Zoology in the Middle East
Supplementum 4, 2012
Advances in Earthworm Taxonomy V (Annelida: Oligochaeta)
Proceedings of the 5th International Oligochaeta Taxonomy Meeting Beatenberg, Switzerland, 11-15 April, 2011.
Price per article (irrespective its length): EUR 10.00 plus handling/postage
Zoology in the Middle East, Supplementum 4, 2012: 3. | Free download (PDF) (OPEN ACCESS)
Robert J. Blakemore
Restating scope of genus Metaphire Sims & Easton, 1972: 40 years on (Oligochaeta)
Abstract. Clarification of correct placement of species in genus Metaphire Sims & Easton, 1972 is required as differentiation from Amynthas Kinberg, 1867 hinges on condition of male pores in their respective type-species. Confusion is due in part to misconception of rules of ICZN and partly because of problems of parthenogenetic degradation of male organs. In summary, Amynthas is the default genus of pheretimoids with superficial or absent male pores, Metaphire species differs by having non-superficial male pores and in Pheretima Kinberg, 1867, which is not known in East Asia, taxa further acquire nephridia on spermathecal ducts. The type of Metaphire, M. javanica (Kinberg, 1867), was misidentified in Australia but M. californica (Kinberg, 1867), its possible synonym, is widespread and is now confirmed in Korea. This latter cosmopolitan is still often confused with Japanese Duplodicodrilus schmardae (Horst, 1883) that has eversible intromittent organs developed much more so than in Metaphire. Figures are provided.
Key words. Taxomonic priority, genera, pheretimoids, Asian earthworms.
Robert J. Blakemore
Japanese earthworms revisited a decade on (Oligochaeta: Megadrilacea)
Abstract. This paper summarizes eco-taxonomic progress since 2001. Five new natives were described and five new exotics recorded plus herein Megascolecid Amynthas rodericensis (Grube, 1879) is retrieved from historical records, Acanthodrilid Microscolex dubius (Fletcher, 1887) is a new Asian record and Lumbricid Dendrodrilus rubidus subrubicundus (Eisen, 1874) morph from Lake Biwa is added to checklist a total of 96 Megadrile earthworm species in eight families from Japan. Approximately 33 are natives and same number are exotics with the balance having insufficient information to decide their origins (e.g. for Eisenia japonica hiramoto Blakemore, 2012). Synonyms are provided especially for parthenogenetically affected morphs which can only be separated via their type DNA. Criodrilid Criodrilus bathybates Stephenson, 1917 is restored as originally described in Criodrilidae and its COI nucleotide barcode is for the first time presented on GenBank.
Key words. Amynthas, Metaphire, pheretimoids, Japan, Megadrile Oligochaeta.
Charlène Briard, Jiang-Ping Qiu, Qi Zhao, Muriel Guernion, Daniel Cluzeau
Phylogenetic study of some Aporrectodea species based on molecular markers (Oligochaeta: Lumbricidae)
Abstract. The Nicodrilus genus is not accepted in the nomenclature according to ICZN; Aporrectodea term is used. However, according to Bouché, Aporrectodea and Allolobophora genera are not homogeneous and are supposed as polyphyletic genera. The aim is to study the phylogenetic structure of Aporrectodea genus in order to verify its cladistic nature and its taxonomical validity. In this work, five species, belonging to the Aporrectodea genus, as the most common in France are studied. First, we use usual morphological characteristics to identify each species. Species life history traits are included among morphological characteristics (e.g. clitellum, puberculum and pores position, Morren’s gland, body size). Then, we pursue a molecular approach on individuals sampled in France. Molecular phylogenetic analyses are based on the sequences of nuclear (rDNA 28S) and mitochondrial (COI, rDNA 16S) gene regions and performed with maximum likelihood and bayesian inference. Phylogenetic analyses revealed two deep sister clades, Aporrectodea caliginosa species complex in one hand, and A. giardi, A. longa and A. nocturna in the other hand. The status of A. nocturna is recognized instead of A. longa and A. giardi ones. Our study cannot confirm the validity of Aporrectodea genus but highlights different lineages within species such as A. longa and A. caliginosa. Markers characterization ensures a molecular genotyping in lumbricids and begins to reveal intraspecific variabilities degrees unsuspected.
