Genetically distinct populations in an Asian soldier-producing aphid, Pseudoregma bambucicola (Homoptera: Aphididae), identified by DNA fingerprinting and molecular phylogenetic analysis

To estimate genetic structure of a soldier-producing aphid, Pseudoregma bambucicola, samples from natural populations throughout southeastern Asia were analyzed by a DNA fingerprinting technique. We unexpectedly found that P. bambucicola comprises two geographic groups, the northern group and the southern group, which are genetically distinct from each other but morphologically almost indistinguishable. Molecular phylogenetic and statistical analyses based on mitochondrial ribosomal DNA sequences demonstrated that the northern and southern groups of P. bambucicola are not closely related but constitute distinct lineages in the genus Pseudoregma. Detailed morphological reexamination revealed that the two groups could be distinguished by the number of setae on the 8th abdominal tergite of 1st instar nymphs and soldiers. From these results, it was suggested that P. bambucicola should be divided into two species. The northern group from Japan, Taiwan, Hong Kong, and northern Vietnam retains the name P. bambucicola, whereas we suggest that the name P. carolinensis (R. Takahashi, 1941, Tenthredo 3, 208-220) should be used for the southern group from Thailand, Malay Peninsula, Java, Irian Jaya, and Micronesia. The morphological resemblance between P. bambucicola and P. carolinensis might be due to shared ancestral characters of the genus Pseudoregma.     

Social structure and the defensive role of soldiers in a eusocial bamboo aphid,Pseudoregma bambucicola (Homoptera: Aphididae): A test of the defence-optimization hypothesis

A complete understanding of the evolution of sociality in aphids requires a detailed knowledge of the patterns of soldier investment in their ecology. The eusocial bamboo aphidPseudoregma bambucicola has a morphologically specialized first-instar soldier caste. The proportion of soldiers was positively correlated with colony size. Within a colony, soldiers were evenly distributed among subcolonies; within each subcolony, however, their distribution was biased toward peripheries which were exposed to many predators. Field experiments introducing natural enemies such asEupeodes confrater (Diptera: Syrphidae) andSynonycha grandis (Coleoptera: Coccinellidae) revealed that the survival rate of these predators was negatively correlated with the density of soldiers, suggesting that soldiers can more or less defend their colonies by killing or removing a range of natural enemies. Observations suggest that large mature colonies attract more predators than newly established small colonies and that, within a colony, the predators attack each subcolony regardless of its position on bamboo shoots. This implies the presence of a positive correlation between colony size and predation risk. Thus, the investment in soldiers seems to reflect the attacking pattern of predators within a colony. These results agree with the defence-optimization hypothesis in soldier investment ofP. bambucicola colonies.     

Social parasitism and behavioral interactions between two gall-forming social aphids

We investigated ecological and behavioral aspects of the interactions between two social aphids, Pseudoregma bambucicola and Astegopteryx bambucifoliae (Hormaphidinae, Cerataphidini), both of which produce second-instar, sterile soldiers in galls formed on Styrax suberifolius, in Taiwan. By censusing their galls, either species was found to invade galls of the other species. Twenty-eight (58%) out of 48 A. bambucifoliae galls contained P. bambucicola, while four (6%) of 69 P. bambucicola galls contained A. bambucifoliae. Furthermore, P. bambucicola behaved like social parasites in galls of A. bambucifoliae. Colonies of P. bambucicola produced much fewer or even no soldiers compared with those in natal galls. Our experiments also revealed that individual aphids of P. bambucicola more successfully intruded into galls of A. bambucifoliae than into conspecific galls, and that guarding soldiers of P. bambucicola effectively prevented aphids from invading their galls and permitted only conspecific soldiers to join their colonies. These differences in behavior provide good explanations for the differences in the frequency of invaded galls found between the two species. Received 4 September 2007; revised 7 January 2008; accepted 17 January 2008.     

Morphological observation and characterization of the Pseudoregma bambucicola with the scanning electron microscope

Both leica microscopic camera system and scanning electron microscopy was used to observe and characterize the feet, back, abdomen, antennae and mouthparts of the Pseudoregma bambucicola from the bamboo, Bambusa multiplex. The possible functions of all the external morphological characteristics of the P. bambucicola were described and discussed in detail, which offers a basis for further enriching the biology, phylogeny and ecological niche of the P. bambucicola. Moreover, the morphological results should contribute to morphological identification and differentiation of the P. bambucicola from other aphids in the same family.     

