate plots of petiole heights vs. petiole lengths and isozyme patterns clearly differentiate the three taxa. Chain lengths and chemical structures of cuticular hydrocarbons are characteristic of each taxon. This congruent evidence shows that there are three different valid species. Ecological hypotheses are discussed to explain the benefits of these species remaining sympatric. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 249–259.     

Species richness and structure of three Neotropical bat assemblages

We compared the assemblages of phyllostomid bats in three Neotropical rainforests with respect to species richness and assemblage structure and suggested a method to validate estimates of species richness for Neotropical bat assemblages based on mist-netting data. The fully inventoried bat assemblage at La Selva Biological Station (LS, 100 m elevation) in Costa Rica was used as a reference site to evaluate seven estimators of species richness. The Jackknife 2 method agreed best with the known bat species richness and thus was used to extrapolate species richness for an Amazonian bat assemblage (Tiputini Biodiversity Station; TBS, 200 m elevation) and an Andean premontane bat assemblage (Podocarpus National Park; BOM, 1000 m elevation) in Ecuador. Our results suggest that more than 100 bat species occur sympatrically at TBS and about 50 bat species coexist at BOM. TBS harbours one of the most species-rich bat assemblages known, including a highly diverse phyllostomid assemblage. Furthermore, we related assemblage structure to large-scale geographical patterns in floral diversity obtained from botanical literature. Assemblage structure of these three phyllostomid assemblages was influenced by differences in floral diversity at the three sites. At the Andean site, where understorey shrubs and epiphytes exhibit the highest diversity, the phyllostomid assemblage is mainly composed of understorey frugivores and nectarivorous species. By contrast, canopy frugivores are most abundant at the Amazonian site, coinciding with the high abundance of canopy fruiting trees. Assemblage patterns of other taxonomic groups also may reflect the geographical distribution patterns of floral elements in the Andean and Amazonian regions.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 617–629.     

The impact of redefined species limits in Palame (Coleoptera: Cerambycidae: Lamiinae: Acanthocinini) on assessments of host, seasonal, and stratum specificity

Assumptions about the host and stratum specificity of tropical insects are routinely incorporated into estimates of global species richness, but few empirical studies reliably assess tropical insect specificity. In French Guiana, cerambycid beetles in the genus Palame reproduce exclusively in trees belonging to the Brazil nut family (Lecythidaceae). During a year-long rearing project, Palame anceps (Bates) emerged exclusively from a single host species, but P. crassimana Bates and P. mimetica Monné appeared to make seasonal changes in host affiliation. In the case of P. crassimana , dry season specimens could be distinguished from rainy season specimens by subtle differences in pubescence, but it was difficult to find diagnostic morphological characters supporting their delimitation as separate species. In order to detect genetic differentiation in Palame , 1049 bp of mitochondrial DNA (COI gene) were sequenced. The three nominal species of Palame yielded six haplotype groups, with sequence divergences among groups ranging from 8.27 to 17.56%. Species with apparent seasonal changes in host use included multiple distinct haplotype groups that I interpret to represent multiple species-level taxa. Insects considered generalized owing to temporal or geographical changes in host association may, with improved resolution in species limits, prove to be assemblages including relatively specialized cryptic species. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 195–209.     

Latitudinal and altitudinal diversity patterns and Rapoport effects in north-west European land snails and their causes

The latitudinal and altitudinal range sizes of north-west European land-snail species increase with increasing latitude/altitude. These Rapoport effects are not caused by northern/high-altitude species with wider latitudinal/altitudinal ranges and southern/low-altitude species with narrower latitudinal/altitudinal ranges, as predicted by the climatic variability hypothesis. They are instead caused mainly by different northern/upper borders of species occurring in the south part of the study area or at low and intermediate altitudes, respectively. This pattern indicates that the observed Rapoport effects are the result mainly of differential northward/upward expansion of species that were restricted to southern/low or intermediate altitude refugia during the glacials. Although all species occurring in a refugium experienced the same climatic conditions, there is stochastic variation in their climatic tolerance. Species with broader climatic tolerance were able to expand farer northwards/upwards postglacial. The altitudinal distribution of species richness in the analysed alpine faunas cannot be explained by the Rapoport-rescue hypothesis, because species richness peaks at intermediate altitudes and because there is no negative correlation between the number of range borders and altitude. The Rapoport-rescue hypothesis alone is probably also insufficient to explain the decrease in species richness with increasing latitude.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 309–323.     

