A systematic appraisal of the Eriophyoidea (Acari Prostigmata)

The diversity of the Eriophyoidea is largely unknown and their systematic study mostly entails alpha-taxonomy which is critically important for these mites. Eriophyoid morphology is almost exclusively studied on slide-mounted specimens, and truly permanent specimen slides cannot be prepared and are eventually lost. Shortcomings in taxon descriptions are persistent, and too few morphological characters are available for systematic use, particularly for phylogenetic studies. The fragile, simplified and minute eriophyoid bodies, and the inadequacy of study methods and technology, including preparation and light microscopy, contribute to these problems. The present eriophyoid classification is widely accepted, relatively stable and useful. The major part of the classification, however, is probably artificial, and some taxon delimitations and identifications are becoming increasingly difficult. Scanning electron microscopy (SEM) is only sporadically used to supplement conventional descriptions of eriophyoid mites, and their phylogeny has hardly been studied. In the present study some aspects of eriophyoid systematics and its improvements by incorporating SEM for morphological study and phylogenetic analyses for testing and improving the naturalness of the present eriophyoid classification, are used and appraised. The morphology of about 64 species, mostly from South Africa, was studied with low-temperature (cryo) SEM. The specimens remained turgid and the shape of the mites largely unaltered. A general overview of the contribution of the SEM study towards systematic morphology of the Eriophyoidea is presented. Discrepancies between species descriptions from slide-mounted specimens and the SEM images were found. These include body form, interpretation of structures, resolution and information on minute morphology, and the presence of secretions. Some of these differences were caused by artefacts introduced with slide-mounting of specimens. The SEM study includes a comparative morphological study of the gnathosoma, including a review and appraisal of characters presently used in eriophyoid systematics. New morphological information was found, including new characters that may be of systematic use. Morphology studied with SEM should be routinely incorporated into eriophyoid descriptions, which is not presently the case. The phylogeny of the Eriophyoidea was studied at genus level, using morphological data, to test the monophyly of the present suprageneric taxa. Three data matrices with 66, 60 and 27 informative characters of 316 (including most Diptilomiopus spp.), 64 and 17 eriophyoid ingroup species respectively were analyzed with parsimony analyses, and trees were searched under different parameters. This was done to find different hypotheses regarding the taxon relationships, to roughly assess the robustness of the tree groups, and to use different approaches: a very comprehensive taxon sample, but with low ratio of characters to taxa; an exemplar species sample to improve the ratio between characters to taxa; and a very small taxon sample with a good ratio between characters and taxa, but very little inclusion of variation found in the Eriophyoidea. Most groups found were supported only by homoplasy, but many made biological sense and various potentially monophyletic groups, additional to taxa in the present classification, are proposed for further study. The robustness and convergence of these groups on monophyly are discussed. The Phytoptidae was found to be polyphyletic. Part of the Nalepellinae is probably positioned outside the remainder of the Eriophyoidea, while the rest of the Phytoptidae were positioned in smaller subgroups among the Eriophyidae. The Phytoptinae and Sierraphytoptinae, including Pentasetacus, may group together. The Eriophyidae never grouped together with much support, and the family is both polyphyletic and paraphyletic. The Diptilomiopidae was largely found to be monophyletic, with a relatively strong phylogenetic structure. The Rhyncaphytoptinae is mainly paraphyletic, and the Diptilomiopinae polyphyletic, but part of the Diptilomiopinae may be monophyletic. Three new Diptilomiopus spp. from South Africa are described as part of the study: D. faurius sp. nov. from Faurea rochetiana (A. Rich.) Pic. Serm. (Proteaceae); D. apobrevus sp. nov. and D. apolongus sp. nov. from Apodytes dimidiata E. Mey. ex Arn. (Icacinaceae). They were leaf vagrants not causing any observable symptoms.Thesis (PhD)--University of Pretoria, 2011.Zoology and Entomologyunrestricte

