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Molecular phylogeny and historical biogeography of Apicotermitinae (Blattodea: Termitidae)
Abstract
Soil-feeding termites are abundant in tropical regions and play an important role in soil bioturbation and in the organic matter cycle. The Apicotermitinae are arguably the most diverse lineage of soil-feeding termites, but they are also the most understudied, probably because many species are soldierless, which makes identification difficult. Although the backbone of the termite phylogenetic tree is now well-resolved, the relationships among representatives of Apicotermitinae are still largely unknown. Here, we present phylogenetic trees inferred from 113 mitochondrial genomes of Apicotermitinae representative of the group diversity. Our analyses confirm the monophyly of the Apicotermitinae and the basal position of soldiered taxa, within which two lineages of soldierless species are nested. We describe two new monotypic genera, whose phylogenetic position appeared of special interest: Koutabatermes gen. n., lies on a long branch among soldiered taxa, and Apolemotermes gen. n., is sister to Adaiphrotermes. We resolved, with high support, the position of Asian genera as sister group of a clade comprising the monophyletic neotropical Anoplotermes-group and the small African clade including Adaiphrotermes and Apolemotermes gen. n.. Our trees cast light on the intergeneric and interspecific relationships within Apicotermitinae and reveal the polyphyly of several genera, including Ruptitermes, Astalotermes and Anoplotermes. Biogeographic reconstructions revealed two dispersal events out of Africa, one to the Oriental realm and one to the Neotropical realm. Overall, the timing of Apicotermitinae diversification and dispersal, following the Eocene–Oligocene boundary, matches that found for other groups of Neoisoptera. Nomenclatural acts are registered in ZooBank:
http://zoobank.org/urn:lsid:zoobank.org:pub:CA1A21B6-573E-4855-8C88-372453C922F7.
... The soldierless termites of the New World form a monophyletic clade (Romero Arias et al. 2021) that comprises 16-47% of the termite diversity in Amazonian ecosystems (Bourguignon et al. 2016b). Although the richness of soldierless taxa is recognized, most have not yet been described (Bourguignon et al. 2016b). ...
... Advances in the taxonomy, phylogeny, biogeography, ecology of the Apicotermitinae are ongoing (Bourguignon et al. 2016b, Romero Arias et al. 2021. One area of study that remains poorly understood is the physiology, especially, digestive physiology and its relationship with the gut morphology of these mostly soil-feeding insects. ...
Chasitermes pax Scheffrahn & Carrijo gen. et sp. nov. is described from workers collected from a single colony in the Northern Range of Trinidad. The shape and texture of the unsclerotized enteric valve, tubular shape of the enteric valve seating, and prominent spherical mesenteric tongue of C. pax are the diagnostic characters for both the genus and species. A Bayesian phylogenetic analysis using the COI gene and including all neotropical Apicotermitinae genera described to date supports the new genus as a distinct terminal.
... (= Hodotermitinae sensu Emerson, 1942) by Grassé (1949) Although the historical biogeography of Neoisoptera and Kalotermitidae has been studied in detail (Bourguignon et al., 2016(Bourguignon et al., , 2017Bucek et al., 2021;Romero Arias et al., 2021;Wang et al., 2019), only a few species of Stolotermitidae, Hodotermitidae and Archotermopsidae have been included in previous termite phylogenies. In this study, we carried out a representative sampling of species belonging to these three families. ...
... In the remainder of this section, we shall refer to the fami- We did not attempt to reconstruct the ancestral range of Stolotermitidae + Hodotermopsidae + Archotermopsidae + Hodotermitidae, particularly given that the many fossils occurring well outside of modern distributions would render meaningless such an estimate based solely on extant taxa. Ancestral range reconstructions have been performed previously for Neoisoptera and Kalotermitidae (Bourguignon et al., 2016(Bourguignon et al., , 2017Bucek et al., 2021;Romero Arias et al., 2021;Wang et al., 2019). However, compared to Teletisoptera, Neoisoptera and Kalotermitidae are diverse and widespread, comprising many extant species whose distribution and phylogenetic relationships can inform on past vicariance and dispersal events, and with most fossils nested within those distributions (Krishna et al., 2013). ...
Termites are social cockroaches distributed throughout warm temperate and tropical ecosystems. The ancestor of modern termites roamed the earth during the early Cretaceous, suggesting that both vicariance and overseas dispersal may have shaped the distribution of early diverging termites. We investigate the historical biogeography of three early diverging termite families –Stolotermitidae, Hodotermitidae and Archotermopsidae (clade Teletisoptera) – using the nuclear rRNA genes and mitochondrial genomes of 27 samples. Our analyses confirm the monophyly of Teletisoptera, with Stolotermitidae diverging from Hodotermitidae + Archotermopsidae approximately 100 Ma. Although Hodotermitidae are monophyletic, our results demonstrate the paraphyly of Archotermopsidae. Phylogenetic analyses indicate that the timing of divergence among the main lineages of Hodotermitidae + Archotermopsidae are compatible with vicariance. In the Stolotermitidae, however, the common ancestors of modern Porotermes Hagen and Stolotermes Hagen are roughly as old as 20 and 35 Ma, respectively, indicating that the presence of these genera in South America, Africa and Australia involved over‐water dispersals. Overall, our results suggest that early diverging termite lineages acquired their current distribution through a combination of over‐water dispersals and dispersal via land bridges. We clarify the classification by resolving the paraphyly of Archotermopsidae, restricting the family to Archotermopsis Desneux and Zootermopsis Emerson and elevating Hodotermopsinae ( Hodotermopsis Holmgren) as Hodotermopsidae ( status novum ).
... While the historical biogeography of Neoisoptera and Kalotermitidae has been studied in detail (Bourguignon et al., 2016(Bourguignon et al., , 2017Wang et al., 2019;Romero Arias et al., 2021;Bucek et al., 2021), only a few species of Stolotermitidae, Hodotermitidae, and Archotermopsidae have been included in previous termite phylogenies. In this paper, we carried out a representative sampling of species belonging to these three families. ...
... We did not attempt to reconstruct the ancestral range of Stolotermitidae + Hodotermopsidae + Archotermopsidae + Hodotermitidae, particularly given that the many fossils occurring well outside of modern distributions would render meaningless such an estimate based solely on extant taxa. Ancestral range reconstructions have been performed previously for Neoisoptera and Kalotermitidae (Bourguignon et al., 2016(Bourguignon et al., , 2017Wang et al., 2019;Romero Arias et al., 2021;Bucek et al., 2021). However, compared to Stolotermitidae + Hodotermopsidae + Archotermopsidae + Hodotermitidae, Neoisoptera and Kalotermitidae are diverse and widespread, comprising many extant species whose distribution and phylogenetic relationships can inform on past vicariance and dispersal events, and with most fossils nested within those distributions (Krishna et al., 2013). ...