Continental scale patterns of biodiversity: can higher taxa accurately predict African plant distributions?

The distribution maps of a total of 3563 species, which represent 8.9% of the known African angiosperm flora, were entered into cells representing a one-degree latitude–longitude grid of Sub-Saharan Africa. The computer programme WORLDMAP was used to explore continental scale patterns of biodiversity. The maps were used to assess the use of higher taxa as a surrogate measure for predicting patterns of species richness. Genera were found to predict species richness distributions most closely, with higher taxa (families, orders, subclasses) exhibiting progressively worse correlations. However in two areas, the Cape Region of South Africa and coastal Cameroon, there was a higher species to genus ratio than in other areas of Africa. In the Cape Region this meant that generic richness failed to predict species richness. Hotspots, defined as the 5% of grid cells with the highest scores for richness and range-size rarity, were identified for species and higher taxa. Whilst a high percentage of species richness hotspots were predicted by higher taxa, there were important exceptions like the Cape Region. Species range-size rarity hotspots were not well predicted by higher taxa. Hotspots of higher taxa (families and orders) do not therefore accurately predict the location of species hotspots. Higher taxa appear to provide a powerful and accurate tool that can be used to predict large scale patterns of species biodiversity in Sub-Saharan Africa. However care must be taken when using taxa higher than genera, especially if selecting areas of highest conservation priority. The special case of the Cape Region indicates the danger of extending predictive generalizations as the ecological mechanisms that promote and retain species may not be the same in all places. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 225–235.     

Testing historical explanations for gradients in species richness in heliconiine butterflies of tropical America

We compiled a large database of 58 059 point locality records for 70 species and 434 subspecies of heliconiine butterflies and used these data to test evolutionary hypotheses for their diversification. To study geographical patterns of diversity and contact zones, we mapped: (1) species richness; (2) mean molecular phylogenetic terminal branch length; (3) subspecies richness and the proportion of specimens that were subspecific hybrids, and (4) museum sampling effort. Heliconiine species richness is high throughout the Amazon region and peaks near the equator in the foothills and middle elevations of the eastern Andes. Mean phylogenetic terminal branch length is lowest in the eastern Andes and tends to be low in species‐rich areas. By contrast, areas of high subspecies richness, where subspecies overlap in range and/or hybridize, are concentrated along the course of the Amazon River, with the eastern Andes slopes and foothills relatively depauperate in terms of local intraspecific phenotypic diversity. Spatial gradients in heliconiine species richness in the Neotropics are consistent with the hypothesis that species richness gradients are driven at least in part by variation in speciation and/or extinction rates, resulting in observed gradients in mean phylogenetic branch length, rather than via evolutionary age or niche conservatism alone. The data obtained in the present study, coupled with individual case studies of recently evolved Heliconius species, suggest that the radiation of heliconiine butterflies occurred predominantly on the eastern slopes of the Andes in Colombia, Ecuador, and Peru, as well as in the upper/middle Amazon basin. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 479–497.     