New pseudotagmic genus of acaricaline mites (Eriophyidae, Acaricalini) from a South African palm Hyphaene coriacea and remarks on lateral opisthosomal spines and morphology of deutogynes in Eriophyoidea

A new genus, Tumescoptella n. g., with two new sympatric species Tumescoptella aculeata n. sp.and T. rotundiscuta n. sp., was found on the indigenous South African palm Hyphaene coriacea, Lala palm. Both species are described and illustrated with the aid of conventional light microscopy, confocal laser scanning microscopy and low temperature scanning electron microscopy. Additionally, a new record of Tumescoptes dicrus Meyer collected from Phoenix reclinata(Arecaceae), near Kirstenbosch Gardens (Cape Town, South Africa), is reported. Morphologically the Tumescoptella n. g. is most similar to Tumescoptes Keifer, but possesses a more reduced chaetome and more derived pseudotagmosis. Pseudotagmosis is one type of body consolidation in vagrant eriophyoids, realized through the formation of dorsal opisthosomal plates, pseudotagmata. Along with previously described pseudotagmata (prodorsum, cervix, postprodorsum, superpostprodorsum, pretelosoma, and telosoma), a new term, anteroscutum, is proposed for the complex pseudotagma formed as a fusion of the three anterior pseudotagmata (prodorsum+cervix+postprodorsum) in several phyllocoptine mites from palms. An anteroscutum is present in monotypic Scolocenus Keifer from coconut and in the new genus Tumescoptella n. g. Protogyne females of Tumescoptella n. g. have uncommonly large lateral opisthosomal spines. Along with tiny spinules, triangular cuticular plates, band-shaped processes, and areas of thickened cuticle, the lateral spines belong to a common group of serial derivatives of dorsal opisthosomal annuli. We give a brief review on variation of spine-like structures across taxa of Eriophyoidea and their morpho-functional evaluation. Contrary to protogyne females, deutogyne females of T.aculeata n. sp. lack lateral spines, and they possess a smoother topography of the anteroscutum. Morphological similarity of these deutogynes with mites of the less transformed genus Tumescoptesis in accordance with a previous hypothesis on the possible correspondence of deutogyne morphology to that of an ancestral taxon. Two hypotheses on the origin of the presumably monophyletic group of Tumescoptes-like phyllocoptines (TP) from palms, comprising genera Scolocenus, Tumescoptella n. g., Tumescoptes, and Pseudotagmus, are proposed based on morphological similarity with two groups of eriophyoid genera: (a) pseudotagmic genera associated with aboriginal Asian dicotyledonous tropical trees, and (b) South American phyllocoptines from arecoid palms.Light microscopy investigation of the mite samples from Africa was supported by research projects of ZIN RAS (#АААА-А17-117030310209-7). CLSM studies were performed at the Research park of St. Petersburg State University in the Center for Molecular and Cell Technologies (IAS 1.53.917.2016, project #109-9164) and Center for Microscopy and Microanalysis (IAS 1.52.1424.2016, project #112-8186). LT-SEM studies were performed at the Laboratory for Microscopy and Microanalysis, University of Pretoria, South Africa.The field survey to collect eriophyoid mites on Hyphaene palm species was funded by the South Africa Department of Science and Technology grant to the Agricultural Research Council in South Africa.https://www.biotaxa.org/saaam2019Forestry and Agricultural Biotechnology Institute (FABI

Eriophyidae (Acari) as potential control agents of South African weeds, with descriptions of a new species of Tegonotus nalepa and of Paraphytoptus nalepa