Key words. Phylogeny, evolution, COI, rDNA16S, rDNA28S, Aporrectodea.
Tomáš Pavlíček, Yarin Hadid, Csaba Csuzdi
Opening Pandora’s box: Clitellum in phylogeny and taxonomy of earthworms (Oligochaeta)
Abstract. The solution to the current contradictions in earthworm taxonomy and phylogeny is a better understanding of the underlying speciation process. The analysis of size and distribution of clitellar segments and that of tubercula pubertatis in the model homoploid genus Lumbricus provides prima facie evidence for the occurrence of the intra-chromosomal autohomoploid hybridization (autohomoploid hybridization = hybridization without changing the ploidy level and taking place between parents from the same panmictic population). The inferred principal mechanism of the autohomoploid hybridization is an unequal crossing recombining paralogous genes organized in a genomic island of divergence (called here “clitellar genomic island of divergence”, CGID). Since clitellar segments constitute a prezygotic reproduction barrier and seem to correspond to the underlying genes at CGID on a one-to-one basis, their analysis helps to illuminate the underlying speciation process. The inferred characteristic features of the autohomoploid hybridization in earthworms are: (1) Presence of CGID; (2) Generation of quantitative changes in CGID leading to speciation by means of unequal crossing over (= separation of the process leading to speciation from the rest of genome); (3) Regulated distribution of breaking points; (4) Intra-lineage hybridizations, and (5) Homeotic character of the CGID genes. As far as we know, this is the first case of autohomoploid hybridization described in animals. Probably, it is not exaggerated to conclude that many earthworm evolutionary lineages (species) originated in the process of the described autohomploid hybridization in the CGID or in the process of inter-chromosomal duplications leading to polyploidization of the whole genome. We do not deny the possible existence of allopatric speciation in earthworms caused, for example, by established geographic or behavioural (e.g., assortative mating) barriers to inter-population gene flow. The major consequences of ignoring autohomoploid hybrid speciation in lumbricid earthworms (and presumably in other earthworm families as well) in phylogenetic analyses are: (i) Incorrect inference of phylogenies by applying bifurcating-like phylogenetic analysis instead of reticulate analyses as often seen in the low statistical supports for different clades in constructed bifurcating-like phylogenetic trees and for trees topologies (frequently not even tested). (ii) Misinterpretation of taxonomy and phylogeny by using genetic distance as the sole defining criterion.
Key words. Erthworms, speciation, clitellum, tubercula pubertatis, Lumbricus, hybridization.
Zoology in the Middle East, Supplementum 4, 2012: 31-46. . | Free download (PDF) (OPEN ACCESS)
Jadwiga Danuta Plisko
Notes on the status of the family Microchaetidae (Oligochaeta)
Abstract. Short historical overview of the variable taxonomic rank of the family Microchaetidae is given. The dispersal of the South African endemic taxa based on paleo-geological evidence is noted. Present familial status and the generic composition are outlined. A correlation between specific features and the species distribution is indicated. Selected familial and generic diagnostic characters and their plesiomorphic and apomorphic conditions are discussed. The possible relationship between Microchaetidae and other families sensu various authors is marked out.
Key words. Microchaetidae, Microchaetus, Geogenia, Tritogenia, Michalakus, Proandricus, Kazimierzus, South Africa, endemic, biogeography.
Zoology in the Middle East, Supplementum 4, 2012: 47-58. | Free download (PDF) (OPEN ACCESS)
Victor V. Pop, Adriana Antonia Pop, Csaba Csuzdi
An annotated checklist of the Romanian earthworm fauna (Oligochaeta, Lumbricidae)
Abstract. This paper updates Pop’s (1938-1964) checklists of Romanian Lumbricidae with the taxa newly described, relocated or modified since then. It comprises a comparative outlook which briefly reflects also the history of the lumbricid systematics during the last seven decades. The checklist is based on a thoroughly re-examined material kept in the Zoological Museum of the Babes-Bolyai University, Cluj-Napoca, and in the Biological Research Institute in Cluj-Napoca. A comparative table shows the taxa accepted by Pop, an up-dated list of the recently described species, and some dubious specie either relocated or eliminated. The earthworm fauna of Romania is quite well known; in the last 45 years, the list was augmented to 71 taxa. As compared with the fauna of the surrounding countries, the lumbricid earthworm fauna of Romania is very rich in number of species and especially in number of endemic elements. The 71 earthworm species found in Romania can be assigned to 12 zoogeographic categories dominated by the Dacian (29%) and Carpathian (14%) endemics.