Pollen morphology and phylogenetic relationships in neotropical Phyllanthus (Euphorbiaceae)

In order to provide new insights into phylogenetic relationships among the neotropical taxa of Phyllanthus , 28 illustrations are provided of the pollen grains of 22 selected species studied from 11 sections of the subgenera represented in the neotropics. Special attention has been given to subgenus Conami because of its variability in pollen morphology: of eight species illustrated, the apertures are diploporate colpi in three species and pores in five species; exine ornamentation is vermiculate in two species and pilate in the other six species. The six species in the neotropical sections Pityrocladus and Microglochidion (subgenus Emblica ) are characterized by prolate grains with an increased number of colpi (4–8). Of particular interest are species in which the pollen exine is clypeate (with exine shields); clypeate pollen grains are illustrated in two species of subgenus Xylophylla and in one species of section Cyclanthera that has unique exine shields with single central pila. The pollen of the one Brazilian phylloclade-bearing species illustrated (in section Choretropsis ) has 3-colporate grains with reticulate exine, typical for subgenus Phyllanthus , and very different from the clypeate grains of the West Indian phylloclade-bearing species in section Xylophylla . This pollen evidence clearly demonstrates homoplasy in the origin of phylloclades in Phyllanthus . Pollen morphological data suggest that the neotropical taxa of Phyllanthus have arisen following colonization from Africa (subgenus Kirganelia ) and Asia (subgenus Emblica ). © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 325–338.     

Improving representativeness of genebank collections through species distribution models, gap analysis and ecogeographical maps

An efficient germplasm collecting method was evaluated using six Lupinus species and the Spanish Lupinus collection as a study case. This method includes the application of geographic information systems, ecogeographical land characterization maps, species distribution models and gap analysis to identify prioritized collecting sites. To evaluate the efficiency of this collecting method, field collecting expeditions were carried out focusing on prioritized sites and the results of these collections were analyzed. Prioritized sites were identified using spatial and ecogeographical gaps, and potential species richness maps. The spatial gaps corresponded to populations non-included in the collection but recorded by other information sources while ecogeographical gaps corresponded to spatial gaps that were located in ecogeographical categories (obtained from the ecogeographical map) that were scarcely represented in the collection. A potential Lupinus species richness map was obtained by adding the information of single maps of Lupinus species distribution models. Subsequently, prioritized sites were obtained in ecogeographical gaps with high potential species richness values. Collecting expeditions were made in Spain in 2006, 2007 and 2008. Results showed that using the efficient germplasm collecting methodology was highly positive not only from a quantitative viewpoint (between 7.8 and 11% increase) but also in qualitative terms, focusing collection efforts in ecogeographical categories with low or null representation in the Spanish Lupinus collection (41% of the new accessions). Phenotypic differences related to adaptation to environment were observed in the field between the populations that grow in low or null represented categories and those that grow in highly represented categories.     

Systematic and comparative anatomy of Cymbidieae (Orchidaceae)

Cymbideae comprise an assemblage of 28 genera nearly all of which are represented in this study. Their anatomy is relatively homogenous with the exception of Govenia , in which roots lack velamen and pseudobulb vascular bundles lack sclerenchyma, conditions that do not obtain in other genera. Marginal fibre bundles in leaves of Grammatophyllum and Porphyroglottis consist of clusters of thicker-walled, narrower, epidermis-facing fibres as well as thinner-walled, wider, mesophyll-facing fibres. This feature also occurs in some species of Maxillaria . Baculate tilosomes appear in the roots of a majority of genera in Cymbidieae, as they do in species of Maxillaria , confirming DNA analyses showing a close relationship between tribes Cymbidieae and Maxillarieae. Govenia is singled out both on anatomical and molecular grounds as being ill-placed in Cymbidieae. Cladistic analysis produces only a few tentative hypotheses of phylogenetic relationships among the 28 genera, showing that anatomical characters are of limited value in assessing affinities within this tribe. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 1–27.     

