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Systematics and biology of termites and about their phoretic associations

They live in eusocial communities, but are not closer related to ants or bees. Termites belong to the cockroaches.

 

Queen, king and castes

 

Usually one queen and one king are reproductive and act as heads of the nest. The different work fields of a nest are executed by infertile specimens, which can show very different and specialized body shapes. The diversity of different castes is in phylogenetically „primitive“ taxa lower than in „higher developed“ termite groups.

 

As example specimens of a deadwood species from Italy

 

This species was found in deadwood of a small forest in Portici (Gulf of Naples, Italy) and might represent the taxon Kalotermitidae. This taxon branches off rather basically  in the systematic tree of termites. Nest work can be taken over by nymphs of later alates.

 

deadwood-termites from Italy, Youtube: copyrights Stefan F. Wirth, April 2020

 

 

How is wood-eating possible?

 

Wood eating termites bear bacteria and protozoans  in their digestive tracts, which perform the digestion of cellulose.

 

Evolution, sister taxon and endosymbionts

 

Termites (Isoptera) evolved within the cockroaches (Blattodea). According to modern systematics (e.g.  Beccaloni & Eccleton, 2011) the cockroach taxon Cryptocercidae is the sister-clade of the termites. But there are controversial theories existing.

According to such reconstructions, the last common ancestor of cockroach taxon Cryptocercidae and termites possessed bacterial and protozoan endosymbionts. Molecular data proved that endosymbionts in both groups are closely related to each other. The last common ancestor of both groups showed in case of their indeed sister-group-relation a tendency towards social communities. Cryptocercidae live temporarily in bigger groups together with their offspring.

 

Subsocial lifestyle in Cryptocercidae

 

Cockroaches of the Cryptocercidae as putative sister taxon of termites live inside galleries in deadwood and feed on wood fibres. At least one parent and its nymphs live subsocially inside their galleries. Cryptocercidae adults and nymphs groom each other, and parents feed juveniles with wood fragments afer these had passed their anus openings.

 

According to recent systematic/ phylogenetic reconstructions the Kalotermitidae belong to the basically branching termite groups. Such basic groups of termites still show a low diversity of castes only.

 

Associates, commensalism and phoresy

 

Like ants or bees, termites share their nests regularly with associates of other groups of animals,

often mites and nematodes. Some of these organisms use termites as carriers for a transport over bigger distances. details of such associations between insects and mites are not well studied yet. But carrier-passenger-situations with transfer („taxi“-) purposes are called phoresy. Phoresy ist mostly considered as a neutral association between different organisms and is thus interpreted as commensalism. Commensalism is differed from strategies like parasiticm or symbiosis and requires that two organisms in association do not harm or noticeably benefit each other. The term commensalism often includes associations, in which the true context for both organism partners is simply not understood yet.

 

Not yet mites of the Gamasina (Parasitiformes) were reared in greater numbers out of my Italian termite substrate. They might represent phoretic cohabitants of those termites. Other mite species of different mite groups (Parasitiformes and Acariformes) were only found in smaller numbers and died out too quickly for collections and determinations under my culture conditions, unfortunately already before the beginning of my shootings. seemingly microclimatic conditions had become too unfavorable.

 

Copyrights Stefan F. Wirth, Berlin 2019 – 2020, all rights reserved

Different wing colors in a harlequin ladybeetle specimen

The ladybeetle Harmonia axyridis is naturally distributed over eastern Asia, but was imported to the United States already at the beginning of the 20th century as pest control. At first, there was no population development in the open fields. These were at first reported from Louisiana in 1988. In 2001 the first free living specimens were for Europe discovered in Belgium. Since then the beetle distributed over several European countries, such as France, entire Germany or Switzerland.

 

Variations of Harmonia axyridis

 

The beetle is well known for its great form variations. Worldwide more than 200 different color pattern forms of thorax and elytrae are described. They are distinctly shaped and maintain in this shape and arrangement of pattern. But four forms dominate within natural populations. Speaking about the elytrae (not the thorax patterns), the reddish form with dark spots, as visible in my film, is one of them.

These distinct different forms must be named a polymorphism and are based on genetic information as well as on environmental conditions, such as temperature, humidity and light intensitivity. According to that even the term polyphenism might be adequate.

 

 

 

 

Transcription factor pannier responsible for color pattern polymorphism

 

According to the work of M. Gautier et al. (the genomic basis of color pattern polymorphism  in the harlequin ladybird, Current biology, 28, 20), the transcription factor pannier is responsible for the genetically based control of this polymorphism. They discovered that different pannier alleles determine the color pattern in the different known forms. The authors furthermore report that pannier was never found before to play a keyfactor role in the pigmentation of insects.