M.Sc. (Zoology)This study centres around the findings on mites (Acari: Eriophyidae) (also referred to as "eriophyids") as potential biological control agents of weeds in South Africa. The use of mites in the biological control of weeds is briefly reviewed. Four species of the family Eriophyidae have been used against weeds in other countries. Seventeen eriophyid species, causing symptoms possibly of significance to biological weed control of 12 plant species regarded as weeds in South Africa, are discussed. Methods to rear and study eriophyids in the laboratory have not yet been fully developed and studying these practically invisible arthropods, very closely associated with their host plants, frequently presents difficulties. Reported methods, and those used for the present study, are reviewed and discussed. The eriophyid, Aceria lantanae (Cook), causing flower and leaf galls on Lantana camara L. could not be successfully established in a quarantine laboratory on potted plants propagated from cuttings of L. camara forms from South Africa. The results obtained during this attempt were inconclusive. The failure of establishment on local L. camara forms could be attributed to a variety of factors, of which rearing methods and host incompatibility are the most likely. Convolvulus arvensis is a troublesome agricultural weed and occurs in especially the Orange Free State and the eastern and southern parts of the Cape Province. Biological control may possibly contribute to curbing this weed in South Africa. The eriophyid, Aceria malherbae Nuzzaci, causing deformation and galling on C. arvensis is regarded as one of the most promising candidates for the biological control of this weed. It was imported and successfully established on potted plants in a quarantine laboratory on biotypes of the weed occurring in South Africa. The biology and host specificity of A malherbae are accounted for, by using reported information and data obtained during this study. The biology of A. malherbae broadly conforms to the biology of other eriophyid species. This species has a narrow host range, being restricted to species very closely related to C. arvensis. The symptoms caused by this eriophyid on C. arvensis are described, and seem especially harmful to the reproductive potential of the plant. Two eriophyid species from L. camara from Paraguay, Tegonotus stefneseri and Paraphytoptus magdalenae, are described as new species. An illustrated as well as a dichotomous key to species (Eriophyoidea) described from L. camara, including the two new species, are provided. A key, and a review of known Tegonotus species, incorporated in a compendium based on the original descriptions of 74 species, are included. Shevtchenkella Bagdasarian is considered to be a junior subjective synonym of Tegonotus Nalepa

Mackiella Keifer 1939

Genus Mackiella Keifer, 1939: 147 Diagnosis. Mackielline mites with broad, thin frontal lobe of prodorsal shield covering medio-proximal part of palpcoxal bases; tibial solenidion φ present, prodorsal shield setae ve and sc directed anteriad or lateroanteriad; dorsal opisthosomal annuli thrice wider than ventral annuli, flattened, with a distinct pattern of prominent longitudinal ridges; ventral annuli convex and rising high on the sides to the lateral edges of dorsal annuli. Type species. Mackiella phoenicis Keifer, 1939: 147 –148, 159 (Fig. XLV) Species included. M. phoenicis, Mackiella reclinata n. sp. Remarks. The mite species Mackiella borasis Mohanasundaram, 1981 is herein excluded from the genus Mackiella (see details under subfamily Phytoptinae below). Distribution and hosts. So far Mackiella mites have been recorded from North America (USA, Keifer 1939), South Africa (this paper) and Iraq (Mohamed & El-Haidari 1965) living on young leaves of Phoenix palms. Remarks. Newkirk and Keifer (1975, p. 568) wrote that M. phoenicis is “widespread on coconut” and “undoubtedly occurring wherever the host grows” without providing a reference to a collecting record. Amrine and Stasny (1994, p. 224) mistakenly mentioned that the original name of Mackiella type species is “ nuciferae Keifer 1939 ” and that M. phoenicis was collected from Cocos nuciferae. This was repeated by Amrine et al. (2003, p. 19) who stated that this mite species is “widespread on Cocos nuciferae ”. Actually, Mackiella mites have never been recorded from coconut palms.Published as part of Chetverikov, Philipp E., Craemer, Charnie, Vishnyakov, Andrey E. & Sukhareva, Sogdiana I., 2014, CLSM anatomy of internal genitalia of Mackiella reclinata n. sp. and systematic remarks on eriophyoid mites from the tribe Mackiellini Keifer, 1946 (Eriophyoidea: Phytoptidae), pp. 261-279 in Zootaxa 3860 (3) on page 266, DOI: 10.11646/zootaxa.3860.3.5, http://zenodo.org/record/22840

New phytoptids (Eriophyoidea, Phytoptidae) from southern African endemic dicotyledons: description of two new species of Solenocristus n. g. and updated key to world genera of the tribe Sierraphytoptini