Soldier differentiation during embryogenesis of a social aphid, Pseudoregma bambucicola

To understand the developmental process of aphid soldier differentiation, we investigated the morphological characters of normal nymphs, soldier nymphs and developing embryos of Pseudoregma bambucicola. Results of morphometric analyses showed that normal and soldier nymphs formed discrete clusters on the basis of several morphological characters, although a small number of intermediate individuals, termed ‘intercaste nymphs’, were present. In late embryonic stages, normal and soldier embryos were morphologically distinguishable. The earlier the embryonic stage, the smaller the morphological differences between them. In early embryos less than 1000 µm in length, normal and soldier embryos were not morphologically distinguishable, suggesting that the onset of soldier differentiation occurs at an early embryonic stage. Throughout embryonic development, morphological differentiation of the soldier caste proceeded gradually. Notably, several morphological characters of soldiers grew remarkably upon larviposition. Observation of embryonic leg cuticle revealed a characteristic folding structure, indicating that some morphological traits of the soldier are exaggerated upon larviposition through expansion of the folded cuticle. We suggest that morphological differentiation of the soldier caste in P. bambucicola comprises two phases: gradual growth during embryogenesis and rapid growth upon larviposition.     

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

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

Phylogenetic relationships and taxonomy of subfamily Zygophylloideae (Zygophyllaceae) based on molecular and morphological data

Phylogenetic analysis of noncoding trnL plastid DNA sequences and morphological data for 43 species of Zygophylloideae, representing most of the morphological and geographical variation in the subfamily, indicates that the currently recognised genera Augea (monotypic, southern Africa), Tetraena (monotypic, China), and Fagonia (c. 30 species, widespread), are embedded in Zygophyllum (c. 150 species, widespread). A generic classification based on six monophyletic and morphologically distinctive entities is proposed: Roepera with c. 60 species in Australia and southern Africa, Zygophyllum with c. 50 species in Asia, Tetraena with c. 40 species in Africa and Asia, Augea with a single species in southern Africa, Melocarpum with two species in the Horn of Africa region, and Fagonia with c. 30 species in both the Old and the New World. Scanning electron microscopy studies of testa structure provided important characters for the delimitation of some genera. New combinations (61) are made in Roepera, a resurrected genus originally described from Australia, one new name is proposed in Zygophyllum, 35 new combinations are made in Tetraena, and two new combinations are made in Melocarpum (previously Zygophyllum sect. Melocarpum).     

Diversification in wild populations of the model organism Anolis carolinensis: A genome‐wide phylogeographic investigation

The green anole (Anolis carolinensis) is a lizard widespread throughout the southeastern United States and is a model organism for the study of reproductive behavior, physiology, neural biology, and genomics. Previous phylogeographic studies of A. carolinensis using mitochondrial DNA and small numbers of nuclear loci identified conflicting and poorly supported relationships among geographically structured clades; these inconsistencies preclude confident use of A. carolinensis evolutionary history in association with morphological, physiological, or reproductive biology studies among sampling localities and necessitate increased effort to resolve evolutionary relationships among natural populations. Here, we used anchored hybrid enrichment of hundreds of genetic markers across the genome of A. carolinensis and identified five strongly supported phylogeographic groups. Using multiple analyses, we produced a fully resolved species tree, investigated relative support for each lineage across all gene trees, and identified mito‐nuclear discordance when comparing our results to previous studies. We found fixed differences in only one clade—southern Florida restricted to the Everglades region—while most polymorphisms were shared between lineages. The southern Florida group likely diverged from other populations during the Pliocene, with all other diversification during the Pleistocene. Multiple lines of support, including phylogenetic relationships, a latitudinal gradient in genetic diversity, and relatively more stable long‐term population sizes in southern phylogeographic groups, indicate that diversification in A. carolinensis occurred northward from southern Florida.     

Logistic model for soldier production in aphids

Summary The number of aphid soldiers in a colony is positively correlated with colony size. This positive correlation has been repeatedly confirmed for the bamboo aphid Pseudoregma bambucicola. To explain this, we present a simple model assuming logistic growth of the aphid colony. The model predicts that a first soldier is more readily produced in large colonies than in small colonies. This is because the productivity of each reproductive decreases as the colony size increases, and also because the efficiency of defense increases as the number of reproductives defended by the soldier increases. The latter effect disappears when the number of reproductives exceeds N_p, the critical number of reproductives that a predator could damage. This argument holds for a second, a third, and an ith soldier in general. Although we assume logistic growth of the aphid colony, the model is applicable to any form of colony growth with minor changes of some premises.Received 12 February 2003; revised 28 March 2003; accepted 22 April 2003.     