 

Ladybeetle species on a meadow in Berlin

 

The specimen in my footage was discovered on an urban meadow in the park area „Nordhafen“ in Berlin. It’s a meadow in autumn predominantly consisting of lucerne and clover, sorrel and yellow field cress. Different ladybeetle species could be in greater numbers found there between September and October 2019. The sevenspot-ladybird, the adonis ladybird (Hippodamia variegata) and most abundant the harlquin ladybird in all its developmental stages.

 

Asymmetrical wing colors and possible explications

 

 

Harlequin beetle specimen from Berlin with asymmetrically colored wings, copyrights Stefan F. Wirth, please like my video also on Youtube

 

The most conspicuous character of „my“ harlequin ladybird specimen was its distinct asymmetrically colored wings (elytrae). One side reddish with black spots, the other side brownish with black spots. During my research about such asymmetries in ladybirds, I didn’t find recent studies, which distinctly focussed on that topic. H. E. Roy et al. reported in their book „ladybirds“ (original version 1989, revised version 2013) about the existance of such differently colored wings in the same specimen. They emphasized that the phenominon was not studied in detail, but assumed different factors being eventually responsible for such a development of a beetle individual: 1) disruption of pigment production, 2) mitotic mutation in early development, 3) environmental conditions, eventually influencing the colors of an originally normal developed young adult (exposed for longer time to different light intensities etc.). The latter might in the case of „my“ specimen being an indeed possible factor, as it is clearly visible that also the brownish wing has at its edges some of the reddish pigments.

 

 

Filming/ photography conditions

 

The beetles was filmed and photographed under artificial conditions in a soil and grass-set in my video lab. There, mites of the Gamasina (Parasitiformes, evtl. mostly Laelapidae) were common. They interestingly showed a phoretic behavior by quickly climbing onto the wings of that ladybeetle. They obviously recognized it as a suitable carrier to new habitats. I assume ladybeetles in the field not being of much attraction for phoretic dispersal, based on their life-cycles and preferred habitats.

 

Berlin, September/ October 2019, Copyrights Stefan F. Wirth

Complex and modified mouthparts in Histiostomatidae mites

Mites represent arachnids, which means that they share characters with much bigger organisms, such as spiders, skorpions or harvestmen. Their bodies consist of specialized bundles of segments, named tagmata.  Two major tagmata are differed from each other in arachnids: prosoma, including legs and mouthparts, and opisthosoma, including for example the digestive and the reproductive systems.

 

Diphyletic origin of mites

 

Mites are not longer just mites. The former two clades of mites, Parasitiformes and Acariformes, originally considered as sister taxa, were in the modern systematics reconstructed to be diphyletic. That means, there was no commor ancestor, from which only those two clades derived.  There is no close relationship between them,each clade is most closely related to different groups of arachnids.  Thus, when I talk about mites, I am talking about the clade Acariformes.

 

Mites of the Acariformes and body plan

 

In these Acariformes mites, the arachnid body construction plan was modified into three visible tagmata: gnathosoma (bearing chelicerae and pedipalps as mouthparts), proterosoma (bearing first two leg pairs) and hysterosoma (bearing last two leg pairs and opisthosoma organs).

 

big male 2 Saarland compost

Male (large morph) of mite Histiostoma feroniarum in dorsal view. Body division in gnathosoma, proterosoma and hysterostoma. Fixation : critical-point-dried, SEM photography, copyrights Stefan F. Wirth

 

 

Mouthparts

 

Let’s talk about mouthparts, as they are an important aspect of my systematic and my function.morphological studies. Originally the gnathosoma consists of a pair of scissor-shaped chelicerae to grasp the food particles and of a pair of leg-shaped pedipalps, which mostly have mechano-sensitive and chemo-sensitive functions. But because mites colonized almost all kinds of existing habitats on earth, they extensively were exposed to the mechanisms of evolution. Acariform mites show a high range of variability regarding their morphology and their life strategies.

 

Mouthparts of Sarcoptiformes

 

Within the clade Sarcoptiformes, consisting of oribatid and astigmatid mites, there evolved a tendency towards miniaturization. Mites of the Astigmata are usually much smaller than one mm. Correspondingly the cuticle becam thinner and softer, perfect adaptations to a life inside very tiny micro habitats, but at the same time also a limitation, namely towards more or less moist habitats due to the lack of a well developed desiccation protection. They appear inside compost, rotting wood or mammal dung, being even there very specifically adapted into very defined micro climatic conditions. They live in a world of complete darkness, which is why light sensory organs are completely lost or reduced to vestigial structures.