Key words. Earthworms, Lumbricidae, Romania, checklist.
Life forms in Oligochaeta: a literature review
Abstract. The main life forms among the Oligochaeta s.l. (= Clitellata) and the related polychaetes are: aquatic (freshwater and marine) sediment-dwellers, inhabitants of the macrovegetation, large and small soil-dwellers, and carnivores. The vegetation-dwellers (Naididae, Pristinidae and Opistocystidae) reproduce mostly in an asexual way; some of them have an ability to swim and posses eyes. A convergent group to the naidid oligochaetes is the aphanoneuran genus Aeolosoma. The smaller Enchytraeidae, and the larger “earthworms” (= Megadrili) Crassiclitellata and Moniligastridae, live in the terrestrial soil. Some Enchytraeidae and Crassiclitellata are secondarily aquatic while some (generally aquatic) tubificids can facultatively live in the soil. Carnivory (as parasitism, commensalism or predation) has been developed in separate genera of several families. A large clade, including the Hirudinea, the Acanthobdellidae and the Branchiobdellidae, is highly adapted for carnivory (suckers, jaws, loss of chaetae, etc.). Two evolutionary trends are evident in different clades: reduction in chaetal number from indefinite to two per bundle, or to complete loss of chaetae, and reduction of the upper tooth in the originally bifid sigmoid chaetae. External gills have appeared at least in four independent cases. There exist many convergencies in the mode of life and morphology of separate Oligochaeta and related “oligochaetoid” polychaetes.
Key words. Oligochaeta, Clitellata, Polychaeta, life forms, characters, convergency.
Zoology in the Middle East, Supplementum 4, 2012: 71-82. | Free download (PDF) (OPEN ACCESS)
Eduardo Boothby Carlo, Sonia Borges, Mónica Alfaro
Abundance and distribution of Pontodrilus litoralis in the shores of the Cabo Rojo Lighthouse, Puerto Rico (Oligochaeta)
Abstract. The structure and dynamics of the populations of the oligochaete Pontodrilus litoralis on the coasts near the Cabo Rojo Lighthouse in Puerto Rico were studied. Sampling was made at low tide on the sandy littoral shores of two beaches (protected and exposed) on each side of the lighthouse. Environmental factors such as substrate temperature, pH, soil moisture and organic matter content, and anoxic sand and surface accumulation of plant debris were registered. Spatial distribution of P. litoralis was determined to be aggregate in clusters which were predominant right at the high tide marking where sand was always humid but never completely submerged under seawater. Organic matter and temperature were the primary factors in the distribution of the oligochaete. Sites with the lowest temperatures recorded seemed to have the most number of individuals. Those with the most accumulation of surface plant debris recorded the lowest temperatures as well as the highest organic matter contents. There were no significant differences in earthworm density between beaches or collection time. New collection sites for the species in Puerto Rico are also included.
Key words. Neotropical, Caribbean, Annelida, Megascolecidae, Acanthodrilidae.
Ghayoumi Razieh, Latif Robabeh
First earthworm records from the Mouteh Wildlife Refuge, Iran (Oligochaeta)
Abstract. A earthworms survey was conducted in the Mouteh Wildlife Refuge at six stations in April 2010. The refuge, encompassing about 220,000 ha of the natural reserve, is located in the northwestern part of the Isfahan province and south of the Markazi province that is arid to semi-arid and warm in summer. The surveyed region is located far from human activities, and we expected that its isolation served as a geographic barrier preventing human-based earthworm introductions. Recorded earthworms belong to three genera of the family Lumbricidae, and include the following five species: Aporrectodea caliginosa (Savigny, 1826), A. rosea (Savigny, 1826), Dendrobaena hortensis (Michaelsen, 1890), D. veneta (Rosa, 1886), Eiseniella tetraedra (Savigny, 1826). In contrast to our expectations, the recorded species, except D. veneta, are introduced in Iran.
Key words. Earthworms, Lumbricidae, Mouteh Wildlife Refuge, Isfahan and Markazi provinces, Iran.
