Molecular phylogeny of Asian termites (Isoptera) of the families Termitidae and Rhinotermitidae based on mitochondrial COII sequences

The families Termitidae and Rhinotermitidae are the most evolved and diverse groups of the social insects, termites (Order Isoptera), showing elaborated morphology and complex behavior. Molecular phylogeny of termites with the emphasis on these families was examined by Bayesian and maximum-likelihood analyses based on DNA sequence of mitochondrial cytochrome oxidase II (COII) gene of 31 genera sampled in Asia (mainly Thailand and Japan) along with those reported previously. Termitidae was monophyletic and originated from within polyphyletic Rhinotermitidae. Among the four subfamilies of Termitidae, Macrotermitinae was monophyletic suggesting a single common origin of fungus-growing habit characteristic for this subfamily, and was placed in the basal position in the family. A group consisting of other subfamilies Termitinae and Nasutitermitinae, though some important groups were still untouched, was the most apical but neither Termitinae nor Nasutitermitinae formed a monophyletic lineage. It was implied that, as defense systems of the soldier castes, the appearance of snapping mandibles has occurred at a single event, but the development of nasus for chemical secretion has probably not. Our tree provides some evidence concerning contradictions in the previously proposed phylogeny of termites.     

Evolutionary trend of phylogenetic diversity of nitrogen fixation genes in the gut community of wood-feeding termites

Nitrogen fixation by gut microorganisms is one of the crucial aspects of symbiosis in wood-feeding termites since these termites thrive on a nitrogen-poor diet. In order to understand the evolution of this symbiosis, we analysed the nitrogenase structural gene nifH in the gut microbial communities. In conjunction with the published sequences, we compared approximately 320 putatively functional NifH protein sequences obtained from a total of 19 termite samples that represent all the major branches of their currently proposed phylogeny, and from one species of the cockroach Cryptocercus that shares a common ancestor with termites. Using multivariate techniques for clustering and ordination, a phylogeny of NifH protein sequences was created and plotted variously with host termite families, genera, and species. Close concordance was observed between NifH communities and the host termites at genus level, but family level relationships were not always congruent with accepted termite clade structure. Host groups examined included basal families (Mastotermitidae, Termopsidae, Kalotermitidae, as well as Cryptocercus), the most derived lower termite family Rhinotermitidae, and subfamilies representing the advanced and highly diverse apical family Termitidae (Macrotermitinae, Termitinae, and Nasutitermitinae). This selection encompassed the major nesting and feeding styles recognized in termites, and it was evident that NifH phylogenetic divergence, as well as the occurrence of alternative nitrogenase-type NifH, was to some extent dependent on host lifestyle as well as phylogenetic position.     

A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology

The first comprehensive combined molecular and morphological phylogenetic analysis of the major groups of termites is presented. This was based on the analysis of three genes (cytochrome oxidase II, 12S and 28S) and worker characters for approximately 250 species of termites. Parsimony analysis of the aligned dataset showed that the monophyly of Hodotermitidae, Kalotermitidae and Termitidae were well supported, while Termopsidae and Rhinotermitidae were both paraphyletic on the estimated cladogram. Within Termitidae, the most diverse and ecologically most important family, the monophyly of Macrotermitinae, Foraminitermitinae, Apicotermitinae, Syntermitinae and Nasutitermitinae were all broadly supported, but Termitinae was paraphyletic. The pantropical genera Termes, Amitermes and Nasutitermes were all paraphyletic on the estimated cladogram, with at least 17 genera nested within Nasutitermes, given the presently accepted generic limits. Key biological features were mapped onto the cladogram. It was not possible to reconstruct the evolution of true workers unambiguously, as it was as parsimonious to assume a basal evolution of true workers and subsequent evolution of pseudergates, as to assume a basal condition of pseudergates and subsequent evolution of true workers. However, true workers were only found in species with either separate- or intermediate-type nests, so that the mapping of nest habit and worker type onto the cladogram were perfectly correlated. Feeding group evolution, however, showed a much more complex pattern, particularly within the Termitidae, where it proved impossible to estimate unambiguously the ancestral state within the family (which is associated with the loss of worker gut flagellates). However, one biologically plausible optimization implies an initial evolution from wood-feeding to fungus-growing, proposed as the ancestral condition within the Termitidae, followed by the very early evolution of soil-feeding and subsequent re-evolution of wood-feeding in numerous lineages.     