Inside their habitats, astigmatid mites need to reproduce, to develop through different nymphal stages until adulthood and of course to feed. Astigmata are no fluid suckers, but feed on particles, such as bacteria, algae, fungi, thus many Astigmata taxa can be named microorganism feeders.

 

Life-strategy of mites of the (family) Histiostomatidae

 

Rollei Digital Camera

Extinct bark beetle fpssil in amber (collection Hoffeins) with phoretic mite deutonymphs. Fixation with hexamethyldisilazane, stereomicroscopic photography, copyrights Stefan F. Wirth

 

One of the largest family within the Astigmata clade is the Histiostomatidae, which I use since many years as model for my scientific studies. These mites are scientifically interesting from different points of view. Their ecology is characterized by life styles, which correspond to the life cycle of insects and other arthropods, to which most species have a close association. Most important aspect of these interactions between mites and other arthropods, commonly insects, is a dispersal strategy named „phoresy“. Mites use their „partners“ as carriers from one habitat to another. These habitats can often be the nests of the corresponding arthropods/ insects.

Habitats, in which mites of the Histiostomatidae develop successfully need to be moist and need to contain a sufficiant amount of microorganisms as food source. It is the most conspicuous feature of these mites to possess  remarkably modified mouthparts compared to the above described standard equipment of an acariform gnathosoma.

 

Mouthparts of the Histiostomatidae

 

 

Mite Histiostoma sp. (sapropel around ponds, female, Berlin) feeding from a substrate surface inside its original habitat. Videography in 4K, copyrights Stefan F. Wirth

 

The character conditions of the gnathosoma were one of the reasons, why I at the beginning of my phd thesis in 2000 decided to put my research focus on this mite family, being worldwide in major still unexplored.

The chelicera modified into a dagger-like structure being formed by the fixed part of the former scissor-like organ, named the digitus fixus. There is a variability of shapes of this digitus fius-chelicera-ending within the Histiostomatidae . It can appear „simple-dagger-like, simple formed with a hook-like ending or having cuticular dentations of specific numbers and sizes along the lower edge of the digitus fixus.

As typical for mites of the big clade Astigmata, the pedipalps are reduced in size and almost immovably ventrally and dorsally connected with each other. In Histiostomatidae, the third pedipalp article is additionally distinctly bent sidewards. Their front sides bear more or less complex arrangements of flexible membraneous structures, which can morphologically differ between taxa or even species, thus giving them a systematic relevance. I named these membrane-organs „palparmembrane“ following the nomenclature, introduced by R. Scheucher in 1957. These membranes can be devided into fringes or being lobe-sphaped and can cover the last pedipalp article dorsally and/or ventrally. My histological analysis from 2006 indicated that these membranes are shaped by the enditesof the pedipalpal coxae.

 

Complex mouthpart apparatus

 

Thus Histiostomatidae possess a bizarre mouthpart apparatus being unique within the Acariformes and representing an amount of characters, which from the phylogenetc point of view  can be reconstructed to have evolved in the stem species of that family (so called apomorphies).

 

Mouthpart apparatus as multifunctional organ

 

 

Mite Histiostoma sp. (male left, female right) feeding from a substrate surface inside its original habitat. Fixation with hexamethyldisilazane, SEM photography, copyrights Stefan F. Wirth

 

This gnathosoma is a multifunctional organ with the main function to select specific microorganism particles out of their liquid environments. When observing a histiostomatid mite with a sufficient high magnification walking along on a smooth water agar surface, on which bacteria and fungi growth was stimulated before, then occasionally trails can be seen around the walking mite, indicating that the gnathosoma was hold mostly leaned downwards towards the ground, pushing the microorganism cover along in front of the mite’s body. I interpreted this as an accumulation of food in order to gain more nutrients all at once. In my early papers, I described this as the typical feeding behavior of histiostomatid mites with the membraneous appendages acting like rubber sliders in the meantime. But as newer analyses showed is that such observations do not describe the full equipment of possible applications of the mite’s complex filter-feeding apparatus.