Earthworm diversity in urban habitats of Basel (Northwestern Switzerland) (Oligochaeta: Lumbricidae)
Abstract. The diversity and ecology of the soil fauna in Central European towns is not yet well known. There are references that earthworms, by their biomass, dominate all other animal groups in cities – surpassed only by man. Basel (11 km²) is, with its 170 000 inhabitants, Switzerland’s third largest community. Open green spaces in the city are scarce; the green belt is narrow. Nevertheless, in more than 60 locations studied between 1999 and 2010, 22 species (12 genera) of Lumbricidae were found. Eleven near-natural habitats could be sampled: private front gardens and backyards, wooded river sides, industrially watered flood plain forest plots, and a Zoological Garden with a large number of green spaces built up from imported soil. Nearly a dozen of these species are quite rare in their natural distribution area and in Basel; many of them are riparian, e.g. Aporrectodea georgii, Fitzingeria platyura depressa, and Helodrilus oculatus – while two are strictly endogeic (Allolobophora satchelli, Murchieona muldali). The sampling method “digging and hand-sorting plus mustard meal suspension” yielded the best possible quantitative results in urban habitats. A maximum of earthworm species (18) was sampled in the Zoological Garden, and the intensely used public lawns had the highest mean density and biomass (450 individuals resp. 280 g fresh mass/m²).
Key words. Quantitative sampling, public parks, private gardens, green belt, woodland, groundwater recharge plants, synanthropic species.
Distribution of earthworms belonging to families Acanthodrilidae, Criodrilidae, and Megascolecidae in Turkey (Oligochaeta)
Abstract. Out of 76 earthworm species recorded in Turkey, 70 species belong to family Lumbricidae, three species to family Megascolecidae (Amynthas corticis (Kinberg, 1867), A. gracilis (Kinberg, 1867), and Metaphire californica (Kinberg, 1867)), two species to family Acanthodrilidae (Microscolex phosphoreus (Dugès, 1837) and M. dubius (Fletcher, 1886)) and Criodrilus lacuum Hoffmeister, 1845 to the family Criodrilidae. The presence and distribution in Turkey of these species is described.
Key words. Earthworms, Fauna, Turkey, Megascolecidae, Acanthodrilidae, Criodrilidae.
Tomáš Pavlíček, Csaba Csuzdi
Earthworm fauna of French Guiana (Oligochaeta)
Abstract. The taxonomic diversity of earthworms in eastern Amazonia is almost unknown. In French Guiana (83,534 km²) 22 species are recorded, of which about 40% are introduced. Despite of the low number of recorded taxa, the taxonomic diversity in earthworms in the humid rain forest could be very high as indicated by the description of five species and a new genus in French Guiana in 2011.
Key words. Amazonia, rain forest, earthworm, species diversity.
Zoology in the Middle East, Supplementum 4, 2012: 107-110. | Free download (PDF) (OPEN ACCESS)
Satyendra M. Singh, Om Prakash
Species richness and density of earthworm populations in grasslands of western Uttar Pradesh, India (Oligochaeta: Megascolecidae, Octochaetidae)
Abstract. This study presents the first survey of earthworm species from the grasslands of Uttar Pradesh, a northern state of India. The study was conducted in all five political regions (e.g., Agra, Bareilly, Meerut, Moradabad and Saharanpur) of western Uttar Pradesh, during different climatic seasons from the year 2007 till 2009. Seven species belonging to two earthworm families were recorded: Metaphire posthuma (Vaillant), Lampito mauritii (Kinberg, 1867) and Perionyx excavatus (Perrier, 1872) from Megascolecidae and Eutyphoeus i.e. E. waltoni (Michaelsen, 1907), E. gigas (Stephenson, 1917), E. orientalis (Beddard, 1883) and E. pharpingianus (Michaelsen, 1907) from Octochaetidae. Moradabad was identified as the region of western Uttar Pradesh showing the highest species richness of earthworms during rainy season (July to October). Species richness and density of worms were assessed the maximum in Moradabad and the minimum density in Saharanpur during peak winter months i.e. from November to February. However, species richness of worms was found the least in Meerut region during winter and summer months and Saharanpur during summers (March to June). M. posthuma was found in all the regions round the years but L. mauritii and E.waltoni have been reported from all the regions only during the rainy season. P. excavatus was next in occurrence pattern, as it was recorded from all the regions except from Meerut. E. pharpingianus was found in all the regions except Meerut. Occurrence of E.orientalis and E. gigas was recorded rarely.