Termites are social cockroaches distributed throughout warm temperate and tropical ecosystems. The ancestor of modern termites (crown-Isoptera) occurred during the earliest Cretaceous, approximately 140 million years ago, suggesting that both vicariance through continental drift and overseas dispersal may have shaped the distribution of early diverging termite lineages. We reconstruct the historical biogeography of three early diverging termite families (Stolotermitidae, Hodotermitidae, and Archotermopsidae) using the nuclear rRNA genes and mitochondrial genomes of 27 samples. Our analyses confirmed the monophyly of Stolotermitidae + Hodotermitidae + Archotermopsidae (clade Teletisoptera), with Stolotermitidae diverging from a monophyletic Hodotermitidae + Archotermopsidae approximately 100.3 Ma (94.3-110.4 Ma, 95% HPD), and with Archotermopsidae paraphyletic to a monophyletic Hodotermitidae. The Oriental Archotermopsis and the Nearctic Zootermopsis diverged 50.8 Ma (40.7-61.4 Ma, 95% HPD) before land connections between the Palearctic region and North America ceased to exist. The African Hodotermes + Microhodotermes diverged from Anacanthotermes, a genus found in Africa and Asia, 32.1 Ma (24.8-39.9 Ma, 95% HPD), and the most recent common ancestor of Anacanthotermes lived 10.7 Ma (7.3-14.3 Ma, 95% HPD), suggesting that Anacanthotermes dispersed to Asia using the land bridge connecting Africa and Eurasia ~18-20 Ma. In contrast, the common ancestors of modern Porotermes and Stolotermes lived 20.2 Ma (15.7-25.1 Ma, 95% HPD) and 26.6 Ma (18.3-35.6 Ma, 95% HPD), respectively, indicating that the presence of these genera in South America, Africa, and Australia involved over-water dispersals. Our results suggest that early diverging termite lineages acquired their current distribution through a combination of over-water dispersals and dispersal via land bridges. We clarify the classification by resolving the paraphyly of Archotermopsidae, restricting the family to Archotermopsis and Zootermopsis, and elevating Hodotermopsinae (Hodotermopsis) as Hodotermopsidae (status novum).
... Since most samples from museums and collections are yet to be studied and there is a lack of inventories in many areas, the geographic gap for these termites is even greater than for other groups. This is also true in the other parts of the world, where the taxonomic gap is not so deep because of the older available literature (Grassé and Noirot 1954;Sands 1972), but the geographical gap still exists due the lack of researchers for a long period (Romero Arias et al. 2021). ...
The neotropical Apicotermitinae is a common and widespread clade of mostly soil-feeding soldierless termites. With few exceptions, species of this group were originally assigned to the genus Anoplotermes Müller, 1873. The application of internal worker morphology coupled with genetic sequencing has recently shed light on the true diversity of this subfamily. Herein, Anoplotermes susanae Scheffrahn, Carrijo & Castro, sp. nov. and four new species in four new genera are described: Hirsutitermes kanzakii Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Krecekitermes daironi Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Mangolditermes curveileum Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., and Ourissotermes giblinorum Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov. Worker descriptions are based mainly on worker gut morphology, including the enteric valve, while imagoes were described based on external characters. A Bayesian phylogenetic tree of New World Apicotermitinae was constructed using the complete mitogenome to infer genera relationships and corroborate the taxonomic decisions. Distribution maps and a dichotomic key to the known Neotropical Apicotermitinae genera are provided.
... The pathways and timing of the spread of termites across continents have been studied in detail in Neoisoptera (Bourguignon et al. 2016, Wang et al. 2019, Romero Arias et al. 2021, with the exception of Madagascar, which has been largely overlooked. The Neoisoptera contain more than 80% of described termite species classified into four families: the Stylotermitidae, the Serritermitidae, the Termitidae and the paraphyletic Rhinotermitidae (Engel et al. 2009, Krishna et al. 2013. ...
Madagascar is home to many endemic plant and animal species owing to its ancient isolation from other landmasses. This unique fauna includes several lineages of termites, a group of insects known for their key role in organic matter decomposition in many terrestrial ecosystems. How and when termites colonised Madagascar remains unknown. In this study, we used 601 mitochondrial genomes, 93 of which were generated from Malagasy samples, to infer the global historical biogeography of Neoisoptera, a lineage containing more than 80% of described termite species. Our results indicate that Neoisoptera colonised Madagascar between 7 and 10 times independently during the Miocene, between 8.4 and 16.6 Ma (95% HPD: 6.1–19.9 Ma). This timing matches that of the colonization of Australia by Neoisoptera. Furthermore, the taxonomic composition of the Neoisopteran fauna of Madagascar and Australia are strikingly similar, with Madagascar harbouring an additional two lineages absent from Australia. Therefore, akin to Australia, Neoisoptera colonised Madagascar during the global expansion of grasslands, possibly helped by the ecological opportunities arising from the spread of this new biome.
... Furthermore, Romero Arias et al. 2021 have identified six additional monotypic apicotermitine genera from the Ethiopian Region using mitochondrial genome sequencing. Herein the worker denoted as 'Apicotermitinae genus L' (Romero Arias et al. 2021) is described as Ebogotermes raphaeli gen. n. sp. ...
Ebogotermes raphaeli gen. n. sp. n., is described from workers collected in Cameroon. This soil-feeding termite is the largest soldierless termite from central Africa and aligns with the Anoplotermes subgroup. The enteric valve armature is weakly armed and, as with most apicotermitine species, is uniquely diagnostic.
While new species of termites are described every year, the description of species distant from every known termite species is rare. In this paper, we describe one such species, Engelitermes zambo sp.n. , an African Termitidae belonging to an entirely new lineage of termites for which we create a new subfamily, Engelitermitinae subfam.n. The subfamily status of Engelitermitinae was supported by termite phylogenetic trees, including sequences from the four existing samples of E. zambo sp.n. , which, albeit with low bootstrap supports, placed Engelitermes gen.n. on a long branch sister to Forficulitermes , the two of which formed the sister group of a clade comprising Cubitermitinae, Nasutitermitinae, Syntermitinae and all other Termitinae. The sister relationship between Engelitermes gen.n. and Forficulitermes is further supported by the similar gut structure of their workers. In contrast, the soldiers of Engelitermes gen.n. resemble those of Cephalotermes . Our phylogenetic analyses, including all clades of Termitinae, call for a global taxonomic revision of the Termitinae subfamily names. Finally, our study highlights that new unique termite lineages are still awaiting to be described.