 

Membraneous structures create an underpressure to incorporate food

 

 

Mite Histiostoma ruehmi mouthpart endings with palparmembrane in ventral view. Fixation with hexamethyldisilazane, SEM photography, copyrights Stefan F. Wirth

 

More recent experiments with a higher videographic resolution and more suitable light conditions than 10 years ago (through-light and up light or one of them depending on the setting) showed that the palpar membrane structures , which more or less surround the entire fore-part (anterior part) of the gnathosoma can act like suckers: When the mite presses its front end of the mouthparts  to the underground, an underpressure can be formed based on these membraneous structures. This seemingly facilitates the incorporation of nutrients in that area. I presented such video footage in one of my former mite videos. To get off from the underground requires a jerky upward movement of the whole mouthpart apparatus (also visible in that older video). As I observed different developmental stages of different species, I could conclude that on a smooth surface with randomly dispersed food supply, regular stops and mouthpart-sucking-activities are seemingly a most common behavior of histiostomatids, while a straight forward walking behavior with the gnathosoma permanently touching the ground in order to push microorganism covers to the body`s front side only than occured, when food supply was uniformly dispersed (under artificial experimental conditions) under uniform moisture conditions.

 

Mite Histiostoma ruehmi and an undetermined species feeding from a smooth artificial substrate surface and performing an underpressure to incorporate food. Videography, copyrights Stefan F. Wirth

 

 

Scanning-electron-microscopic experiments

 

Mite Histiostoma feroniarum feeding from a substrate mount inside its original habitat. Fixation with hexamethyldisilazane, SEM photography, copyrights Stefan F. Wirth

 

Mite Bonomoia opuntiae feeding from the surface of a substrate mount inside its original habitat. Rounded particles might represent yeast bodies. Fixation with hexamethyldisilazane, SEM photography, copyrights Stefan F. Wirth

 

 

In my early postdoc-years, still at the FU Berlin, I performed experiments in order to fix mite activities inside their original substrates by filling such a mite-substrate-setting up with 1,1,1,3,3,3-hexamethyldisilazane and warming the corresponding small experimental dish, until the chemical was vaporized. I then sputtered the conserved setting with gold and studied the details on it via scanning-electron-microscopy. Occasionally, mites were shrinkled or deformed after this procedure, but sometimes they stayed in shape and did seemingly still remain in their last activity positions. I several times could take SEM photos, showing that (well visible only in adult mites due to their size) mite specimens can insert their (distal) chelicerae-endings into bigger heaps of substrate (obviously full of nutrients) and use the entire laterally bent pedipalpal articles, including the connected palparmembranes, to lean it against the substrate surface, either to stabilize the chelicerae movents or even to support the incorporation of nutrients again by forming a slight underpressure, or both.

 

Mite species Bonomoia opuntiae

 

Early observations during times of my phd-thesis on the mite Bonomoia opuntiae could show that the mouthpart apparatus of this terrestrial/semiaquatic mite works well also under water or inside a watery juce of decomposing cactus pieces. There even a filter function comparable with a fishing net was hypothesised, but so far was never studied in detail. The very distinct fringes along the palparmembrane lobes in this mite species might support this theory. I also studied the semiaquatic mite Sarraceniopus nipponensis feeding inside watery environments (normally the digestive fluids of Sarracenia pitchers), again never focussing in detail in how excactly the feeding mechanism works.

 

A putatively new species

 

The herewith presented video shows behaviors of  a female of the putative new species Histiostoma sp. , which I discovered in beginning of 2019 in sapropel around ponds inside an old gravel pit area in the Berlin forest Grunewald. The footage is presented in slow motion. The question was about how motile the whole gnathosoma apparatus in a histiostomatid species can be and what kinds of movements occured. As the settings, which I in early years of my mite studies used for videographic studies, were simplyfied and thus unnatural (smooth agar surfaces), I thought it being necessary and important to capture behaviors in a complexly sculptured habitat, namely surfaces of decomposing potato pieces (on which most histiostomatid species use to develop well).

 

It was visible, based on the specimens of my video of this species, that histiostomatid mites can be able to lift up their entire gnathosomas on a sometimes even higher position than the levels of the rest of their bodies. Additionally the gnathosoma can be turned to the right and to the left. Up and down as well as sideward movements of the whole feeding apparatus were often performed and represented obviously flexible reactions of the mite to the surface structure of the substrate and to the availability of suitable nutrients. In this context I was also interested in details of the movements of the chelicera tips themselves.

