Nomenclatural changes in <Emphasis Type="Italic">Lithospermum</Emphasis> (Boraginaceae) and related taxa following a reassessment of phylogenetic relationships

Lithospermum (Boraginaceae) comprises approximately 40 species in both the Old and New Worlds, with a center of diversity in the southwestern United States and Mexico. Using ten cpDNA regions, a phylogeny of Lithospermum and related taxa was reconstructed. Lithospermum (including New World and Old World species) and related New World members of Lithospermeae form a monophyletic group, with Macromeria, Onosmodium, Nomosa, Lasiarrhenum, and Psilolaemus nested among species of Lithospermum. New World Lithospermeae also is a monophyletic group, with Eurasian species of Lithospermum sister to this group. Because Lithospermum is not monophyletic without the inclusion of the other New World genera, species from these genera are transferred to Lithospermum, and appropriate nomenclatural changes are made. New combinations are Lithospermum album, Lithospermum barbigerum, Lithospermum dodrantale, Lithospermum exsertum, Lithospermum helleri, Lithospemum leonotis, Lithospermum notatum, Lithospermum oaxacanum, Lithospermum pinetorum, Lithospermum rosei, Lithospermum trinverium, and Lithospermum unicum; new names are Lithospermum chiapense, Lithospermum johnstonii, Lithospermum macromeria, Lithospermum onosmodium, Lithospermum rzedowskii, and Lithospermum turneri.     

Resource availability and population dynamics of Nicrophorus investigator, an obligate carrion breeder

1. Food resources for rearing young may influence insect populations. This is particularly true for insects that breed obligately on rare, ephemeral resources such as dung, fungi, or carrion. 2. Beetles in the genus Nicrophorus bury small vertebrate carcasses for rearing their young. Studies reviewed by Scott (1998) have found a positive relationship between carcass mass and total brood size. It is likely that access to carcasses suitable for breeding, and not food or mates, limits reproduction in both male and female Nicrophorus. Thus, small mammal densities could determine Nicrophorus population sizes. 3. The work reported here examined the relationship between Nicrophorus investigator (Coleoptera: Silphidae) population size and small mammal abundance at two sites over a 4‐year period. 4. Nicrophorus investigator buried and reared young on all the native small rodent species trapped at two sites in south‐western Colorado, U.S.A. (Peromyscus maniculatus, Microtus montanus, Zapus princeps, Tamias minimus, Thomomys talpoides). They preferred to bury and reproduce on rodent carcasses weighing between 16 and 48 g; rodents of this size represented 82% of captures. 5. Population sizes of N. investigator and small rodents were estimated simultaneously using mark‐recapture censuses over a 4‐year period. Considering only rodents within the size range used by N. investigator, the estimated small mammal biomass per hectare in one year and the beetle population size in the following year were correlated significantly.     

Molecular evidence for naturalness of genera in the tribe Antirrhineae (Scrophulariaceae) and three independent evolutionary lineages from the New World and the Old