Termites feed on vegetal matter at various stages of decomposition. Lineages of wood- and soil-feeding termites are distributed across terrestrial ecosystems located between 45°N and 45°S of latitude, a distribution they acquired through many transoceanic dispersal events. While wood-feeding termites often live in the wood on which they feed and are efficient at dispersing across oceans by rafting, soil-feeders are believed to be poor dispersers. Therefore, their distribution across multiple continents requires an explanation. Here, we reconstructed the historical biogeography and the ancestral diet of termites using mitochondrial genomes and δ13C and δ15N stable isotope measurements obtained from 324 termite samples collected in five biogeographic realms. Our biogeographic models showed that wood-feeders are better at dispersing across oceans than soil-feeders, further corroborated by the presence of wood-feeders on remote islands devoid of soil-feeders. However, our ancestral range reconstructions identified 33 dispersal events among biogeographic realms, 18 of which were performed by soil-feeders. Therefore, despite their lower dispersal ability, soil-feeders performed several transoceanic dispersals that shaped the distribution of modern termites.
Machadotermes is one of the basal Apicotermitinae genera, living in tropical West Africa. Old observations suggested the presence of a new gland, the intramandibular gland, in Machadotermes soldiers. Here, by combining micro-computed tomography, optical and electron microscopy, we showed that the gland exists in Machadotermes soldiers only as an active exocrine organ, consisting of numerous class III cells (bicellular units made of secretory and canal cells), within which the secretion is produced in rough endoplasmic reticulum, and modified and stored in Golgi apparatus. The final secretion is released out from the body through epicuticular canals running through the mandible cuticle to the exterior. We also studied three other Apicotermitinae, Indotermes, Duplidentitermes, and Jugositermes, in which this gland is absent. We speculate that the secretion of this gland may be used as a general protectant or antimicrobial agent. In addition, we observed that the frontal gland, a specific defensive organ in termites, is absent in Machadotermes soldiers while it is tiny in Indotermes soldiers and small in Duplidentitermes and Jugositermes soldiers. At last, we could also observe in all these species the labral, mandibular and labial glands, other exocrine glands present in all termite species studied so far.
GetOrganelle is a state-of-the-art toolkit to accurately assemble organelle genomes from whole genome sequencing data. It recruits organelle-associated reads using a modified “baiting and iterative mapping” approach, conducts de novo assembly, filters and disentangles the assembly graph, and produces all possible configurations of circular organelle genomes. For 50 published plant datasets, we are able to reassemble the circular plastomes from 47 datasets using GetOrganelle. GetOrganelle assemblies are more accurate than published and/or NOVOPlasty-reassembled plastomes as assessed by mapping. We also assemble complete mitochondrial genomes using GetOrganelle. GetOrganelle is freely released under a GPL-3 license (https://github.com/Kinggerm/GetOrganelle).
Termites are one of the key ecosystem engineers in tropical forests where they play a major role in decomposition rates, both above and belowground. The interest in termite ecology and biogeography has increased in the last few decades; however, the lack of comparable data has limited the wider impact of termite research. For Ecuador, termite studies are relatively rare and comparable data that are collected using standardized sampling methods are missing. In this study, we aim to 1) provide comparable data of termite species and feeding-group diversity from two primary forests in Ecuador and 2) explore the differences in termite species and feeding-group diversity between the two forest sites. Sampling took place in the national parks of Yasuní and Podocarpus where three belt transects (100 × 2 m) following Jones and Eggleton (2000) were conducted in each forest. We found that termite species richness was higher in Yasuní (56 species) than in Podocarpus (24 species) and that 57% of the sampled termite genera had never previously been recorded in Ecuador. The inter-site species dissimilarity was almost complete (Bray Curtis (±SD), 0.91 ± 0.01), which may have been linked to the difference in tree density and species richness in the two forests. Termite feeding-groups diversity was significantly higher in Yasuní than in Podocarpus with the exception of soil-feeding termites which may have been due to competition between humus-and soil-feeding species.
With the continual progress of sequencing techniques, genome-scale data are increasingly used in phylogenetic studies. With more data from throughout the genome, the relationship between genes and different kinds of characters is receiving more attention. Here, we present version 4 of RASP, a software to reconstruct ancestral states through phylogenetic trees. RASP can apply generalized statistical ancestral reconstruction methods to phylogenies, explore the phylogenetic signal of characters to particular trees, calculate distances between trees, and cluster trees into groups. RASP 4 has an improved graphic user interface and is freely available from http://mnh.scu.edu.cn/soft/blog/RASP (program) and https://github.com/sculab/RASP (source code).
With the rapid increase of sequenced metazoan mitochondrial genomes, a detailed manual annotation is becoming more and more infeasible. While it is easy to identify the approximate location of protein-coding genes within mitogenomes, the peculiar processing of mitochondrial transcripts, however, makes the determination of precise gene boundaries a surprisingly difficult problem. We have analyzed the properties of annotated start and stop codon positions in detail, and use the inferred patterns to devise a new method for predicting gene boundaries in de novo annotations. Our method benefits from empirically observed prevalances of start/stop codons and gene lengths, and considers the dependence of these features on variations of genetic codes. Albeit not being perfect, our new approach yields a drastic improvement in the accuracy of gene boundaries and upgrades the mitochondrial genome annotation server MITOS to an even more sophisticated tool for fully automatic annotation of metazoan mitochondrial genomes.
Anenteotermescherubimi Scheffrahn, sp. n. is described from workers and male imagos collected in Cameroon and Republic of the Congo. This is the smallest soldierless termite known from Africa. As with many soldierless and soil-feeding termite species, the enteric valve morphology is a robust and essential diagnostic character for An.cherubimi . Preserved workers display pre-autothysis morphology and the effects of abdominal autothysis.