 

Chelicera endings (digitus fixus)

 

Although they can be used dagger-like and be accurately inserted into muddy substrate mounts, chelicera tips will also appear in a very fragile and seemingly careful way, when palpating the surface of the substrate underneath. Such chelicera movements are visible in the footage of this video, presented in slow motion (about 25 percent of original speed) and in a digital magnification. I interpret this visible fragility caution of the chelicerae as one option to discover suitable food sources. Other important organs perceive the mite’s environment chemically, modified setae, namely the so called solenidia, which might additionally recognize profitable microorganism sources.

 

Fig. 2

Mite Histiostoma feroniarum feeding from substrate mounts inside its original habitat (A-F). Rounded particles might represent yeast bodies. D = distal chelicera endings (digitus fixus), holding food particles, fixation with hexamethyldisilazane, SEM photography, copyrights Stefan F. Wirth

 

Berlin, September 2019

 

Copyrights Stefan F. Wirth

Phoretic Mites waiting on Ant Pupae

Greater numbers of pupae from a nest of the myrmecine ant Myrmica rudinodis are attached by phoretic mites, which wait for these pupae to hatch. They would then attach the newly developed ants to be carried around and dispersed this way. They this way had already occupied their later ants before, namely during their pupal stage, one could call this phenomen „pupa-guarding“. In my samples, I discovered two species of mites performing this pupa guarding behavior. Most abundant were deutonymphs of the mite Forcellinia wasmanni (Astigmata). But also individuals of a mite species of the Gamasina were repeatedly discovered sitting on pupae, where they were hiding between head, ventrum and limbs of the pupa. They even seemed to defend their pupae, when they felt disturbed, e.g. by my filming activities.

 

Ant pupa guarding by mitees, looking for a carrier for dispersal

 

These pupa guarding-findings concerning this ant and with these corresponding mite species might be new to science (so far I didn’t found literature indications) and thus need to be studied closer in the future in order to understand the whole context of behaviors. In the footage, two types of pupae are visible, pupae of the winged alates and those of workers. Mites generally prefered both, but especially the deutonymphs of Forcellinia wasmanni seemed to appear more often on the pupae of later workers. Most pupae had at least one deutonymph attached, rarely, there were found up to four individuals. This is different to what could be found on older workers. They on their ventral side can have 4-6 deutonymphs. Many workers seem to be covered with the deutonymphs, but I didn’t check more workers until now, so I can’t say, how many were without mites. It is unknown, how deutonymphs come to the pupae, whether they simply leave older workers for the pupa-guarding or whether they were waiting in the soil for the pupae to arrive (due to the brood caring activitoes of the ants).

Mite-Life inside an ant nest. Copyrights Stefan F. Wirth 2015/18

 

Astigmatid mite with a strict relationship to ants

 

The mite Forcellinia wasmanni is known to be strictly associated with ants (e.g. Türk & Türk 1957). It is clear that attaching young female alates would secure the dispersial of the mite into a new ant nest. It is not clear, which function the transport via ant workers can have. But Türk & Türk (1957) mention that the free living instars of Forcellinia wasmanni would feed on dead ants. Such a kind of microhabitat for the development is not unique in astigmatid mites. Some species within the Astigmata are known to have such preferences for decaying cadavers, but are then feeding on microorganisms, which grow on these (insect) cadavers. Ant workers might be ideal to carry mite deutonymphs to new cadavers, where they would leave and develop. Ants generally have a very well developed hygienic behavior. This guarantees the mites to get access to cadavers regularly. I do not know any other video footage, showing living deutonymphs attached to their carriers on such a magnification level as visible in this film. The original footage of these deutonymphs is much longer.

 

Morphology and behavior of the dislersal-instar, the so called „deutonymph“

 

The function of the proterosoma (dorsal shield of the forebody) is acting as a flexible structure, protecting the mouthpart-area (non-functional in deutonymphs) and the fore-legs, but being very motile and being easily pushed backwards (under the following hyterosoma-shield), when the mite lifts up from the surface of the ant pupa. I cannot state much more concerning the second mite, found on pupae, which is a species of the Gamasina. I discovered this phenomenon only on three of my pupae. Ant nests represent complex communities of organisms, to which fungae, other insects, mites and nematodes can belong. The samples visible in this film were collected in July 2015 on the German island Usedom inside a forest area between the villages Zinnowitz and Karlshagen. The ant nest was quite small. An ant hill was not visible.

 

Complexity of life in ant nests

 

The complexity of life within ant nests is a result of evolution. I am an enemy of creationistic movements, including all modern faces of creationism. Creationism stimulates carelessness und illiteracy in the believing people.