Termite‐induced heterogeneity in African savanna vegetation: mechanisms and patterns

Objectives: To (1) assess the strength of evidence for the role of termites in vegetation heterogeneity in African savannas, and (2) identify the mechanisms by which termites induce such heterogeneity. Location: African savannas. Methods: We conducted a review of the literature, a meta‐analysis and qualitative systems analysis to identify mechanisms to explain the observed patterns. Results: The review provided evidence for termite‐induced heterogeneity in floristic composition and vegetation patterning in savannas across Africa. Termites induced vegetation heterogeneity directly or indirectly through their nest‐building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire. The literature reviewed indicated that termite mounds essentially act as islands of fertility, which are responsible for ecosystem‐level spatial heterogeneity in savannas. This was supported by the meta‐analysis, which demonstrated that mounds of Ancistrotermes, Macrotermes, Odontotermes (family Macrotermitinae), Cubitermes (family Termitinae) and Trinervitermes (Nasutitermitinae) are significantly enriched in clay (75%), carbon (16%), total nitrogen (42%), calcium (232%), potassium (306%) and magnesium (154%) compared to the surrounding savanna soil. Conclusions: Termite activity is one of the major factors that induce vegetation patterning in African savannas. The implications of this are discussed and research questions for future studies and modelling efforts are indicated.     

Diet composition of an escaped captive‐born southern tamandua (Tamandua tetradactyla) in a nonnative habitat in Asia

Studies on the role of natural predatory instincts in captive‐born mammalian myrmecophagy are rare. Consequently, researchers rely extensively on case reports to learn more about the contexts in which predatory behavior occurs among such animals. In this study, we recorded an uncommon case of a captive‐born southern tamandua (Tamandua tetradactyla) that accidentally escaped from a zoo into a nonnative habitat in Asia. The southern tamandua was found alive 3 months later. Two fresh fecal samples were obtained, and the diet composition was examined. Three termite species (one family, three genera), and 14 ant species (four subfamilies, nine genera) were identified in the fecal samples. The studied southern tamandua preyed on terrestrial and arboreal ants and termites, as the wild populations of its species do. Ants of the subfamily Myrmicinae and termites of the subfamily Nasutitermitinae were the most abundant prey items in the samples, which is consistent with related reports on the wild populations. Soldier ants constituted <1% of the prey items in the fecal samples, suggesting that the southern tamandua likely avoided preying on ants of the soldier caste. Fungus‐growing termites Odontotermes (Isoptera: Macrotermitinae), which are not native to neotropical regions, were also ingested by the southern tamandua. This study provides information on how a captive‐born mammalian myrmecophagy applies its natural feeding instincts in nonnative natural settings.     

Comparative morphology of epicortical roots in Old and New World Loranthaceae with reference to root types, origin, patterns of longitudinal extension and potential for clonal growth

The large mistletoe family, Loranthaceae, contains 75 genera, three of which are terrestrial root parasites. The remaining 72 genera are aerial parasites. Four basic haustorial system types are found in aerial genera: epicortical roots (ERs), wood roses, clasping unions and bark strands. The focus of this report is on genera in which ERs are present. Presence of ERs is based on our worldwide collection of haustoria and from literature sources. Our collections include 78% of all aerial genera and 72% of genera with ERs. Collections were analyzed using comparative morphological methods. Of the 72 aerial genera 40 (56%) have ERs and 75% of these are Old World. ERs are the most common haustorial type for Loranthaceae on every major landmass except Africa. Three ER types are described, basal, cauline and adventitious. Basal and adventitious ERs occur in both the Old and New World, whereas cauline ERs are exclusively New World. Adventitious ERs form in a few species in response to injury or epiparasitism. Significant differences occur between basal and cauline ERs in the extent and pattern of elongation, frequency of lateral root formation, and production of haustoria and shoots. Three patterns of axis extension of ERs are recognized, the monochasial sympodium, dichasial sympodium and monopodium. Marked differences in patterns of axis elongation occur between the Old and New World genera analyzed. In Old World taxa 94% of lateral roots contributed to a monochasial sympodium, whereas in New World taxa 84% of root extension was monopodial. Two strategies of resource procurement occur in genera with ERs; the “phalanx” strategy is found in species with basal ERs only, the “guerilla” strategy in New World species with cauline ERs. Species with ERs have the potential for clonal growth through fragmentation of stems, ERs, or both, but the extent of clonal growth in nature is unknown. The large number and wide distribution of genera with ERs add support to the hypothesis that the presence of ERs is an ancestral trait for aerial Loranthaceae.     