Taxonomic, nomenclatural and chorological notes on several taxa of the genus Echinodorus (Alismataceae)

Taxonomic, nomenclatural and chorological notes on several taxa of the genus Echinodorus are given. A new species E. maculatus is described. The name Alisma intermedium (basionym of the name E. intermedius) is lectotypified. Echinodorus major and E. martii should be treated as two distinct taxa. The name E. xinguensis was invalidly published. It is also confirmed that E. africanus, E. veronikae and E. viridis (all belonging to the E. uruguayensis group) do not originate from Africa. The name E. bleherae should be written with a female suffix (-ae); this name is considered here as a synonym of the earlier name E. grisebachii.     

Phenetic investigation of non-nodulating African species ofAcacia (Leguminosae) using morphological and molecular markers

Morphological and RAPD markers were used to assess the relationships among nodulating and non-nodulating species of AfricanAcacia. Non-nodulating species of AfricanAcacia are only found within subg.Aculeiferum sect.Monacanthea. African species of sect.Monacanthea examined were found to form a group distinct from the other African species examined on a morphological and molecular basis. All lack the ability to nodulate, suggesting that non-nodulation may be used as a taxonomic tool. The species of sect.Aculeiferum were separated by RAPD and morphological analysis into two groups depending on whether they were armed with prickles in pairs and/or prickles in threes, or solitary. A third group of species was identified within sect.Acacia: further subdivision of this group was achieved into subsectt.Pluriseriae andUniseriae. The position ofA. albida relative to other AfricanAcacia species was found to be distinct but not totally independent of the genus. The partitioning and distribution of the genetic variability within the genus is further elucidated by the RAPD analysis of populations ofAcacia species. A population analysis ofA. polyacantha demonstrated geographical and site-specific variation.     

Morphological variation inLolium (Poaceae) as a measure of species relationships

Analysis of morphological and phenological data for determining the genetic variation within sevenLolium species led to the recognition of two groups within this genus. One group, containing the two inbreeding speciesL. temulentum andL. persicum, was clearly distinct from all other species. Strong morphological and phenological intergradation was found between both species. The cross-breeding species,L. perenne, L. rigidum, andL. multiflorum, formed another group. Little differentiation was found between these species, though they were distinct. Two inbreeding species,L. loliaceum andL. remotum, were clearly distinct from each other and the two groups.L. loliaceum had an isolated position and was most related toL. rigidum. L. remotum had an intermediate position between the cross-breeding and inbreeding species, and was almost equally distant from all three cross-breeding species.Genetic variation inLolium spp. I.     

Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers

We studied mitochondrial divergence in 27 individuals of colubrid snakes of the genus Madagascarophis Mertens from most of its distribution area in Madagascar. Combined analyses of 16S rRNA and cytochrome b sequences identified six major clades which only partly agreed with previously proposed classifications. Analysis of nuclear DNA sequences of the c-mos gene as well as of ISSR fingerprints revealed consistent differences only among three clades which we consider as distinct species: a widespread Madagascarophis colubrinus (Schlegel), with M. citrinus (Boettger) as a junior synonym, a southern M. meridionalis Domergue, and a presumably undescribed species from the extreme north of Madagascar. The species M. ocellatus Domergue was not available for our study. Within M. colubrinus there are two populations from the north-west, each showing two divergent haplotypes with pairwise divergences of up to 5.2% in the cytochrome b gene. Maximum divergence in this gene within M. colubrinus was 7.1%. These high values emphasise that caution needs to be applied before genetic distance values are used for species delimitation. Phylogeographically, most of the genetic variation in M. colubrinus is found in northern Madagascar, indicating that the species might have originated in this region. Later one haplotype clade colonised western and eastern Madagascar, with a putative secondary introgression into north-western populations.     

A reevaluation of the genus Malassezia by means of genome comparison

Results of a study of the genus Malassezia on the basis of genome characters confirm that two species should be maintained, M. furfur and M. pachydermatis. The two forms associated with skin disease, frequently referred to as Pityrosporum orbiculare and P. ovale, were found to be synonymous, the name M. furfur having priority. Malassezia pachydermatis, hitherto regarded as a strictly zoophilic species, may also be found on humans.     