 

 

Berlin August 2015/ December 2018, copyrights Stefan F. Wirth

Habitat compost: Mite Histiostoma sachsi carries old cuticle and dirt as camouflage

My parents have a compost area in their backyards. I use it as reference habitat for two mite species of the family Histiostomatidae (Astigmata): Since I began my research in 2000, the compost regularly contained Histiostoma feroniarum with its typical male dimorphism. Since summer 2017 another species appears additionally regularly: Histiostoma sachsi. Both species do not appear under the same conditions. While H. feroniarum prefers fresher decaying material, H. sachsi on visibly older decomposed tissue. There mite be even more mites of the Histiostomatidae exist in this complex compost habitat, but under my laboratory conditions, only the two named species were so far successfully reared out of samles always again. Regarding the determination of H. sachsi on a species level, I was more careful in my comments to a former video (June 17), in which I named it Histiostoma cf. sachsi due to doubts about a correct identification. Meanwhile, also due to the morphology of the deutonymph, I determine „my“ compost mite as Histiostoma sachsi Scheucher, 1957. But it is still to emphasize that Scheucher described H. sachsi from cattle dung, not from compost. But generally, both habitats can sometimes share the same inhabitants.

 

Adult females carry their old cuticles and „dirt“ on their backs as tactile comouflage

 

Biologically conspicuous is darkish material, which especially adult females carry on their backs. Unlike males, females posses elongated setae on their backsides. These setae support the holding of material such as old cuticle and soil particles. In slide preparations, this cover usually appears amorphic and contains substrate from the mite’s environment. My video footage indicates that the basis of this cover is a retained old cuticle from the former nymphal instar . That this cannot easily be proven with the light microscope is due to the very soft and fine character of the cuticles in these mites. Remnants might become decomposed by microorganisms after a while.

Compost: the habitat of the mite Histiostoma sachsi Scheucher, 1957 (Acariformes, Astigmata, Histiostomatidae). Copyrights Stefan F. Wirth, please like my video also on youtube, in case you like it.

 

The phoretic dispersal instar, named deutonymph, in mites of the Astigmata controls its body position due to sticky leg endings and suckers on their undersides

 

Deutonymphs of H. sachsi represent one of my resent models to study mite-dispersal behavior. My research focus since a while concerns ultrastructure and function morphology of the deutonympal suckerplates and other structures to attach to insects for dispersal (this dispersal strategy is called phoresie). The anterior front-suckers on the suckerplate of the mite’s underside is extendable and very flexible, not only to find a suitable position on the insect carrier. When falling, the deutonymphs use it to lift their bodies up into a proper position again. Additionally they will try to get hold using „sticky“ lobe-shaped setae on the endings of legs I and II. Both is visible in my footage. The forelegs seem generally to make the first contact, when trying to get on a suitable carrier.

 

Deutonymphs of Histiostoma sachsi take a ride on other mites (Oribatida)

 

The suitable carrier of H. sachsi is unknown to me. Some astigmatid species have even a range of carrier-„hosts“. In my samples, deutonymphs at least attach to other mites, especially to mites of the Oribatida. This is in a very short scene visible in my video too.

 

Copyrights Stefan F. Wirth, Berlin December 2018

Isopoda: locomotion and aggregation behavior

Isopods represent Arthropoda. But they are no insects, no arachnids and no myriapods, although the glomerids or pill-millipedes can have a similar shape. Isopods are indeed crustaceans.  And the shape of pill-millipedes is a result of independent evolution (convergent evolution).

Isopoda represent a clade of the Malacostraca, whose members are originally native to aquatic habitats. Also isopods from the phylogenetic point of view represent aquatic/ marine organisms. But one clade, the Oniscidea, also named terrestrial isopods, evolved mechanisms to survive ashore.

The footage shows a species of Oniscidea of genus Porcellio from mediterranean habitats in Croatia. Land isopods retained their gills and thus usually prefer moist habitats. My model-isopods are reared in a terrarium, were they use to aggregate under moist pieces of bark.

As most terrestrial isopods, they are destruents and feed on decaying organic material. I add regularly pieces of fruits or vegetables and thus create conditions comparable with compost. Most of my footage is presented in slow motion.

I thank Jana Bedek, Croatian Biospeleological Society, for her determination of the isopods on a genus level.

Copyrights: Stefan F. Wirth, Berlin May 2017/ November 2018

Was sind sogenannte „Errata“?