The tribe Antirrhineae consists of 29 genera distributed in the New World and the Old. Phylogenetic analyses of ITS and ndhF sequences served to recognize six main lineages: Anarrhinum group (Anarrhinum, Kickxia); Linaria group (Linaria); Maurandya group (Cymbalaria, Asarina, Maurandella, Rhodochiton, Lophospermum); Schweinfurthia group (Pseudorontium, Schweinfurthia); Antirrhinum group (Antirrhinum, Pseudomisopates, Misopates, Acanthorrhinum, Howeliella, Neogarrhinum, Sairocarpus, Mohavea, Galvezia); Chaenorrhinum group (Chaenorrhinum, Albraunia, Holzneria). Parsimony (cladistics), distance-based (Neighbor-Joining), and Bayesian inference reveal that: (1) the tribe is a natural group; (2) genera such as Linaria, Schweinfurthia, Kickxia, and Antirrhinum also form natural groups; (3) three Antirrhineae lineages containing genera from the New and Old World are the result of three intercontinental disjunctions displaying similar levels of ITS-sequence divergence and differentiation times (Oligocene-Miocene); (4) evolution of flower shapes is not congruent with primitiveness of personate flowers; (5) both polyploidy and dysploidy appear to be responsible for most variation in chromosome number in the six main lineages. Nuclear and chloroplast evidence also supports the split of American and Mediterranean species of Antirrhinum into different genera, a result that should be contemplated in the interest of a more natural (monophyletic) taxonomy. Nucleotide additivity causes poor resolution in the ITS analysis of 22 species of Mediterranean Antirrhinum and lead us to interpret extensive hybridization in the Iberian Peninsula.This research was supported by the Spanish Dirección General de Investigación Científica y Técnica (DGICYT) through the project Flora Iberica (PS91-0070-C03-01). We thank Jorge Martínez and Emilio Cano for his assistance in the lab, Rafael Rubio and Omar Fiz for helping on the Bayesian inferences, the curators/collection managers of BCF, E, JACA, MA, SALA, SALAF, SEV, TARI, VAL for loans and access to specimens, and two anonymous reviewers for improving the quality of this publication.     

Chloroplast DNA restriction site variation in theVernonieae (Asteraceae), an initial appraisal of the relationship of New and Old World taxa and the monophyly ofVernonia

Chloroplast DNA restriction site variation was examined for 35 taxa in theVernonieae and four outgroup tribes, using 17 restriction enzymes mapped for ca. 900 restriction sites per species; 139 mutations were found to be phylogenetically informative. Phylogenetic trees were constructed using Wagner and weighted parsimony, and evaluated by bootstrap and decay analyses. Relationships of Old and New World taxa indicate complex geographical relationships; there was no clear geographic separation by hemisphere. The relationships between Old and New World Vernonias found here support prior morphological analyses. The sister group to all New and most Old World taxa was composed of a small group of Old World species including yellow-flowered, trinervate-leaved species previously postulated to be basal in the tribe. The majority of both New and Old World taxa are derived from a lineage beginning with the monotypic genusStokesia, an endemic of the southeastern United States. The genusVernonia was also found to be paraphyletic within both the New and Old World. Available data do not support either the separation ofVernonia or the tribeVernonieae into geographically distinct lineages. The pattern of relationships within theVernonieae for taxa from North America, Asia, Africa, Central and South America is most similar to that of several other groups of both plants and animals with a boreotropical origin, rather than an origin in Gondwanaland. Such a pattern of distribution suggests more ancient vicariant events than are routinely postulated for theAsteraceae.     

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.     

Evidence for Gondwanan vicariance in an ancient clade of gecko lizards

Aim Geckos (Reptilia: Squamata), due to their great age and global distribution, are excellent candidates to test hypotheses of Gondwanan vicariance against post‐Gondwanan dispersal. Our aims are: to generate a phylogeny of the sphaerodactyl geckos and their closest relatives; evaluate previous phylogenetic hypotheses of the sphaerodactyl geckos with regard to the other major gecko lineages; and to use divergence date estimates to inform a biogeographical scenario regarding Gondwanan relationships and assess the roles of vicariance and dispersal in shaping the current distributions of the New World sphaerodactyl geckos and their closest Old World relatives. Location Africa, Asia, Europe, South America, Atlantic Ocean. Methods We used parsimony and partitioned Bayesian methods to analyse data from five nuclear genes to generate a phylogeny for the New World sphaerodactyl geckos and their close Old World relatives. We used dispersal–vicariance analysis to determine ancestral area relationships among clades, and divergence times were estimated from the phylogeny using nonparametric rate smoothing. Results We recovered a monophyletic group containing the New World sphaerodactyl genera, Coleodactylus, Gonatodes, Lepidoblepharis, Pseudogonatodes and Sphaerodactylus, and the Old World Gekkotan genera Aristelliger, Euleptes, Quedenfeldtia, Pristurus, Saurodactylus and Teratoscincus. The dispersal–vicariance analysis indicated that the ancestral area for this clade was North Africa and surrounding regions. The divergence between the New World spaherodactyl geckos and their closest Old World relative was estimated to have occurred c. 96 Myr bp . Main conclusions Here we provide the first molecular genetic phylogenetic hypothesis of the New World sphaerodactyl geckos and their closest Old World relatives. A combination of divergence date estimates and dispersal–vicariance analysis informed a biogeographical scenario indicating that the split between the sphaerodactyl geckos and their African relatives coincided with the Africa/South America split and the opening of the Atlantic Ocean. We resurrect the family name Sphaerodactylidae to represent the expanded sphaerodactyl clade.     