A new phytoptid genus and two new species, Solenocristus karooensis n. g. & n. sp. and S. searsius n. sp. (Phytoptidae, Sierraphytoptinae, Sierraphytoptini), were collected in the Eastern and Western Cape provinces of South Africa, from southern African endemic dicotyledonous trees Schotia afra (Fabaceae) and Searsia lucida (Anacardiaceae) respectively. They are described and illustrated with the aid of conventional light microscopy and low temperature scanning electron microscopy. Females of both new species possess a distinct pregenital plate divided into two parts by a subcuticular medial ridge. Additionally, in S. karooensis n. sp. the movable digit of the male chelicerae varies in length: among six observed males, three males had a movable digit (md) twice shorter than the fixed digit (fd), in one male it was slightly shorter than fd and in the remaining two males md and fd were of the same length. An updated key to the world genera of the tribe Sierraphytoptini is given. The key incorporates data on seven sierraphytoptine genera including three genera (Neoprothrix Reis & Navia, Solenoplatilobus Chetverikov & Craemer and Solenocristus n. g.) which were described since the last generic key of Eriophyoidea by Amrine et al. (2003) was published.Partly supported by Russian Foundation for Basic research (16-04-01292) and research project of ZIN RAS (АААА-А17-117030310209-7) to the first author.https://www.biotaxa.org/saaam2019Forestry and Agricultural Biotechnology Institute (FABI

Integrative Taxonomy of the Gall Mite <i>Nothopoda todeica</i> n. sp. (Eriophyidae) from the Disjunct Afro-Australasian Fern <i>Todea barbara</i>: Morphology, Phylogeny, and Mitogenomics

Eriophyoidea is a group of phytoparasitic mites with poorly resolved phylogeny. Previous studies inferred Eriophyidae s.l. as the largest molecular clade of Eriophyoidea, and Nothopodinae as the basal divergence of Eriophyidae s.l. We investigate the morphology and molecular phylogeny of Nothopoda todeica n. sp. (Nothopodinae, Nothopodini), associated with a disjunct Afro-Australasian fern Todea barbara (Osmundaceae) from South Africa. Our analyses (1) determine new erroneous sequences (KF782375, KF782475, KF782586) wrongly assigned to Nothopodinae instead of Phyllocoptinae, (2) confirm the basal position of Nothopodinae in Eriophyoidea s.l., (3) question the monophyly of the Colopodacini and Nothopodini tribes, and (4) show the nested position of African fern-associated Nothopoda within a clade dominated by Asian nothopodines from angiosperms, which implies (a) a secondary association of nothopodines with ferns and (b) no relation between geography (continents) and the phylogenetic relationships of Nothopodinae species. Finally, we obtained a first complete mitochondrial genome for Nothopodinae and revealed a new gene order in the mitogenome of N. todeica n. sp., notably deviating from those in other investigated eriophyoids. Our results contribute to resolving the phylogeny of Eriophyoidea and provide an example of an integrative study of a new taxon belonging to an economically important group of acariform mites

CLSM anatomy of internal genitalia of Mackiella reclinata n. sp. and systematic remarks on eriophyoid mites from the tribe Mackiellini Keifer, 1946 (Eriophyoidea: Phytoptidae)

Chetverikov, Philipp E., Craemer, Charnie, Vishnyakov, Andrey E., Sukhareva, Sogdiana I. (2014): CLSM anatomy of internal genitalia of Mackiella reclinata n. sp. and systematic remarks on eriophyoid mites from the tribe Mackiellini Keifer, 1946 (Eriophyoidea: Phytoptidae). Zootaxa 3860 (3): 261-279, DOI: 10.11646/zootaxa.3860.3.