A new species of Mesopropithecus (Primates, Palaeopropithecidae) from Northern Madagascar

We diagnose new subfossil specimens belonging to the extinct palaeopropithecid, Mesopropithecus,from several caves at the Ankarana massif in northern Madagascar. They include three partial crania with mandibles, and the first vertebrae, ribs, clavicle, pelvis, ulna, and hand and foot bones of Mesopropithecusever found. Several other postcranial elements are known for this northern variant of Mesopropithecusand for previously described species from central and southern Madagascar. The new materials cannot be accommodated in either currently recognized species of this genus and are,hereby, given the name Mesopropithecus dolichobrachion. M. dolichobrachionis the second new species of extinct subfossil lemur to be found in the caves of Ankarana. It is unlike the two previously described species of this genus primarily in its larger size, different limb proportions, and aspects of its long bone morphology. Most notable is the fact that this is the only species of Mesopropithecuswith the forelimb longer than the hind limb— hence the name, the “long-armed” Mesopropithecus.Various characteristics place M. dolichobrachionphenetically closest (of species belonging to Mesopropithecus)to Babakotia radofilai, Archaeoindris,and Palaeopropithecus.If these similarities are derived specializations rather than primitive for palaeopropithecids, a new generic name will be required.     

Life history characteristics and behaviour of the bamboo aphid,Pseudoregma bambucicola (Hemiptera: Pemphigidae), having sterile soldiers

Summary Life history and behavioural characteristics of the bamboo aphid,Pseudoregma bambucicola (Takahashi), which has sterile soldiers, were studied in the laboratory. The stadium of normal (fertile) first instar larvae was two times longer than that of second instar larvae, and the stadium of soldier-type (sterile) first instar larvae was much longer (max. 116 days) than the stadium of normal first instar, suggesting that soldiers are able to take nutrition from bamboo. Stimulation of larvae with breath, vibration of bamboo shoots or disturbing the larvae with the tip of a fine brush induced significantly more defensive acts by soldiers than those by normal larvae — the latter usually fled. Soldiers did not attack non-kin conspecific intruders or even aphids of different species, suggesting that, in this species, kin-recognition ability is low.     

Genetic differentiation of Pseudoregma bambucicola population based on mtDNA COII gene

mtDNA COII gene sequences were identified and analyzed using different types of software, namely, MEGA5.0, DNAMAN, and DnaSP5.0 in four Chinese provinces, namely, Sichuan, Zhejiang, Guizhou and Shanghai. Analysis of molecular genetic variation and its genetic structure and differentiation, combined with NJ tree, MP tree analysis and analysis of molecular variance (AMOVA), at Fst = 0.0582 conclude that the genetic differentiation is low, gene flow is Nm = 8.0911, and gene exchange is sufficient. However, for the geographic populations of Pseudoregma bambucicola in the four provinces, their gene exchange is relatively weak at Nm = 0.8284, whereas the genetic differentiation is high at Fst = 0.3764. Based on the data, total nucleotide diversity between the populations is 0.00158 ± 0.00021. The results showed that the total population of Tajima’s D and Fu’s Fs results are D = ?0.885 and Fs = 0.226, respectively. The experimental numerical results showed that this total population is not significant (P > 0.10), indicating that nine different geographic populations are short-term. No expansion occurred in the internal population. This study provided a theoretical and practical basis for the comprehensive prevention and control of P. bambucicola.     

Phylogeography of colour polymorphism in the coral reef fish Pseudochromis fuscus, from Papua New Guinea and the Great Barrier Reef

Body colour has played a significant role in the evolution of coral reef fishes, but the phylogenetic level at which colour variation is expressed and the evolutionary processes driving the development and persistence of different colour patterns are often poorly understood. The aim of this study was to examine the genetic relationships between multiple colour morphs of Pseudochromis fuscus (family Pseudochromidae), both within and among geographic locations. Pseudochromis fuscus is currently described as a single species, but exhibits at least six discrete colour morphs throughout its range. In this study, P. fuscus from Papua New Guinea (PNG) and the Great Barrier Reef (GBR), Australia, formed three genetically distinct clades based on mitochondrial DNA (control region) sequence data: (1) yellow and brown morphs from the GBR and southern PNG, as well as an orange morph from southern PNG; (2) a pink morph from southern PNG; and (3) all three morphs (pink, orange and grey) found in Kimbe Bay, northern PNG. The three groups showed deep levels of divergence (d=14.6–25.4%), suggesting that P. fuscus is a complex of polychromatic species, rather than a single widespread species with many different colour morphs. Population genetic analyses indicate that the three clades have experienced unique evolutionary histories, possibly from differential effects of sea level fluctuations, barriers to gene flow and historical biogeography.