The higher termites (Termitidae) are keystone species and ecosystem engineers. They have exceptional biomass and play important roles in decomposition of dead plant matter, in soil manipulation, and as the primary food for many animals, especially in the tropics. Higher termites are most diverse in rainforests, with estimated origins in the late Eocene (∼54 Ma), postdating the breakup of Pangaea and Gondwana when most continents became separated. Since termites are poor fliers, their origin and spread across the globe requires alternative explanation. Here, we show that higher termites originated 42-54 Ma in Africa and subsequently underwent at least 24 dispersal events between the continents in two main periods. Using phylogenetic analyses of mitochondrial genomes from 415 species, including all higher termite taxonomic and feeding groups, we inferred 10 dispersal events to South America and Asia 35-23 Ma, coinciding with the sharp decrease in global temperature, sea level, and rainforest cover in the Oligocene. After global temperatures increased, 23-5 Ma, there was only one more dispersal to South America but 11 to Asia and Australia, and one dispersal back to Africa. Most of these dispersal events were transoceanic and might have occurred via floating logs. The spread of higher termites across oceans was helped by the novel ecological opportunities brought about by environmental and ecosystem change, and led termites to become one of the few insect groups with specialized mammal predators. This has parallels with modern invasive species that have been able to thrive in human-impacted ecosystems.
This paper gives a historical overview of the African rain forest from its origins, towards, the end of the Cretaceous period. The areas around the Gulf of Guinea, in particular from Ivory Coast to Nigeria and especially Cameroon, Gabon and Congo, appear to have been already occupied at this time by wet tropical forest formations mainly composed of Angiosperms. In the course of the Tertiary period the combined effect of the equator being situated further north than now and the development of the Antarctic ice cap favoured the development of wet tropical conditions over a large part of North Africa. Towards the end of the Tertiary, the equator reached its present position and the northern hemisphere ice caps appeared, and these phenomena resulted in the disappearance of the forest formations spread across the north of Africa, and the concentration of these formations near the equatorial zone around the Gulf of Guinea and in the Congo-Zaire basin. From 800 000 years ago onwards the marked glacial variations at middle and high latitudes in both hemispheres, lowered temperatures in equatorial areas and brought arid climates at times of maximum glacial extension. -Author
Soldierless termites are extremely abundant in South America where they account for up to one-third of the termite species richness. However, their diversity has mostly been addressed through scattered, indecisive descriptions, leaving large gaps in our understanding of termite diversity. Here, to progress towards a more complete knowledge of this group, we redescribe the genus Grigiotermes and describe five new genera, Amplucrutermes, Humutermes, Hydrecotermes, Patawatermes, and Rubeotermes, comprising a total of nine species. We reassign four species to new genera, synonymize two species, and provide extensive descriptions and diagnoses for all taxa included here. We also provide a molecular phylogeny of selected Neotropical and Old World Apicotermitinae and provide a key to the genera of Neotropical workers to facilitate identification.
Termites are among the most important animals
in tropical ecosystems where they often make up over 10 %
of the total animal biomass and enhance ecosystem productivity.
While termites in general have been the focus of a
reasonable amount of work, this effort is not equally distributed
among taxonomic groups. The soil-feeding
Apicotermitinae, in particular, have received less attention
than other taxonomic groups. In terms of species diversity
and abundance, the Apicotermitinae dominate African and
Neotropical rainforests, where they generally feed on soil
organic fractions. Whereas basal Apicotermitinae possess
soldiers, this caste is missing in a large cluster of species,
collectively called the Anoplotermes-group, which possibly
constitutes a monophyletic lineage. These soldierless Apicotermitinae
evolved alternative defensive strategies, such
as defensive body rupture through autothysis or dehiscence.
As species identification in termites is commonly based on
characters of soldiers, the Anoplotermes-group has long
been neglected by taxonomists, but alternative diagnostic
characters, derived from the worker gut topology and enteric
valve structures, are now routinely used. Although species
identification based solely on worker characters is feasible
and new molecular techniques greatly facilitate taxonomic
studies, the biology of soldierless Apicotermitinae remains poorly known. The main objective of this paper is to increase the awareness and understanding of this dominant
soil arthropod through a comprehensive review of their
lifestyle and ecological importance.
A new fossil termite species, the first Rhinotermitinae in amber, Dolichorhinotermes dominicanus, is described based on a single imago trapped in a Dominican amber. The fossil indicates that a species of the Rhinotermes-complex (Rhinotermes, Dolichorhinotermes, Acorhinotermes) was present in the Caribbean region during the Miocene period.
While de novo assembly graphs contain assembled contigs (nodes), the connections between those contigs (edges) are difficult for users to access. Bandage (a B: ioinformatics A: pplication for N: avigating De novo A: ssembly G: raphs E: asily) is a tool for visualising assembly graphs with connections. Users can zoom in to specific areas of the graph and interact with it by moving nodes, adding labels, changing colours and extracting sequences. BLAST searches can be performed within the Bandage GUI and the hits are displayed as highlights in the graph. By displaying connections between contigs, Bandage presents new possibilities for analysing de novo assemblies that are not possible through investigation of contigs alone.
Source code and binaries are freely available at https://github.com/rrwick/Bandage. Bandage is implemented in C++ and supported on Linux, OS X and Windows.
rrwick@gmail.comSupplementary Information:A full feature list and screenshots are available at Bioinformatics online and http://rrwick.github.io/Bandage.
© The Author(s) 2015. Published by Oxford University Press.
Termites are frequently dominant invertebrate decomposers and bioturbators in lowland tropical forests and therefore strongly influence ecosystem processes favouring soil stability, porosity and nutrient retention. In this study, we provide the first spatially replicated dataset on termite assemblage composition, abundance and biomass in a Peruvian rainforest by sampling six separate plots. In addition, two alternative sampling methods (transect method-TM and quadrat method-QM), providing termite species density data, were compared among the plots. The relationships between a range of environmental and spatial variables and species composition were examined using canonical correspondence analysis variation partitioning. We found that the TM captured a higher proportion of the known species in the site (82 %) compared with the QM (66 %). In addition, 56 % of the species sampled by TM were common between the plots while only 18 % of species overlapped using the QM. The QM may therefore potentially have undersampled the species pool. Environmental variables were shown to explain a larger proportion of the species patterns than the spatial variables with elevation, soil temperature and distance to the river being the most important. We discuss the impacts of the environmental and spatial variables on termite species composition.