Atypically Shaped Setae in Gall Mites (Acariformes, Eriophyoidea) and Mitogenomics of the Genus <i>Leipothrix</i> Keifer (Eriophyidae)

The setae in Eriophyoidea are filiform, slightly bent and thickened near the base. Confocal microscopy indicates that their proximal and distal parts differ in light reflection and autofluorescence. Approximately 50 genera have atypically shaped setae: bifurcated, angled or swollen. These modifications are known in the basal part of prosomal setae u′, ft′, ft″, d, v, bv, ve, sc and caudal setae h2. We assessed the distribution of atypically shaped setae in Eriophyoidea and showed that they are scattered in different phylogenetic lineages. We hypothesized that the ancestral setae of eriophyoid mites were bifurcated before later simplifying into filiform setae. We also proposed that hypo-furcating setae are a synapomorphy that unites Eriophyoidea with Nematalycidae. We analyzed four new mitochondrial genomes of Leipothrix, the largest genus with bifurcated d, and showed that it is monophyletic and has a unique mitochondrial gene order with translocated trnK. We exclude Cereusacarus juniperensis n. comb. Xue and Yin, 2020 from Leipothrix and transfer five Epitrimerus spp. to Leipothrix: L. aegopodii (Liro 1941) n. comb., L. femoralis (Liro 1941) n. comb., L. geranii (Liro 1941) n. comb., L. ranunculi (Liro 1941) n. comb., and L. triquetra (Meyer 1990) n. comb

A New Webbing Aberoptus Species from South Africa Provides Insight in Silk Production in Gall Mites (Eriophyoidea)

Arthropods include a high diversity of lineages adapted for silk production. Several species of microscopic phytophagous mites of the hyperdiverse superfamily Eriophyoidea spin web; however, the origin of their silk is unknown. We described a new web-spinning mite, Aberoptus schotiae&nbsp;n. sp., collected from leaves of Schotia brachypetala (Fabaceae) in South Africa and showed that it has a complex life cycle, including two morphotypes of adults and nymphs. Molecular phylogenetic analyses and 28S sequence comparison showed conspecificity of heteromorphic females and rejected synonymy of Aberoptus and Aceria proposed by previous authors. For the first time, we provided SEM images of the web nests and, using a set of different microscopic techniques, described the silk-producing anal secretory apparatus (ASA) of Aberoptus. It comprises two pairs of anal glands (hypertrophied in web-spinning females), three cuticular sacs and a rectal tube leading to the anal opening. This is a unique case (analogy) of anal silk secretion in Chelicerata previously reported only in Serianus (Pseudoscorpiones). Recent findings of rudimentary ASA in distant eriophyoid lineages and the results of this study transform the current paradigm of exoticism of web-spinning eriophyoid taxa into the concept of synapomorphic specialization of the hindgut for excreting the anal gland secretions in Eriophyoidea

A New Webbing <i>Aberoptus</i> Species from South Africa Provides Insight in Silk Production in Gall Mites (Eriophyoidea)

Arthropods include a high diversity of lineages adapted for silk production. Several species of microscopic phytophagous mites of the hyperdiverse superfamily Eriophyoidea spin web; however, the origin of their silk is unknown. We described a new web-spinning mite, Aberoptus schotiae n. sp., collected from leaves of Schotia brachypetala (Fabaceae) in South Africa and showed that it has a complex life cycle, including two morphotypes of adults and nymphs. Molecular phylogenetic analyses and 28S sequence comparison showed conspecificity of heteromorphic females and rejected synonymy of Aberoptus and Aceria proposed by previous authors. For the first time, we provided SEM images of the web nests and, using a set of different microscopic techniques, described the silk-producing anal secretory apparatus (ASA) of Aberoptus. It comprises two pairs of anal glands (hypertrophied in web-spinning females), three cuticular sacs and a rectal tube leading to the anal opening. This is a unique case (analogy) of anal silk secretion in Chelicerata previously reported only in Serianus (Pseudoscorpiones). Recent findings of rudimentary ASA in distant eriophyoid lineages and the results of this study transform the current paradigm of exoticism of web-spinning eriophyoid taxa into the concept of synapomorphic specialization of the hindgut for excreting the anal gland secretions in Eriophyoidea