Phylogenetic patterns of diversification in a clade of Neotropical frogs (Anura: Aromobatidae: Mannophryne)

We used partial sequences of mitochondrial 16S and cytochrome oxidase I genes to perform a phylogenetic study of collared frogs (Anura: Aromobatidae: Mannophryne ), a genus endemic to Venezuela and the islands of Trinidad and Tobago. We analysed 1.2 kb from 13 of the 15 described species of Mannophryne . Maximum parsimony, maximum likelihood and Bayesian analyses support the monophyly of Mannophryne . Mannophryne consists of three deeply differentiated clades that split from each other in a relatively short period of time. The diversification of Mannophryne occurred well before the glacial-interglacial periods of the Quaternary. Our data support the taxonomic validity of M. olmonae , a species endemic to Tobago Island. Mannophryne olmonae is more closely related to the continental species Mannophryne riveroi than to the Trinidad island endemic Mannophryne trinitatis . As in most tropical clades of frogs, molecular evidence indicates that species richness in Mannophryne is largely underestimated and, consequently, current priorities for conservation are inadequate.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 185–199.     

Hymenaea mexicana sp. nov. (Leguminosae: Caesalpinioideae) from Mexican amber indicates Old World connections

Hymenaea mexicana sp. nov. is described in amber originating from mines of Chiapas, Mexico. The species is characterized by clawed showy petals with cordate bases and glabrous, verrucose ovaries with long hirsute hairs at their bases and margins on one side. This species is closely related to the extinct Hymenaea protera from Dominican amber, and together with it, is placed in the primitive section Trachylobium of the genus Hymenaea in the caesalpinoid legumes. The closest extant species to H. mexicana is H. verrucosa , a relict surviving in East Africa. These results challenge previous views that Mexican amber was produced by the South American species H. courbaril – H. intermedia , which belong to the advanced section of the genus.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 125–132.     

Late Miocene Carnivora from Chad: Lutrinae (Mustelidae)

We describe extensive Late Miocene fossil Lutrinae from Toros-Menalla, Chad. A minimum of four species are present in this fossiliferous area. Such a large number of species, diverse in size and dental morphology, is significant and unique in the fossil record of the subfamily Lutrinae in the Neogene of Africa. Two new taxa are described, Sivaonyx beyi sp. nov. and Djourabus dabba gen. nov. sp. nov. ; the two other species, which are represented by more fragmentary remains, are assigned to Lutrinae indet. aff. Torolutra sp. and Lutrinae indet. aff. Aonyx sp. Sivaonyx beyi is the best known of the species. It is represented by many dental remains and a subcomplete skeleton. Postcranial characters of this species are not particularly specialized. It is interpreted here as a terrestrial predator with poorly developed aquatic adaptations. Fossils of otters are known from ten different localities at Toros-Menalla, each of which yielded a single individual of one or two species. The presence of four morphologically distinct otters in the area is indicative of a palaeoenvironment with a relatively well-developed freshwater system of lakes and/or rivers.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 793–846.     

Historical legacies in the geographical diversity patterns of New World palm (Arecaceae) subfamilies

The extent to which species richness patterns of the major palm subfamilies in the Americas are controlled by lineage history was studied. Based on the fossil record, we suggest that the subfamily Coryphoideae has followed a boreotropical dispersal route into Central and South America, whereas Calamoideae (tribe Lepidocaryeae), Ceroxyloideae and Arecoideae have Gondwana/South America-biased histories. However, Arecoideae has been present and diverse in both South and Central America at least since the early Tertiary. We used regression analyses to evaluate the relative importance of environmental factors and spatial variables (as substitutes for historical or other non-environmental factors) as determinants of geographical variation in species richness for each subfamily. Given the different lineage histories, we hypothesized that: (1) coryphoid richness should be least strongly controlled by the modern environment and exhibit a strong non-environmental bias towards Central and North America, reflecting its boreotropical invasion route, (2) calamoid species richness should exhibit a non-environmental bias towards South America, reflecting its long African–South American history, and (3) arecoid species richness should be most strongly environmentally determined, reflecting the long arecoid residency in both Central and South America. The regression analyses confirmed the hypothesized effects of lineage history on the geographical patterns in species richness. Hence, modern species richness patterns in the New World palm subfamilies strongly reflect their divergent biogeographical histories.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 113–125.     