Termites have colonized many habitats and are among the most abundant animals in tropical ecosystems, which they modify considerably through their actions. The timing of their rise in abundance and of the dispersal events that gave rise to modern termite lineages is not well understood. To shed light on termite origins and diversification, we sequenced the mitochondrial genome of 48 termite species and combined them with 18 previously sequenced termite mitochondrial genomes for phylogenetic and molecular clock analyses using multiple fossil calibrations. The 66 genomes represent most major clades of termites. Unlike previous phylogenetic studies based on fewer molecular data, our phylogenetic tree is fully resolved for the lower termites. The phylogenetic positions of Macrotermitinae and Apicotermitinae are also resolved as the basal groups in the higher termites, but in the crown termitid groups, including Termitinae + Syntermitinae + Nasutitermitinae + Cubitermitinae, the position of some nodes remains uncertain. Our molecular clock tree indicates that the lineages leading to termites and Cryptocercus roaches diverged 170 Ma (153-196 Ma 95% confidence interval [CI]), that modern Termitidae arose 54 Ma (46-66 Ma 95% CI), and that the crown termitid group arose 40 Ma (35-49 Ma 95% CI). This indicates that the distribution of basal termite clades was influenced by the final stages of the breakup of Pangaea. Our inference of ancestral geographic ranges shows that the Termitidae, which includes more than 75% of extant termite species, most likely originated in Africa or Asia, and acquired their pantropical distribution after a series of dispersal and subsequent diversification events. © The Author 2014. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: [email protected]
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We present a new open source, extensible and flexible software platform for Bayesian evolutionary analysis called BEAST 2. This software platform is a re-design of the popular BEAST 1 platform to correct structural deficiencies that became evident as the BEAST 1 software evolved. Key among those deficiencies was the lack of post-deployment extensibility. BEAST 2 now has a fully developed package management system that allows third party developers to write additional functionality that can be directly installed to the BEAST 2 analysis platform via a package manager without requiring a new software release of the platform. This package architecture is showcased with a number of recently published new models encompassing birth-death-sampling tree priors, phylodynamics and model averaging for substitution models and site partitioning. A second major improvement is the ability to read/write the entire state of the MCMC chain to/from disk allowing it to be easily shared between multiple instances of the BEAST software. This facilitates checkpointing and better support for multi-processor and high-end computing extensions. Finally, the functionality in new packages can be easily added to the user interface (BEAUti 2) by a simple XML template-based mechanism because BEAST 2 has been re-designed to provide greater integration between the analysis engine and the user interface so that, for example BEAST and BEAUti use exactly the same XML file format.
Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community.
I present some of the most notable new features and extensions of RAxML, such as, a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX, and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date, 50 page user manual covering all new RAxML options is available.
The code is available under GNU GPL at https://github.com/stamatak/standard-RAxML.
Alexandros.Stamatakis@h-its.org.
A brief overview and dichotomous key is provided for those termites (Isoptera) occurring in middle Eocene (Lutetian) Baltic amber. Ten species in seven genera are presently docu- mented from Baltic amber, these being as follows: Garmitermes succineus n. gen. n. sp., Ter- mopsis bremii (HEER), T. ukapirmasi n. sp., Archotermopsis tornquisti ROSEN, Proelectroter- mes berendtii (PICTET-BARABAN), Electrotermes affinis (HAGEN), E. girardi (GIEBEL), Reti- culitermes antiquus (GERMAR), R. minimus SNYDER, and Parastylotermes robustus (ROSEN). Garmitermes succineus n. gen. n. sp. is the first Mastotermes-like termite discovered in Baltic amber. Hemerobites GERMAR, a senior synonym of Reticulitermes HOLMGREN, is designated as a nomen oblitum under the ICZN rules. Maresa GIEBEL, another senior synonym of Reti- culitermes, is tentatively suppressed pending a petition to the ICZN to preserve the latter name.
Sampling using a replicated standard transect method in an extensive undisturbed primary hill rain forest in Congo (Brazzaville) revealed a very high species richness of termites, especially soil-feeding soldierless termites (Apicotermitinae). The assemblage, as estimated by the transects, resembled that previously characterized in comparable forest in southern Cameroon, but with some species turnover reflecting the gamma diversity of the Guinean-Congolese block as a whole. Species richness was three or four times that shown by a previous study addressed to mound populations alone, emphasizing the importance of sampling soil-feeding termites neither building nor living in mounds.
Modern attempts to produce biogeographic maps focus on the distribution of species, and the maps are typically drawn without
phylogenetic considerations. Here, we generate a global map of zoogeographic regions by combining data on the distributions
and phylogenetic relationships of 21,037 species of amphibians, birds, and mammals. We identify 20 distinct zoogeographic
regions, which are grouped into 11 larger realms. We document the lack of support for several regions previously defined based
on distributional data and show that spatial turnover in the phylogenetic composition of vertebrate assemblages is higher
in the Southern than in the Northern Hemisphere. We further show that the integration of phylogenetic information provides
valuable insight on historical relationships among regions, permitting the identification of evolutionarily unique regions
of the world.
We report a major update of the MAFFT multiple sequence alignment program. This version has several new features, including
options for adding unaligned sequences into an existing alignment, adjustment of direction in nucleotide alignment, constrained
alignment and parallel processing, which were implemented after the previous major update. This report shows actual examples
to explain how these features work, alone and in combination. Some examples incorrectly aligned by MAFFT are also shown to
clarify its limitations. We discuss how to avoid misalignments, and our ongoing efforts to overcome such limitations.
Five forest plots of differing disturbance levels were sampled qualitatively for termites in the Mbalmayo Forest Reserve, southern Cameroon, using 100 m belt transects. Results showed a large reduction in species richness in two severely disturbed plots compared with a Near Primary plot, but little difference in two less disturbed regenerating plots; there is some evidence for a slight increase in species richness in the regenerating plots. Soil-feeders predominate in the primary and regenerating plots, but are greatly reduced in the severely disturbed plots. Wood-feeders appear to be more resilient to disturbance than soil-feeders, although their species richness is low in the most disturbed plots. The Termitinae are the dominant taxonomic group in the Near Primary plot, the Apicotermitinae in the less disturbed regenerating plots and the Macrotermitinae in the clear felled areas. There appears to be no secondary invasion of plots by savanna-associated species, and the small apparent increase in species richness in the less disturbed regenerating plots appears to be due to the influx of forest species usually associated with natural gaps.
Gastrotermes spinatus gen. n. sp. n is described from workers of a single foraging group collected in Cameroon. This soil-feeding termite aligns with the Labidotermes subgroup (Apicotermes group) because of its non-protruding and symmetrical enteric valve armature, its short P1, and its globular P3a. An asymmetrical field of robust sclerotized spines at the opening of the P3a is unique among the other Labidotermes subgroup genera.