On the elevated intestinal pH of higher termites (Isoptera: Termitidae)

Summary The pH of the gut contents was measured in 52 species of higher termites (Termitidae), representing 36 genera in all four subfamilies. A statistically significant trend was shown from lower termites with low mean gut pH through to the Termitinae with higher mean gut pHs. Elevation of the pH occurred principally in the first and third proctodaeal segments, reaching values as high as 10.5 in 8 soil-feeding genera and 1 wood-feeding genus of Termitinae. Elevation of gut pH within the Termitidae appears to be independent of the general nature of the feeding substrate.     

A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology

? Premise of the study: The Malpighiaceae include ～1300 tropical flowering plant species in which generic definitions and intergeneric relationships have long been problematic. The goals of our study were to resolve relationships among the 11 generic segregates from the New World genus Mascagnia, test the monophyly of the largest remaining Malpighiaceae genera, and clarify the placement of Old World Malpighiaceae. ? Methods: We combined DNA sequence data for four genes (plastid ndhF, matK, and rbcL and nuclear PHYC) from 338 ingroup accessions that represented all 77 currently recognized genera with morphological data from 144 ingroup species to produce a complete generic phylogeny of the family. ? Key results and conclusions: The genera are distributed among 14 mostly well-supported clades. The interrelationships of these major subclades have strong support, except for the clade comprising the wing-fruited genera (i.e., the malpighioid+Amorimia, Ectopopterys, hiraeoid, stigmaphylloid, and tetrapteroid clades). These results resolve numerous systematic problems, while others have emerged and constitute opportunities for future study. Malpighiaceae migrated from the New to Old World nine times, with two of those migrants being very recent arrivals from the New World. The seven other Old World clades dispersed much earlier, likely during the Tertiary. Comparison of floral morphology in Old World Malpighiaceae with their closest New World relatives suggests that morphological stasis in the New World likely results from selection by neotropical oil-bee pollinators and that the morphological diversity found in Old World flowers has evolved following their release from selection by those bees.     

Fungus-growing termites originated in African rain forest

Fungus-growing termites (subfamily Macrotermitinae, Isoptera) cultivate fungal crops (genus Termitomyces, Basidiomycotina) in gardens inside their colonies. Those fungus gardens are continuously provided with plant substrates, whereas older parts that have been well decomposed by the fungus are consumed (cf.). Fungus-growing termites are found throughout the Old World tropics, in rain forests and savannas, but are ecologically dominant in savannas. Here, we reconstruct the ancestral habitat and geographical origin of fungus-growing termites. We used a statistical model of habitat switching repeated over all phylogenetic trees sampled in a Bayesian analysis of molecular data. Our reconstructions provide strong evidence that termite agriculture originated in African rain forest and that the main radiation leading to the extant genera occurred there. Because extant savanna species are found in most genera, this moreover suggests that the savanna has repeatedly been colonized by fungus-growing termites. Furthermore, at least four independent "out-of-Africa" migrations into Asia, and at least one independent migration to Madagascar, have occurred. Although fungus growing by termites is ecologically most successful under the variable, unfavorable conditions of the savanna, it seems to have evolved under the more constant and favorable conditions of the rain forest.     