Ziel der Forschungsarbeit eines Biologen ist stets die Publikation. Es ist dem Wissenschaftler dabei angeraten, in Journals zu publizieren, die ein ordentliches und hochwertiges Begutachtungsverfahren durchführen. Auf diese Weise ist sicher gestellt, dass die Arbeit auch tatsächlich zeitgemäßen wissenschaftlichen Standards entspricht.

Dennoch kann es auch in hochrangigen Journals geschehen, dass Fehler sowohl durch die Autoren wie auch durch die Gutachter übersehen werden. Selbst der Editor , der das Manuskript zuletzt auf die Eignung zur Publikation hin erneut überprüft, bemerkt nicht jede inhaltliche Unkorrektheit, jeden Druckfehler oder jede falsche Bildunterschrift.

Eine Publikation ist im Nachhinein jedoch nicht rückgängig zu machen. Auch Korrekturen können in der Regel nachträglich nicht mehr eingefügt werden. Sollten also Fehler nach Erscheinen der Publikation ins Auge springen, ist zu erwägen, ob der Leser diese leicht als Fehler erkennen kann oder ob ein sogenanntes Erratum (Lat. von errare = sich irren) notwendig ist, um Missverständnisse bei künftigen Lesern zu verhindern

Es handelt sich dabei um eine winzige eigenständige Publikation in derselben Zeitschrift, die in möglichst sehr knapper Form die Fehler benennt und korrigiert. Der Autor kann dann nur hoffen, dass Leser seiner Arbeit das zugehörige Erratum, das häufig erst in einer späteren Ausgabe des Journals erscheint, auch tatsächlich entdecken.

Korrekturwürdig ist beispielsweise auch, wenn ein Zoologe, der dabei ist, eine neue Tierart zu beschreiben, zu unvorsichtig ist, vor Erscheinen seiner Artbeschreibung Tiermaterial an Kollegen herauszugeben. In seltenen Fällen kann es so geschehen, dass der betreffende Kollege das Material in seine aktuelle Forschung mit einbezieht und seine Ergebnisse publiziert, noch bevor die eigentliche Artbeschreibung durch den Entdecker der Art auf den Weg gebracht wurde.

Im schlimmsten Falle nennt der Kollege in seiner vorgegriffenen Publikation einen Artnamen für das Tier. Er könnte somit unter Umständen zum offiziellen Erstbeschreiber werden. Im harmloseren Falle wird die Art als neu oder unbekannt (Gattung + n. sp. oder Gattung + sp.)angegeben. Jedoch ist es für spätere Leser beider Publikationen dann trotzdem eventuell nicht ersichtlich, dass die zuerst publizierten Forschungsergebnisse sich auf die erst danach publizierte neue Tierart beziehen.

In einem solchen Falle kann ein Erratum, um wissenschaftliche Zusammenhänge klarzustellen, angeraten sein.

Stefan F. Wirth

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Stefan Friedrich Wirth is a freelance German biologist, zoologist, evolutionary biologist and acarologist, living in Berlin.

– born in 1972 in the South-West of Germany.

– studies at the FU (Free University) Berlin 1994-2000

– phd thesis at the FU Berlin 2000-2004

– since 2004 research in the fields of systematics, evolution und ecology of mites (Histiostomatidae, Astigmata, Acari)  in cooperation with different international scientific institutions and  videografie/ macro-Videography as documentary contributions, for example to the „arte“-channel documentaries „Voyage sous nos pieds“ by the French director Vincent Amouroux.

selected publications:

Wirth, S. (2003): Das Stammartmuster der Histiostomatidae (Acari) und Beschreibung der durch zwei Männchen-Typen charakterisierten Histiostoma palustre n. sp.. Acarologia 42, 3: 257-270.

Wirth, S. (2004): Phylogeny, biology and character transformations of the Histiostomatidae (Acari, Astigmata). Promotionsarbeit. Internet Publikation, URL:http://www.diss.fu-berlin.de/2004/312.

Wirth, S. (2004): Phylogeny, Morphology and habitats of the Histiostomatidae (Astigmata). Proceedings of the V Symposium of the European Association of Acarologists. Phytophaga, XIV: 389-407.

Wirth, S. (2005): Description of a new species Bonomoia opuntiae (Histiostomatidae, Astigmata) with observations on the function of its eyes. Acarologia, vol. 45, no 4: 303-319. (URL:http://cat.inist.fr/?aModele=afficheN&cpsidt=18276055)

Wirth, S. (2005): Transformations of copulation structures and observations on the male polyphenism in the phylogeny of the Histiostomatidae. Internat. J. Acarol., Vol. 31, No. 2: 91-100.