The anatomy of the workers' digestive tube is essential in taxonomical studies of soil-feeding Apicotermitinae termites, especially in soldierless lineages. Two structures, the mesenteric-proctodeal junction and the enteric valve, have long been important to distinguish genera and species. By contrast, the gizzard (proventriculus) has been almost ignored by taxonomists because of its generally regressed state in soil-feeding termites. In this study, we document in detail for the first time the sclerotized structures and ornamentations in the gizzard in the Apicotermitinae subfamily. We identified two main clusters of species: those without ornamentations and those exhibiting a sclerotized pulvillar armature, which may include spicules or spines of diverse sizes, numbers and dispositions. The latter group comprises the majority of African soldierless species, a widely diverse and dominant group in tropical forests and savannas. We outline the potential role of the anatomy of the gizzard in the taxonomy of Apicotermitinae based on the interspecific anatomical variation of the pulvillar armatures. We suggest that sclerotized ornamentations regulate the flow of food particles and break or lacerate aggregates to facilitate the access of enzymes in the midgut.
Termitidae comprises ∼80% of all termite species [1] that play dominant decomposer roles in tropical ecosystems [2, 3]. Two major events during termite evolution were the loss of cellulolytic gut protozoans in the ancestor of Termitidae and the subsequent gain in the termitid subfamily Macrotermitinae of fungal symbionts cultivated externally in "combs" constructed within the nest [4, 5]. How these symbiotic transitions occurred remains unresolved. Phylogenetic analyses of mitochondrial data previously suggested that Macrotermitinae is the earliest branching termitid lineage, followed soon after by Sphaerotermitinae [6], which cultivates bacterial symbionts on combs inside its nests [7]. This has led to the hypothesis that comb building was an important evolutionary step in the loss of gut protozoa in ancestral termitids [8]. We sequenced genomes and transcriptomes of 55 termite species and reconstructed phylogenetic trees from up to 4,065 orthologous genes of 68 species. We found strong support for a novel sister-group relationship between the bacterial comb-building Sphaerotermitinae and fungus comb-building Macrotermitinae. This key finding indicates that comb building is a derived trait within Termitidae and that the creation of a comb-like "external rumen" involving bacteria or fungi may not have driven the loss of protozoa from ancestral termitids, as previously hypothesized. Instead, associations with gut prokaryotic symbionts, combined with dietary shifts from wood to other plant-based substrates, may have played a more important role in this symbiotic transition. Our phylogenetic tree provides a platform for future studies of comparative termite evolution and the evolution of symbiosis in this taxon.
A new genus, Tonsuritermes Cancello & Constantini gen. nov., is described from South America. The main morphological features of the new genus are a remarkable frontal gland and protibia with two rows of spine-like bristles. Two new species of Tonsuritermes are described: T. tucki Cancello & Constantini sp. nov. and T. mathewsi Cancello & Constantini sp. nov. Comparisons, measurements, a map, histology of the frontal gland, and illustration of all fundamental morphological aspects are provided.
After more than fifteen years of existence, the R package ape has continuously grown its contents, and has been used by a growing community of users. The release of version 5.0 has marked a leap towards a modern software for evolutionary analyses. Efforts have been put to improve efficiency, flexibility, support for 'big data' (R's long vectors), ease of use, and quality check before a new release. These changes will hopefully make ape a useful software for the study of biodiversity and evolution in a context of increasing data quantity.
Availability:
ape is distributed through the Comprehensive R Archive Network: http://cran.r-project.org/package=apeFurther information may be found athttp://ape-package.ird.fr/.
• Miombo woodlands cover extensive areas in southern and southeastern Africa, but are under high anthropogenic pressure. A conspicuous feature of this ecosystem is the presence of high termitaria (>2 m high) built by fungus‐growing termites (Macrotermitinae).
• Despite the fundamental role of termites in African ecosystems, miombo termite communities remain poorly known. We suspected high termitaria to play a structuring role for the whole termite assemblage of miombo woodlands in southern Burundi. In a formerly cleared area, in regeneration since 2009, we expected the termite assemblage to be highly impoverished. The newly growing termitaria of fungus growers might provide suitable bases for recolonisation by soil‐feeding species.
• We recorded 25 species from more than 1000 termite samples. Fungus growers were abundant everywhere. In preserved miombo, high termitaria were home to secondary soil‐feeding termite species, seldom encountered in the matrix between mounds. Bushes growing on termitaria also sustained wood‐feeding species. The most remarkable feature of the assemblage was the high frequency and diversity of soil‐feeding soldierless Apicotermitinae, especially in the matrix. Besides scarce Coptotermes wood feeders, Macrotermitinae were the sole inhabitants of the regenerating area.
• Our results emphasize the importance of high termitaria for particular soil‐feeding species and wood feeders associated with the vegetation growing on termitaria. They raise the question of the ecological factors allowing the coexistence of a rich assemblage of soldierless Apicotermitinae species in the nutrient‐poor matrix between mounds. The observations from regenerating miombo confirm the vulnerability of soil‐feeding termites to habitat degradation and provide baseline data for future studies of ecosystem restoration.
The left mandible of termite workers possesses just in front of the molar plate a characteristic 'premolar tooth' that, in most species, is partly or wholly hidden under the mandible. The position, structure and size of this tooth were observed and compared from a functional point of view in 46 wood-feeder species belonging to all termite families and in 33 soil-feeder species belonging to 4 different clades of Termitidae. In all wood- and other plant matter-feeder species observed the premolar tooth resembles the chisel of a carpenter's plane-like device. It is suited to cut superficial fragments out of the wood before ingestion. In all soil-feeder species observed the premolar tooth has lost one or several functional features that characterise wood feeders. It assists the other teeth in the gathering of soil particles towards the mouth before ingestion. In the left mandible of termite workers, the premolar tooth thus shows clear morphological adaptations to the species' diet.
The most diverse and best-preserved paleofauna of the higher termites heretofore known, all found in Miocene amber of the Dominican Republic, is described. The imago of Coptotermes priscus Emerson is redescribed, and the soldier of C. priscus, the first known fossil soldier of this genus, is described. The fauna includes the following 29 new species, all in existing genera, with Krishna and Grimaldi as authors of each: in the Rhinotermitidae, two new species based on imagoes of each-Coptotermes hirsutus and C. paleodominicanus; in the Termitidae, 23 new species based on imagoes-Amitermes lucidus, Anoplotermes bohio, A. cacique, A. carib, A. maboya, A. naboria, A. nitaino, A. quisqueya, A. taino, Atlantitermes antillea, A. caribea, A. magnoculus, Microcerotermes insulanus, M. setosus, Nasutitermes amplioculatus, N. incisus, N. magnocellus, N. medioculatus, N. pilosus, N. seminudus, Subulitermes hispaniola, S. insularis, and Termes primitivus; in the Nasutitermitinae four new species based on nasute soldiers-Caribitermes hispaniola, Nasutitermes rotundicephalus, Parvitermes longinasus, and Velocitermes bulbus. This brings the total termite fauna in Dominican amber to four families, 17 genera, and 39 species, a number that exceeds that of the present-day fauna of Hispaniola. Biogeographical, paleoecological, and phylogenetic implications of the Dominican amber termites are discussed.