Phylogenetic Framework for Trema (Celtidaceae)

We used ITS and trnL sequence data, analyzed separately and combined by MP, to explore species relationships and concepts in Trema (Celtidaceae), a pantropical genus of pioneer trees. Whether Trema is monophyletic or includes Parasponia is still unresolved. Three clades within Trema received moderate to high support, one from the New World and two from the Old World, but their relationships were not resolved. In the New World, specimens of T. micrantha formed two groups consistent with endocarp morphology. Group I, with smaller brown endocarps, is a highly supported clade sister to T. lamarckiana. Group II, with larger black endocarps, is poorly resolved with several subclades, including the highly supported T. integerrima clade. Both Old World clades contain Asian and African species, with three or more species in each region. Trema orientalis is not monophyletic: specimens from Africa formed a highly supported clade sister to T. africana, while those from Asia were sister to T. aspera from Australia.     

Phylogeny of the Celastreae (Celastraceae) and the relationships of Catha edulis (qat) inferred from morphological characters and nuclear and plastid genes

The phylogeny of Celastraceae tribe Celastreae, which includes about 350 species of trees and shrubs in 15 genera, was inferred in a simultaneous analysis of morphological characters together with nuclear (ITS and 26S rDNA) and plastid (matK, trnL-F) genes. A strong correlation was found between the geography of the species sampled and their inferred relationships. Species of Maytenus and Gymnosporia from different regions were resolved as polyphyletic groups. Maytenus was resolved in three lineages (New World, African, and Austral-Pacific), while Gymnosporia was resolved in two lineages (New World and Old World). Putterlickia was resolved as nested within the Old World Gymnosporia. Catha edulis (qat, khat) was resolved as sister to the clade of Allocassine, Cassine, Lauridia, and Maurocenia. Gymnosporia cassinoides, which is reportedly chewed as a stimulant in the Canary Islands, was resolved as a derived member of Gymnosporia and is more closely related to Lydenburgia and Putterlickia than it is to Catha. Therefore, all eight of these genera are candidates for containing cathinone- and/or cathine-related alkaloids.     

Differentiation in Searching Behavior for Carcasses Based on Flight Height Differences in Carrion Beetles (Coleoptera: Silphidae)

Flight ability allows insects to search for food and locate oviposition sites across a wide range of environments and, thus, allows insects to locate scarce and patchy resources such as carcasses. We examined differentiation in searching for carcasses, based on capture height differences in carrion beetles, and found that capture height differed among species. For Nicrophorus quadripunctatus and Oiceoptoma nigropunctatum, the height of abundance peaks differed among sites and was related to species composition at each site, indicating that shifts in flight height can occur plastically, whereas those for Nicrophorus maculifrons and Nicrophorus investigator did not differ among sites, indicating that the flight heights of these species are largely genetically determined. Thus, the present differentiation in capture height is primarily determined by current plastic shifts and on some genetic basis.     