A Phylogeny‐Based Comparison of Tarantula Spider Anti‐Predator Behavior Reveals Correlation of Morphology and Behavior

We studied anti‐predator strategies in nine species of tarantulas from Asia, Africa, and the Americas. Tarantulas in the New World possess urticating hairs which induce inflammation when in contact with vertebrate mucus membranes and skin. In contrast, tarantulas from the Old World lack this defense but are observed to exhibit a much greater willingness to escalate to an active defense when provoked. We had three goals: (1) describe the behaviors exhibited by each taxon in response to two levels of provocation, (2) look for the presence of alternative classes of anti‐predator strategy as predicted by the Old World–New World dichotomy in aggressive defense, and (3) examine the evolution of these behaviors in the context of the phylogeny of the group. We compared the response of nine different mygalomorph spider genera to two levels of aversive stimuli: puffs of air and prodding. We found that the overall structure of the defensive behavior was similar between the different taxa, consisting of fleeing, rearing, striking, and biting. Some genera did exhibit unique behaviors such as stridulating (Hysterocrates) or rocking (Haplopelma and Selenocosmia). We found that the genera from the New World exhibited low levels of escalation in their defense behaviors, while those from the Old World readily escalated to striking and biting. Our findings are consistent with the hypothesis that the possession of urticating hairs is associated with very low levels of active defense behaviors such as striking and biting. The phylogenetic analysis indicates that both the notable levels of aggression displayed by the African taxa tested and the relative passivity of the New World tarantulas each represents a synapomorphy.     

Speices diversity and the systematics of some New World leaf litter Thysanoptera (Phlaeothripinae; Glyptothripini)

Abstract More than fifty species of closely related glyptothripine Thysanoptera
have been described from leaf litter at one site in southern Brazil. The ecological and evolutionary significance of this species swarm are discussed, and the relationships of the New World glyptothripine genera are reassessed. A key is produced to the world genera of the Glyptothripini, and a check list is provided to the fifteen genera and 119 species from the New World. Nomenclatural changes include thirty-five new combinations, five new generic synonyms and one new specific synonym.     

Inferring the phylogenetic position of Boa constrictor among the Boinae

Snakes of the subfamily Boinae are found in Madagascar, the Papuan-Pacific Islands, and the Neotropics. It has been suggested that genera within each of these particular areas do not form monophyletic groups. Further, it was proposed that the New World Boa constrictor is more closely related to boine genera in Madagascar than to boines in the Neotropics. Along with inferring the relationship of all boine genera using data from the cytochrome b gene and morphology, the placement of Boa was also examined. Phylogenetic inferences using maximum likelihood and Bayesian (BI) methods for combined data analyses and separate analyses of DNA sequence and morphological data were conducted. Priors, parametric bootstraps, and the Shimodaira-Hasegawa test were used to examine the previously proposed placement of Boa with Madagascan taxa using these DNA data. DNA data and combined data analyses strongly reject the hypothesis that Boa is more closely related to Old World genera than to other New World genera. Additionally, strong tree support suggests that all species within Madagascar, the Papuan-Pacific Islands, and the Neotropics each form a monophyletic group with respect to their geographic region.     

Retropositional events consolidate the branching order among New World monkey genera

Due to contradicting relationships obtained from various morphological and genetic studies, phylogenetic relationships among New World monkey genera are highly disputed. In the present study, we analyzed the presence/absence pattern of 128 SINE integrations in all New World monkey genera. Among them, 70 were specific for only a single genus, whereas another 18 were present in all New World monkey genera. The 40 remaining insertions were informative to elucidate phylogenetic relationships among genera. Several of them confirmed the monophyly of the three families Cebidae, Atelidae and Pitheciidae as well as of the subfamily Callithrichinae. Further markers provided evidence for a sister grouping of Cebidae and Atelidae to the exclusion of Pitheciidae as well as for relationships among genera belonging to Callithrichinae and Atelidae. Although a close affiliation of Saimiri, Aotus and Cebus to Callithrichinae was shown, the relationships among the three genera remained unresolved due to three contradicting insertions.     

On the loss of uricolytic activity during primate evolution--I. Silencing of urate oxidase in a hominoid ancestor

Urate oxidase activity is not detectable in liver homogenates from the gibbon, orangutan, chimpanzee, gorilla and human. Liver homogenates from five genera of Old World and two genera of New World monkeys have easily detectable levels of urate oxidase activity. There is no evidence for extant detectable intermediate steps in the loss of urate oxidase activity in the hominoids. Urate oxidase activity from Old World and New World monkeys is stable, a simple observation which debunks a long-standing myth. Urate oxidase activity was silenced in an ancestor to the five living genera of hominoids after divergence from the Old World monkeys.     