The taxonomy of Ruptitermes Mathews 1977 is revised. Nine new species are described: R. araujoi, sp. n. (from Brazil), R. atyra, sp. n. (from Brazil, Peru, and Panama), R. bandeirai, sp. n. (from Brazil), R. cangua, sp. n. (from Brazil), R. kaapora, sp. n. (from Paraguay and Brazil), R. krishnai, sp. n. (from Trinidad), R. maraca, sp. n. (from Brazil), R. piliceps, sp. n. (from Brazil), and R. pitan, sp. n. (from Brazil). Ruptitermes franciscoi (Snyder 1959), new combination, is trans-ferred from Anoplotermes Fr. Müller to Ruptitermes. Ruptitermes proratus Emerson 1949 is placed under the synonymy of R. reconditus (Silvestri, 1901). The genus is redescribed to accommodate all the species included. Ruptitermes ar-boreus, R. reconditus and R. xanthochiton are redescribed based on the examination of large series. A key to the 13 species of the genus based on the external morphology of workers is provided. The enteric valve armature is described and illus-trated for all species except R. kaapora. Distribution maps are presented for all species.
Fossil insect taxa from the Ischichuca Formation (late Middle Triassic to early Late Triassic), La Rioja Province (Argentina) are described. One new genus and six new species are proposed: Hermosablatta pygmaea sp. nov. (Blattoptera, Mancusoblattidae), Ademosyne umutu sp. nov., lschichucasyne cladocosta sp. nov. (Coleoptera, Permosynidae), Argentinosyne ischichucaensis sp. nov. (Coleoptera, Schizocoleidae), Babuskaya elaterata gen. et sp. nov. (Coleoptera, Elateridae?) and Argentinocupes sara sp. nov. (Coleoptera, Cupedidae?). Other previously described Triassic insect species, such as Ademosyne punctuada MartinsNeto and Gallego, Argentinosyne frenguellii Martins-Neto and Gallego, and Argentinocupes pulcher MartinsNeto and Gallego are reported for the first time from this formation.
The history of megathermal (currently ‘‘tropical’’) rainforests over the last 30 kyr is now becoming relatively well-understood, as demonstrated by the many contributions in this volume. However, our perception of their longer-term history remains highly fragmentary. There is a real need for a better understanding of rainforest history on an evolutionary time scale, not only to have a better idea of the biological, geological, and climatic factors which have led to the development of the most diverse ecosystem ever to have developed on planet Earth, but also since the implications of rainforest history on an evolutionary time scale are inextricably linked to a plethora of other issues currently receiving wide attention. Determining the place and time of origin and/or radiation of angiosperms (which overwhelmingly dominate present day megathermal rainforests), establishing patterns of global climate change, clarifying the nature of global temperature gradients through time, understanding the successive switching from greenhouse to icehouse climates, global warming, patterns of dispersal of megathermal plants and animals, higher rank (ordinal) taxonomy and the nature of controls on global diversity gradients are but some issues which are being clarified with the better understanding of the long-term history of megathermal rainforests.
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing
methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform
(FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid
residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy
of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length.
Two different heuristics, the progressive method (FFT‐NS‐2) and the iterative refinement method (FFT‐NS‐i), are implemented
in MAFFT. The performances of FFT‐NS‐2 and FFT‐NS‐i were compared with other methods by computer simulations and benchmark
tests; the CPU time of FFT‐NS‐2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT‐NS‐i is over
100 times faster than T‐COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
A comprehensive compendium on the taxonomy and biology of the 3106 living and fossil species of the worlds termites is presented, along with reviews of Isoptera morphology and evolution, identification keys, the history of termite systematics, and summary of the worlds 363 significant pest species. A complete bibliography is provided of nearly 5000 references covering virtually all aspects of termite taxonomy and biology through December 2011.The morphology of Isoptera is thoroughly reviewed and illustrated with original scanning electron micrographs and photomicrographs, covering the cuticular anatomy and those internal organs that are taxonomically and phylogenetically significant, including several new character systems. Terminology is presented for the systems of tibial spines and spurs so as to establish homologs. Keys are presented to the nine living families of termites, and the world subfamilies and genera of Archotermopsidae, Hodotermitidae, Kalotermitidae, Mastotermitidae, Rhinotermitidae, Serritermitidae, Stolotermitidae, and Stylotermitidae. A key to subfamilies of the Termitidae is included. A detailed morphological diagnosis for each family and subfamily is provided, along with images of exemplar species. The history of isopteran research in taxonomy, systematics, morphology, paleontology, and biology is reviewed from 1758 to the present, with emphasis on transformative workers such as Holmgren, Silvestri, Emerson, Roonwal, Noirot, and Sands. Evolution of the Isoptera is reviewed, including the diversity and natural history of genera and species in all Zoogeographic regions, major patterns in social biology, the phylogeny of Recent and fossil genera and families, and 135 million years of fossils preserved as compressions, mineralized replicas, and in amber. The definitive sister group to the Isoptera is the monogeneric family of wood roaches, Cryptocercidae (Cryptocercus), so the taxonomic ranks of the two groups are now Infraorder Isoptera and Infraorder Cryptocercoidea within Order Blattaria (roaches and termites).The compendium summarizes the taxonomic history, nomenclature, distribution, type locality, and repository, and all significant aspects of natural history and biology for each species of the world, exclusive of pest control and colony inquilines (termitophiles). The classification of Recent and fossil lower termites (all those exclusive of family Termitidae) used in the compendium is from Engel et al. (2009), which is based on morphology and largely congruent with molecular studies. Rhinotermitidae s.s. (exclusive of Stylotermitidae) may be paraphyletic with respect to Termitidae, although the six traditional subfamilies of the former are used here. A separate section summarizes the nomenclatural changes made in the compendium, including new synonymies, new combinations, status novus, lectotype selection etc. A detailed list is provided of museums and other institutional collections that house type specimens. An index is included. The Treatise is intended to provide an authoritative foundation for taxonomic work on the Isoptera, present and future.