Stelar structure ofAsplenium obtusifolium and its allied species in the New World tropics,with comparison to the Asian members ofAsplenium sect.Hymenasplenium

The stelar structure ofAsplenium obtusifolium and its related species (A. repandulum, A. riparium, A. triquetrum, A. volubile, A. purpurascens, A. ortegae, A. delitescens, A. hoffmannii, andA. laetum) in the New World tropics was observed and compared to that of Asian species ofAsplenium sect.Hymenasplenium. Both of the groups were found to share peculiar stelar structures: steles with two meristeles, a broader ventral and a narrower dorsal, each providing one of the two leaf traces; leaf gaps arranged in two rows between the dorsal and ventral strands, which are connected by thin meristeles, delimiting the leaf gaps. These structures are distinct from the radial symmetrical ones general inAsplenium. Together with cytological evidence, this strongly indicates that the New and Old World groups are closely related. Thus, these Neotropical species should be included in sect.Hymenasplenium.     

Oriole pollination ofErythrina breviflora (Leguminosae): Evidence for a polytypic view of ornithophily

Erythrina breviflora is visited by large numbers of passerine birds of which orioles (Icterus: Icteridae) are the primary pollinators. The flowers produce large quantities of nectar but they are rarely visited by hummingbirds. Inflorescence and floral morphology, and low levels of sucrose in the nectar probably explain the rarity of foraging hummingbirds. A comparison of Old WorldErythrina and their pollinators with New World species pollinated by orioles and hummingbirds suggests that parallel evolution has occurred. When the comparison is expanded to include other species pollinated by orioles, it is clear that various New WorldIcteridae, Thraupidae, etc. are ecological equivalents of Old WorldOriolidae, Pycnonotidae, Sturnidae, etc. and that flowers pollinated by these birds have similar characteristics.     

Molecular systematics of the Old World Astragalus (Fabaceae) as inferred from nrDNA ITS sequence data

This study represents a nuclear rDNA ITS-based phylogenetic analyses of a greater sampling of the Old WorldAstragalus compared to our previous work (212 vs. 134 taxa). Phylogenetic relationships among 212 species (213 accessions) of the Old WorldAstragalus, including newly segregated monotypic genusPodlechiella, the two aneuploid New WorldAstragalus, and five related genera, were inferred from analyses of nuclear rDNA ITS sequences using maximum parsimony. A total of 658 nucleotide sites and four binary characters for indels were analyzed. The results of phylogenetic analyses suggest sect.Phyllolobium, comprising mostly the Chinese species, is placed outside of the so-calledAstragalus s. str. and is a well-supported monophyletic group. The monotypic annual segregate genusThlaspidium (≡Astragalus sect.Thlaspidium, A. thlaspi), is clearly nested withinAstragalus s. str. Among the many sections analyzed here, only sects.Cenanthrum, Caraganella, Eremophysa, Incani, Laxiflori, andLotidium are strongly supported as monophyletic. Our analysis, in agreement with previous studies, shows that the North American euploidAstragalus species are scattered throughout the Old World groups of the genus.     

Observations on pollination inSarcostemma (Asclepiadaceae)

Field observations on pollination in New World species of the genusSarcostemma R. Br. sensuHolm are reported. Morphological and anatomical comparisons of pollinated flowers ofSarcostemma subg.Ceramanthus Kuntze (New World) andSarcostemma subg.Sarcostemma (Old World) are presented.     