Wirth, S. (2006): Development of the prelarva and larval behavior to open the eggshell in the Histiostomatidae (Astigmata). Abh. Ber. Naturkundemus. Görlitz 78,1: 93-104.

Wirth, S. (2006): Morphology and function of the gnathosoma in the Histiostomatidae (Astigmata). Acarologia,  vol. 46, no. 1-2: 103-109. (URL:http://cat.inist.fr/?aModele=afficheN&cpsidt=18695493)

Wirth, S. (2007): Phylogeny and characteristic transformations of the Histiostomatidae. In: J.B. Morales-Malacara, V. Behan-Pelletier, E. Ueckermann, T.M. Pérez, E.G. Estrada-Venegas and M. Badii (Eds.), Acarology XI. Proceedings of the XI International Congress of Acarology: 607-615, México.

Wirth, S. & Moser, J.C. (2008): Interactions of histiostomatid mites (Astigmata) and leafcutting ants. In: M. Bertrand, S. Kreiter, K.D. McCoy, A. Migeon, M. Navajas, M.-S. Tixier, L. Vial (Eds.), Integrative Acarology. Proceedings of the 6th Congress of the European Association of Acarologists: 378-384; EURAAC 2008, Montpellier, France.

Wirth S. (2009): Necromenic life style of Histiostoma polypori (Acari, Histiostomatidae). Experimental and applied acarology. DOI number: 10.1007/s10493-009-9295-6. URL:http://www.springerlink.com/openurl.asp?genre=article&id=doi:10.1007/s10493-009-9295-6

Wirth, S. (2010): Food competition and feeding behaviour and its implications for the phylogeny of the Histiostomatidae (Astigmata). – In: Sabelis, M. W. & Bruin, J. (eds.). Trends in Acarology: 37-40.

Wirth S. & Moser J. C. (2010): Histiostoma blomquisti N. SP. (Acari: Histiostomatidae) A phoretic mite of the Red Imported Fire Ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Acarologia 50(3): 357-371.

DunlopJ. A., Wirth1 S., Penney2 D.,  McNeilA., Bradley3R.S.,  Withers3 P. J.,Preziosi2 R. F. (2011): A tiny phoretic mite deutonymph in Baltic amber recovered by X-ray computed tomography. Biology letters doi:10.1098/rsbl.2011.0923.

Krüger J. & Wirth S. (2011): Life cycle of  Sarraceniopus nipponensis (Histiostomatidae: Astigmata) from the fluid-filled pitchers of Sarracenia alata (Sarraceniaceae). Acarologia 51(2): 259-267.

Koller L., Wirth S. and Raspotnig G. (2012): Geranial-rich oil gland secretions: a common phenomenon in the Histiostomatidae (Acari, Astigmata)? International journal of Acarology 38(5-38): 420-426.

Pernek M.1,2, Wirth S.3, Blomquist S. R.4, Avtzis D. N.5, Moser J. C.4 (2012): New associations of phoretic mites on Pityokteines curvidens (Coleoptera, Curculionidae, Scolytinae). Central European Journal of Biology. Volume 7, Issue 1: 63-68.

Pernek M. (1),  Novak Agbaba S.(1), Lackovic N. (1), Dod(1) N., Lukic I. (2), Wirth S. (3) (2012): The role of biotic factors on pine (Pinus spp.) decline in north dalmatia (croat: uloga biotičkih čimbenika u sušenjuborova (Pinus spp.) na područjusjeverne dalmacije). Šumarski list, 5–6, cxxxvi: 343–354.

Wirth S. (1), Pernek M. (2) (2012): First record of the mite Histiostoma ulmi in silver fir and indication of a possible phoretic dispersal by the longhorn beetle Acanthocinus reticulates. Šumarski list, 11–12, CXXXVI: 597–603.

Wirth S. & Garonna A. P.  (2015): Histiostoma ovalis (Histiostomatidae, Acari) associated with Ips sexdentatus (Scolytinae, Curculionidae, Coleoptera): ecology and mite redescription on the basis of formerly unknown adults and nymphs . International Journal of Acarology DOI: 10.1080/01647954.2015.1050062

WIRTH S., WEIS O., PERNEK M. (2016): A comparison of phoretic mites associated with bark beetles Ips typographus and Ips cembrae from Central Croatia. Šumarski list, in press.

WIRTH S. (2016): Description of developmental instars of Bonomoia sibirica n. sp. (Astigmata: Histiostomatidae) with ecological observations and phylogenetic conclusions. Acarina, December issue; in press.