Phylogenetic relationships in Termitomyces (Family Agaricaceae) based on the nucleotide sequence of ITS: a first approach to elucidate the evolutionary history of the symbiosis between fungus-growing termites and their fungi

Termitomyces constitutes a very poorly known genus of fungi whose essential characteristic is that all representatives of the genus are cultivated by termites (Macrotermitinae) in their nest and that all the fungi cultivated by termites belong to this genus. For the first time, the phylogenetic relationships of several African Termitomyces species was studied by the sequencing of their internal transcriber spacer region (ITS1--5.8S--ITS2). It appeared that this group is clearly monophyletic and belongs to the Tricholomataceae family. The total homology of the ITS zone of several Termitomyces symbionts of different termite genera indicated that the specific diversity of this group is in fact less important than previously supposed. Finally, the comparison between the Termitomyces phylogenetic tree and the taxonomic tree of Macrotermitinae showed that if for certain genera the hypothesis of termite/fungus coevolution is acceptable, it should not be applied for all symbiosis.     

Phylogenetic analysis of the New World Ptininae (Coleoptera: Bostrichoidea)

A phylogenetic analysis of the New World Ptininae (Anobiidae) was conducted with representatives of nine of ten New World genera, several Old World genera and seven more of the ten subfamilies of Anobiidae. One hundred and two characters (forty‐three multistate) from thirty‐four taxa were used. The single cladogram shows Ptininae as monophyletic and the sister group of the remaining Anobiidae, supporting their placement as subfamilies of a monophyletic Bostrichidae. Genus Niptus Boieldieu is polyphyletic supporting recognition of Pseudeurostus Heyden and the creation of a new genus to encompass the remaining New World species of Niptus. Flightlessness has evolved a minimum of three times within Ptininae and myrmecophily has probably evolved three times within just the New World taxa. The classifications of Ptininae and the remaining Anobiidae are examined and the evolution of feeding habits, myrmecophily and wing loss are discussed.     

Molecular phylogeny of a cosmopolitan group of woodpeckers (genus Picoides) gased on COI and cyt b mitochondrial gene sequences.

Picoides is the largest genus of woodpeckers and member species are found on most major land masses. Current systematic arrangement of this group, based on morphological, behavioral, and plumage characters, suggests that New World species evolved from a single invasion by a Eurasian common ancestor and that all New World species form a monophyletic group. No clear link has ever been established between the relationships of Old World and New World species other than to infer that the most primitive species is Eurasian. This study employs DNA sequences for two protein-coding mitochondrial genes, cytochrome oxidase I and cytochrome b, to reconstruct phylogenetic relationships among all New World species and several Eurasian representatives of the genus Picoides. A well-resolved mitochondrial gene tree is in direct conflict with proposed species relationships based on nongenetic characters; monophyly among New World species is rejected, the evolution of New World species likely resulted from as many as three independent Eurasian invasions, and Picoides is paraphyletic with two other woodpecker genera, Veniliornis and Dendropicos. These results strongly suggest that this large, cosmopolitan genus is in need of systematic revision in order to reflect evolutionary history.     

Polymorphism and oligogyny in the ponerine antCentromyrmex bequaerti (Formicidae: Ponerinae)

Summary Centromyrmex bequaerti is a ponerine ant of the African equatorial forest which nests in the termitaries of diverse Termitinae and Macrotermitinae, occupying several chambers and galleries of the termitaries and preying on their guests. This species presents four traits convergent with termitolestic Myrmicinae: a parasitic nesting habit (but shelters directly in the cavities of the termitaries and not in the walls), blind workers, a strong dimorphism between the queens (20 mm in length) and the workers (from 5.75 to 12 mm total length) and polymorphism in the worker caste. We recorded two other traits considered as evolutionarily-derived: oligogyny, a kind of polygyny (the societies include up to 13 queens, but each queen lives in a separate cavity of the termitary) and large colon size. Behavioural observations show that there is a relationship between worker size and polyethism in this species. Although workers of all size classes participate in hunting, the media are more often engaged in this activity and the largest workers guard the nest entrance.