MONOPHYLY OF ANEUPLOID ASTRAGALUS (FABACEAE): EVIDENCE FROM NUCLEAR RIBOSOMAL DNA INTERNAL TRANSCRIBED SPACER SEQUENCES

Evolutionary relationships within Astragalus L. (Fabaceae) were inferred from nucleotide sequence variation in nuclear ribosomal DNA of both New World and Old World species. The internal transcribed spacer regions (ITS) of 18S–26S nuclear ribosomal DNA from representatives of 26 species of Astragalus, three species of Oxytropis DC., and two outgroup taxa were analyzed by polymerase chain reaction amplification and direct DNA sequencing. The length of the ITS 1 region within these taxa varied from 221 to 231 bp, while ITS 2 varied in length from 207 to 217 bp. Of the aligned, unambiguous positions, approximately 34% were variable in each spacer region. In pairwise comparisons among Astragalus species and outgroup taxa, sequence divergence at these sites ranged from 0 to 18.8% in ITS 1 and from 0 to 21.7% in ITS 2. Parsimony analyses of these sequences resulted in a well-resolved phylogeny that is highly concordant with previous cytogenetic and chloroplast DNA evidence for a major phylogenetic division in the genus. These data suggest that the New World aneuploid species of Astragalus form a monophyletic but morphologically cryptic group derived from euploid species of Old World (Eurasian) origin, which are consequently paraphyletic.     

Phylogenetic implications of comparative primate growth rates

Growth data from a number of species of Old and New World primates have been analyzed by calculating instantaneous relative growth rates. Species discussed are the New World species Saimiri sciureus and Saguinus nigricollis, and the Old World species Pan troglodytes and Macaca mulatta. The analysis of the perinatal growth data indicated that differences in relative growth rates are present during early periods of growth. More specifically, it was found that the closer taxonomically a species is to man the greater the deceleration of growth during the first postnatal year. It is suggested that this may be a general primate trend.     

Integrative analyses of Nervilia (Orchidaceae) section Linervia reveal further undescribed cryptic diversity in Thailand

The delimitation of cryptic species is necessary to accurately classify and appropriately conserve biodiversity. Integrative analyses can be incisive in detecting and circumscribing cryptic diversity, especially in species complexes whose members are delineated by minor or overlapping morphological variation. We adopt an integrative approach to assess species relationships and resolve species boundaries in the taxonomically difficult Nervilia adolphi/punctata species alliance of N. sect. Linervia, an Old World complex of reduced, one-flowered terrestrial orchids that is both species-rich and poorly known in tropical and warm temperate Asia. We sampled 12 of the 27 known species of the alliance in Asia, including all four species reported from Thailand and a further 20 plants collected in that country that could not be satisfactorily identified using morphology alone. Phylogenetic analyses using one nuclear (ITS) and two plastid (matK and trnL-F) markers confirmed both N. sect. Linervia and the alliance itself as monophyletic, and corroborated 11 of the 12 sampled species; N. punctata proved polyphyletic, with the Thai samples referred to this Indonesian species falling sister to the Himalayan N. mackinnonii. The 20 unidentified Thai samples formed three distinct, strongly supported clades. STACEY, a Bayesian coalescence approach to species delimitation, resolved the same three clusters, but provided evidence suggesting that one comprised two distinct sub-clades. Building on this genetic evidence, we identify subtle morphological differences and invoke a diagnosable species concept to circumscribe three previously unrecognized cryptic species from Thailand. This objective approach to species delimitation validates ostensibly minor morphological differences as a basis for differentiating species within the alliance, paving the way for a global analysis of species boundaries throughout the genus as a whole.     

Plastome sequences of two New World bamboos—Arundinaria gigantea and Cryptochloa strictiflora (Poaceae)—extend phylogenomic understanding of Bambusoideae

• Premise of the study: Two New World species of Bambusoideae, Arundinaria gigantea and Crytpochloa strictiflora, were investigated in a phylogenomic context. Complete plastome sequences have been previously determined and analyzed for nine bambusoid species that exclusively represent Old World lineages. The addition of New World species provides more complete information on relationships within Bambusoideae. • Methods: Plastomes from A. gigantea and C. strictiflora were sequenced using Sanger methods. Phylogenomic and divergence estimate analyses were conducted on both species with 23 other Poaceae. • Key Results: Phylogenomic and divergence analyses suggested that A. gigantea diverged from within Arundinarieae between 1.94–3.92 mya and that C. strictiflora diverged as the sister to tropical woody species between 24.83 and 40.22 mya. These results are correlated with modern relative diversities in the two lineages. • Conclusions: The two New World bamboos show unique plastome features accumulated and maintained in biogeographic isolation from Old World taxa. The overall evidence for A. gigantea is consistent with recent dispersal, and that for C. strictiflora is consistent with vicariance.     