Key words. Earthworm, species richness, population density, grassland, climatic factors, Uttar Pradesh state.
Alexandra D. Solomou, Athanassios I. Sfougaris, Evangelia M. Vavoulidou, Csaba Csuzdi
The effects of farming practices on earthworm dynamics in olive groves of central Greece (Oligochaeta)
Abstract. We compared species richness, species composition, density and biomass of earthworm communities in organically and conventionally managed olive groves in central Greece. Earthworm biomass and density was significantly higher (p<0.05) in organic olive groves than in conventional olive groves. The highest species richness was recorded in the organic olive groves. Six species were recorded in total: Octodrilus complanatus, O. croaticus, Dendrobaena byblica, D. veneta, Aporrectodea caliginosa, and Microscolex phosphoreus. Only four of these were found in conventional olive groves, where D. veneta and M. phosphoreus were absent. The most abundant species were O. complanatus, D. veneta, and D. byblica.
Key words. Olea europaea, earthworms, organic farming, conventional farming, soil properties.
Ghanshyam Tripathi, Kesu Ram Panwar
Biotic interaction of earthworm in arid pedoecosystem (Oligochaeta)
Abstract. Presence of a faunal group may influence the density of other fauna in pedoecosystem in arid environment. Therefore, season-dependent impacts of earthworms on populations of some important groups of soil fauna were studied. Earthworm associated alterations in populations of Collembola, mites, beetles, woodlice and pseudoscorpions showed remarkable results. Abundances of these soil faunal groups were higher at the sites with earthworms as compared to the sites without earthworms throughout the year. On the one hand, the highest abundance of Collembola and beetles were recorded in July. On the other hand, mite, woodlouse and pseudoscorpion showed their highest abundances in August at the sites with earthworms. The present findings suggest earthworms-induced enrichment of below-ground faunal biodiversity in a desert pedoecosystem.
Key words. Earthworm, Pedoecosystem, Seasonality, Biota interaction, Desert.
Ghanshyam Tripathi, Kesu Ram Panwar
Earthworm fauna of Indian Thar Desert (Oligochaeta)
Abstract. Studies were carried out to explore earthworm fauna in a part of Indian Thar Desert. In addition to species diversity, the qualitative composition, density, biomass, biodiversity indices and habitat relationship of earthworms were studied. There were apparently abundant, moderate and low distribution of earthworm in different pedoecosystems. Maximum density of total earthworm species was in grassland, while it was minimum in cultivated field. Biomass was also maximum in grassland but it was minimum in bare land. Species richness varied from 4-6 in arid environment. Evenness and species diversity indices were highest under natural plantation and lowest in bare land. Some kind of species-habitat relationship appeared in relation to physicochemical characteristics of soil system. In spite of a low population density of earthworm in desertic soil system, species diversity was appreciable in desert region.
Key words. Earthworm, biomass, density, species diversity, habitat-relationship, desert.
Qi Zhao, Daniel Cluzeau, Charlène Briard, Jing Sun, Jibao Jiang, Muriel Guernion, Jiang-Ping Qiu
Hainan earthworm community and the comparison with other East and Southeast Asia countries for geographic distribution and endemic rate (Oligochaeta)
Abstract. This paper presents an up-to-date list of the terrestrial earthworms of Hainan Island, which is the second largest island of China and separated by Qiongzhou Strait from the Leizhou Peninsula, Guangdong Province. Actually, 45 species are already known in this island, 60% of which is native species. Ten new species which were collected in the fieldwork in 2006 are also included in the list. Of these 45 species, 82% belongs to the family Megascolecidae, with the dominant genus Amynthas (70%). Some of the species also disperse in other regions of the Southeast Asia and China mainland, except 5 species only have a limited distribution. The endemism rate of Hainan (60%) is similar with that of China mainland (66%) and Taiwan (58%), but its endemism rate of Amynthas (77%) is much higher. All of these results maybe have some links with the geographical history of Hainan Island.
Key words. Earthworm, biodiversity, distribution, endemism rate, Hainan Island.