I present a new statistical method for analysing the relationship
between two discrete characters that are measured across a group of
hierarchically evolved species or populations. The method assesses
whether a pattern of association across the group is evidence for
correlated evolutionary change in the two characters. The method takes
into account information on the lengths of the branches of phylogenetic
trees, develops estimates of the rates of change of the discrete
characters, and tests the hypothesis of correlated evolution without
relying upon reconstructions of the ancestral character states. A
likelihood ratio test statistic is used to discriminate between two
models that are fitted to the data: one allowing only for independent
evolution of the two characters, the other allowing for correlated
evolution. Tests of specific directional hypotheses can also be made.
The method is illustrated with an application to the Hominoidea.
Soil-feeding termites are dominant members of the soil fauna in lowland tropical rainforests. As ecosystem engineers, they have a profound effect on their environment, particularly through modification of the vast quantities of soil that they ingest. There is growing evidence that the processing of the soil in the gut is influenced by the enteric valve—an extremely well-developed feature in the hindgut of the Termitidae, consisting of six ridges, variously armed with spines, teeth and scales. Although this valve has been extensively used in morphological work, little is known of its function. Scanning electron microscopy has been used in this study to better understand the three-dimensional structure of the enteric valve in the Afrotropical Apicotermitinae, a group in which these valves are everted into the following chamber of the hindgut. This configuration lends itself to scanning electron microscopy in a way that it does not in other soil-feeders, since in those species the armature is obscured within the gut lumen. It seems plausible that this structure is instrumental in filtering material through the gut in such a way that the coarse, organic-poor portion of the soil is channeled fast through the centre, while directing the fine, organic-rich portion to the edges for lengthier digestion of recalcitrant plant material. Thus, the enteric valve would appear to be important for the termites to survive on this nutrient-poor but super-abundant food resource.
Summary A new dragonfly is described from the Lower Cretaceous limestones of the Crato Forma- tion (Brazil): Araripechlorogomphus muratai n. gen. n. sp. (Araripechlorogomphidae n. fam.) which clearly belongs to the chlorogomphid clade within Anisoptera. It is the first fossil record of Chlorogomphida and also the first New World record of this dragonfly group that is now exclusively distributed in East Asia. The phylogenetic and biogeographic implications of this new discovery are discussed. As consequence of the phylogenetic analysis the new higher taxa Cristotibiata, Paucipostnodalia, Eubrachystigmata, Neobrachystigmata, Paneury- palpidomorpha, Eurypalpidomorpha, and Eurypalpidiformia are introduced. A monotypic family Juracorduliidae n. fam. is established for the genus Juracordulia, and the genus Prohe- meroscopus is transferred from Nannogomphidae to a new monotypic family Prohemero- scopidae n. fam. within Paucipostnodalia as sistergroup of Eubrachystigmata.
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform (FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length. Two different heuristics, the progressive method (FFT-NS-2) and the iterative refinement method (FFT-NS-i), are implemented in MAFFT. The performances of FFT-NS-2 and FFT-NS-i were compared with other methods by computer simulations and benchmark tests; the CPU time of FFT-NS-2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT-NS-i is over 100 times faster than T-COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
A new subfamily, genus, and species, Archeorhinotermitinae, Archeorhinotermes rossi, from Burmese amber, dated as Turonian-Cenomanian (90–100 mya) of the Cretaceous period, are described and figured. Comparisons are made between the other subfamilies of the Rhinotermitidae and the new subfamily. This is the first fossil record of the family Rhinotermitidae from the Cretaceous.
Termites are an important component of tropical rain forests, and have been included in many studies focusing on the influence of human disturbance. Their distribution among primary rain forest has, however, rarely been investigated. Here we studied the termite fauna in seven mostly undisturbed forest sites, representing several rain forest types. Overall, approximately 70 percent of species were soil-feeders and 25 percent were wood-feeders, the remaining 5 percent being classified here as litter-feeders. Termite species richness did not differ significantly among sites, but sites differed in termite abundance. The palm swamp and the low forest situated on the foothills of an inselberg, hosted different termite communities to the other sites. These two sites presented a singular physiognomy suggesting that forest type is an important factor influencing species composition. We found no correlation between termite species composition and distance between sites, highlighting that at the scale of our study (about 100 km), forest sites share a similar species pool.
The gut of the Termitidae is much more diverse than that of the previously studied "lower" termites (NOIROT, 1995). Nevertheless the monophyly of this family is confirmed. The characters of the mesentero-proctodeal junction (presence and morphology of the mixed segment) and of the Malpighian tubules (attachment, fusion, closure, presence of a "Malpighian knot") seem especially significant. The gut allows the definition of twelve groups of genera. The Macrotermes group is equivalent with the subfamily Macrotermitinae. The Apicotermes, Anoplotermes and Speculitermes groups make the Apicotermitinae, the monophyly of which is also confirmed. Among the Nasutitermitinae the genera with mandibulate soldiers (Syntermes group) are very distinct from the genera with nasute soldiers (Nasutitermes and Subulitermes groups), which suggests a paraphyly of the Nasutitermitinae. The genera now included in the subfamily Termitinae can be distributed into five groups. The Cubitermes group is very homogeneous. The genera having soldiers with "snapping" mandibles could make two distinct groups: Termes and Pericapritermes. The Amitermes group seems more heterogenous. The gut of the Foraminitermes group is idiosyncratic (new subfamily?). Thus the Termitinae don't make a clade. At least the Foraminitermes group must be removed. Finally a phylogeny of the Termitidae is proposed. As it is based essentially on the gut characters it must not be regarded as more than a working hypothesis.
The history of megathermal (currently ‘‘tropical’’) rainforests over the last 30 kyr is now becoming relatively well-understood,
as demonstrated by the many contributions in this volume. However, our perception of their longer-term history remains highly
fragmentary. There is a real need for a better understanding of rainforest history on an evolutionary time scale, not only
to have a better idea of the biological, geological, and climatic factors which have led to the development of the most diverse
ecosystem ever to have developed on planet Earth, but also since the implications of rainforest history on an evolutionary
time scale are inextricably linked to a plethora of other issues currently receiving wide attention. Determining the place
and time of origin and/or radiation of angiosperms (which overwhelmingly dominate present day megathermal rainforests), establishing
patterns of global climate change, clarifying the nature of global temperature gradients through time, understanding the successive
switching from greenhouse to icehouse climates, global warming, patterns of dispersal of megathermal plants and animals, higher
rank (ordinal) taxonomy and the nature of controls on global diversity gradients are but some issues which are being clarified
with the better understanding of the long-term history of megathermal rainforests.