Molecular evidence for the hybrid origin of a new endemic species of Stylosanthes Sw. (Fabaceae) from the Mexican Yucatán Peninsula

Stylosanthes aff. calcicola is a formally undescribed tetraploid species from the Mexican Yucatán Peninsula, showing morphological similarities to the diploid species S. calcicola , but distinct in a number of characters. We used uni- and biparentally inherited molecular markers to infer the hybrid origin of this species in relation to known diploid species of Stylosanthes . Molecular characterization was based on length and/or DNA sequence variation of nuclear sequence-tagged site (STS) markers, the internal transcribed spacer (ITS) region of nuclear rDNA and the trnL intron of chloroplast DNA (cpDNA). Stylosanthes aff. calcicola contains a distinct cpDNA haplotype and nuclear DNA fragment, with closest relationship to the diploid species S. calcicola . In contrast, the DNA sequences of two nuclear loci reveal a closer relationship to the diploid species S. angustifolia , S. hispida , S. humilis , S. leiocarpa and S. viscosa . The majority of the STS markers showed additivity of PCR fragments in S. aff. calcicola , representing the combination of two genetically different genomes. We postulate that S. aff. calcicola is a distinct species of allotetraploid origin that appears to have originated once from hybridization between two divergent genomes, of which the maternal and paternal parent are closely related to, or derived from, a member of the lineages represented by S. calcicola and S. viscosa , respectively.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 140 , 1–13.     

A new species of Acantholimon Boiss. (Plumbaginaceae) from Turkey

A new species, Acantholimon avanosicum Dogan & Akaydin in the Plumbaginaceae is described and illustrated. The species grows on argillaceous slopes in Central Anatolia, Turkey. The diagnostic morphological characters that distinguish it from closely related taxa are discussed. A revised key to Acantholimon species in Turkey with densely distichous simple spikes is presented. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 365–368.     

Phylogeny of a paradigm lineage: the Drosophila melanogaster species group (Diptera: Drosophilidae)

Although Drosophila melanogaster is a paradigm eukaryote for biology, relationships of this species and the other 174 species in the melanogaster species group are poorly explored and ambiguous. Gene regions of Cytochrome oxidase II (mt:CoII ), Alcohol dehydrogenase ( Adh ) and hunchback ( hb ) were sequenced and analysed phylogenetically to test prior hypotheses of relationships for the group based on chromosomes, morphology, and 28S rRNA gene sequences. A simultaneous cladistic analysis of the three newly sequenced gene regions produced a single well-resolved phylogeny for 49 exemplar species representing eight subgroups. Monophyly of each of the ananassae , melanogaster , montium , and takahashii subgroups is supported; the suzukii subgroup is polyphyletic. This phylogeny is consistent with variation in significant morphological structures, such as the male sex comb on the fore tarsus. The broad range of morphological variation among these species is interpreted and the applicability to evolution and developmental investigations is discussed. This phylogeny facilitates comparative investigations, such as gene family evolution, transposable element transmission, and evolution of morphological structures. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 21–37.     

Checklist of the Caesalpinioideae (Leguminosae) of Equatorial Guinea (Annobón, Bioko and Río Muni)

This study provides a checklist of the Caesalpinioideae (Leguminosae) present in Equatorial Guinea, comprising 52 genera and 124 taxa. Seven species are known from Annobón, 33 from Bioko and 109 from Río Muni. The best represented genus is Senna with eight species. In addition, bibliographic references for Caesalpinioideae (Leguminosae) from Equatorial Guinea have been gathered and checked. Fourteen species are included based on literature records, because their distribution ranges suggest they may occur in Equatorial Guinea, 11 introduced species could be naturalized, and 45 taxa are recorded for the first time from the country. This represents an increase of over 35% in the floristic knowledge of Caesalpinioideae from Equatorial Guinea. A statistical summary is presented at the end of the checklist.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 541–562.