First record of reproduction by fragmentation in the genus Marionina (Oligochaeta: Enchytraeidae)
Abstract. Hermaphrodite enchytraeids usually breed sexually, but a few species can also multiply asexually by fragmentation (architomy). These species can alternate sexual breeding with asexual breeding by fragmentation. In Hungary, a Marionina species was found in the botanical garden of University of Szeged in 209, which is also able to reproduce asexually by architomy. No mature specimens of this species have been found so far. Fragmentation was observed rarely in these individuals and a strategy comparable to other fragmenting species is not known.
Key words. Enchytraeidae, Marionina, architomy, green houses.
Geographic differences in the setal pattern of Eisenia fetida (Savigny, 1826) in Iran (Oligochaeta: Lumbricidae)
Abstract. The intra-specific variations of the setae arrangements was evaluated in three populations of Eisenia fetida (Savigny, 1826), e.g., to apply and test of the usefulness of setal formula at the intra-specific (population) level. The results confirm the role of the expected geographic isolation on the inter-population variability, and the results showed that variability in the setal formula (variability in paired setae groups aa, ab, and dd) could be used in the study of the intra-specific variations.
Key words. Iran, earthworm, setae, Eisenia.
Robert J. Blakemore
Call for a Census of Soil Invertebrates (CoSI)
Abstract. A case is argued for urgent reappraisal of biodiversity of soils in order to mitigate their rapid global decline (MEA). A first step is to compile a “stock inventory” of soil fauna thought to number around 210,000 species. Basic knowledge is yet wanting and even earthworms are poorly known despite being the major component as well as key "environmental-engineers" and vital links in all terrestrial food webs (including in waterways, hence their excellence yet trivial use as bait). That biodiversity of earthworms is disproportionately underappreciated is surprising as, with 10,000 already named and many more expected, they are no less species-rich than marine polychaetes, for example, that number ca. 8,000 valid taxa. A model for CoSI is the 10 yr, $1 billion global Census of Marine Life (CoML) that concluded with 250,000 total ocean taxa, but since 2 million species are already catalogued and estimates of diversity are of 10 million, this represents 12.5% of described species and just 2.5% of a probable total. Even claims that oceans occupy two thirds of the planet overlooks that land is hilly and the relative surface areas are perhaps 50:50. Socio-economic arguments flounder in context of 99% of the total worldwide human food supply produced on land, whereas oceans and other aquatic ecosystems provide a paltry 0.6% (FAO). Thus it seems timely and appropriate to advocate a sea change to firmly ground eco-taxonomic studies on our diminishing soils that support all life on Earth and, via runoff, provision or pollute the oceans too.
Key words. Biodiversity inventory, Annelida, Oligochaeta, extinction, taxonomy.
Maxim P. Shashkov
Lumbricus – database on earthworms ranges (Oligochaeta)
Abstract. The great volume of information on earthworms’ fauna and population has been accumulated in literature by now. It is necessary to organize these data into an information system for faunistic, ecological, taxonomical and other investigation. The subject of the paper is to develop versatile system for storing, systematization and processing of data on earthworms’ geographical distribution and ecological ranges. Open source database and GIS solutions were used to develop the system further referred to as Lumbricus database. The goal of this implementation is to cover blank spaces on Russian territory observed in data on earthworm’s biodiversity available in World Wide Web. Lumbricus database provides some information on geographical distribution and ecological characteristics of lumbricid species of Russia both in English and Russian. It is not a comprehensive literature data collection, but just a prototype for further developing into worldwide accessible system of full value. There are more than 700 records in the database now, but it is widening permanently. The Lumbricus database is available online since December, 2011 on www.biocenos.org website.
Key words. Earthworm range, database, open-source software, biodiversity.
Emma Sherlock, Louise Berridge
History of the earthworm collections at the Natural History Museum, London (Oligochaeta)
Abstract. The Natural History Museum London houses one of the largest and most historic earthworm collections in the world. Despite some periods of inactivity, largely due to international events and a lack of general enthusiasm by collectors worldwide in some eras, it has continued to grow. Periods of rapid expansion of the collection coincide with eras of resident Oligochaete researchers at the Museum, however, the general growth can be in part attributed to Oligochaetoligists world wide and the assistance they provided each other and their respective institutions, something they continue to do to this day. This paper highlights the periods of growth, explains the times of inactivity and highlights the collection for future depositions and further utilisation by researchers worldwide.
Key words. Natural History Museum, collections, earthworms, history, oligochaete.