Historical evidence for a pre-Columbian presence of <Emphasis Type="Italic">Datura</Emphasis> in the Old World and implications for a first millennium transfer from the New World

Datura (Solanaceae) is a small genus of plants that, for long, was thought to occur naturally in both the New and Old Worlds. However, recent studies indicate that all species in the genus originated in the Americas. This finding has prompted the conclusion that no species of Datura could have been present in the Old World prior to its introduction there by Europeans in the early 16th century CE. Further, the textual evidence traditionally cited in support of a pre-Columbian Old World presence of Datura species is suggested to be due to the misreading of classical Greek and Arabic sources. As a result, botanists generally accept the opinion that Datura species were transferred into the Old World in the post-Columbian period. While the taxonomic and geographic evidence for a New World origin for all the Datura species appears to be well supported, the assertion that Datura species were not known in the Old World prior to the 16th century is based on a limited examination of the pre-Columbian non-Anglo sources. We draw on old Arabic and Indic1 texts and southern Indian iconographic representations to show that there is conclusive evidence for the pre-Columbian presence of at least one species of Datura in the Old World. Given the systematic evidence for a New World origin of the genus, the most plausible explanation for this presence is a relatively recent but pre-Columbian (probably first millennium CE) transfer of at least one Datura species, D. metel, into the Old World. Because D. metel is a domesticated species with a disjunct distribution, this might represent an instance of human-mediated transport from the New World to the Old World, as in the case of the sweet potato (Ipomoea batatas).     

Cuticular Extracts of Five Common Mantids (Mantodea: Mantidae) of the Eastern United States

We undertook a preliminary investigation of the cuticular extracts of five common mantid species in the eastern United States: Tenodera sinensis (Saussure), T. angustipennis (Saussure) and Mantis religiosa (Linnaeus) introduced from the Old World and Stagmomantis carolina (Johannson) and Bruneria borealis (Scudder), which are New World species. The major components of these mixtures were normal alkanes, predominately hentriacontane, or in the case of the parthenogenic species B. borealis, tritriacontane. Tricontanal was detected in the extracts of all five species, and smaller amounts of other aldehydes and n-tricontanol were detected in some species. Complex mixtures of methyl and dimethylalkanes also were present in these extracts. The composition of the cuticular hydrocarbons of these mantids may be an adaptation for reduction of evaporative water loss in these insects that inhabit open fields.     

Systematic implications of chloroplast DNA VARIATION IN Jaltomata and selected physaloid genera (Solanaceae)

Chloroplast DNA restriction site data were used to assess relationships among the solanaceous genera Jaltomata, Hebecladus. Old and New World Physalis, Chamaesaracha, Leucophysalis, Margaranthus, Nicandra, and Saracha, and to assess interspecific relationships within Jaltomata. Cladograms rooted with Nicotiana tabacum were constructed with Wagner and Dollo parsimony. Strict consensus trees indicate that Hebecladus originated from within Jaltomata; together these genera are monophyletic and constitute the recently circumscribed genus Jaltomata. There are two primary clades in Jaltomata: one a morphologically diverse group confined to western (largely Andean) South America, the Greater Antilles, and the Galapagos Islands; and the other a morphologically homogeneous group widely distributed from the southwestern United States to Bolivia. The controversial Leucophysalis viscosa, formerly treated as Jaltomata viscosa, is related to Leucophysalis, Physalis, Chamaesaracha, and Margaranthus; it does not group with any of the sampled species of Jaltomata. Physalis appears to be polyphyletic since P. alkekengi of the Old World branches off prior to a clade including Chamaesaracha, Margaranthus, and the two New World Physalis species sampled.