biologe

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Tag: Histiostoma feroniarum

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.

Discussed diphyletic origin of mites

Mites are according to some acarological scientists eventually not longer just mites. The former two clades of mites, Parasitiformes and Acariformes, originally considered as sister taxa, were in some modern systematics reconstructed to be diphyletic. That would mean, there was no commor ancestor, from which only those two clades derived, the two major clades would be polyphyletic with no close relationship between them, each clade is assumed being closely related to different groups of arachnids (e.g. Psedoscorpions and Opiliones).  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 mites, Endeostigmata (seemingly paraphyletic) and astigmatid mites, there evolved a tendency towards miniaturization. Mites of the Astigmata are usually much smaller than one mm. Correspondingly the cuticle became 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.

Note from January 2020: In retrospect, I do not consider it sensible to superficially describe the feeding behavior using the palpar membrane at the edge. A precise videographic analysis of individual images exists and is currently being developed into a scientific paper.

Aspects of the histiostomatid feeding behavior, including using the membranous components at the anterior end of the mouthparts (pedipalps), can partly be seen in the video below.

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 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

Male and female of Histiostoma sachsi and unsuccessful mating with a „stranger“

Mites of the Acariformes vary in very different forms and life-strategies. One taxon of very tiny and soft-skinned mites is named Astigmata. Within them the familiy Histiostomatidae is especially rich of species, most of them surely not yet described or discovered.

 

Modyfied mouthparts and a specific mode of dispersal

 

These mites feed on microorganisms using a complex mouthpart-apparatus with multifunctional abilities. They can be found in habitats, which dry out quickly. When it’s getting too dry, a specific instar of the mites takes a ride on insects or other bigger arthropods for dispersal to a new and fresh habitat ( strategy called Phoresy).

Histiostoma sachsi is one of numerous (often closely related) long haired (in females) species. It was originally in 1957 described from cattle-dung. I found it in compost.

 

Long upper-setation in females and tactile camouflage (mimesis)

 

Adult females are characterized by a long setation on their uppersides. They use them to hold parts of the old nymphal cuticle and soil particles on their backs. This seems to be due to a strategy named mimesis or camouflage. It’s a tactile camouflage as an optical sense in this kind of microhabitats plays almost no role.

 

Normal and unusual copulation position, trial of an interspecific copulation

 

Males mate their females via a dorsal copulation opening and thus need to ride on them. In H. sachsi, that copulation opening is located very close to the hind-edge of the body. That way it is even despite of the camouflage cover accessible. It seems even slightly being elevated out of the body surface in order to surmount adjacent soil particles. This is an adaptation of this particular species. It might share such morphological characters only with very closely related (not yet described) species In other members of genus Histiostoma, the copulation opening is usually more centered related to the hind body.

The copulation position requires that males insert their aedeagus („penis“) into the copulation opening. They additionally use their legs to grasp into the females body. That kind of leg arrangement and thus the whole copulation position can differ from species to species.

This is why copulations between members of different species already fail, because the right copulation setting does not fit, nor does the shape of the aedeagous. Nevertheless the phenomenon of unsuccessful trials for interspecific copulations can sometimes be observed in laboratory cultures. Such a trial is also visible in this video, where a male of Histiostoma feroniarum (also appears in my compost samples regularly) tries to mate a female of H. sachsi. It cannot even almost get in a proper copulation position and seems to hold on to the dorsal camouflage cover of the female. it could only remain in a transverse position related to the female body and thus not get access to the copulation opening, normal would be a longitudinal position with the sameame orientation of female and male.

Adult mites of the family Histiostomatidae (Astigmata) and a „false“ copulation. Copyrights Stefan F. Wirth, Berlin December 2018. Please like my video also at Youtube, in case you like it.

 

Chemical communication and chemo-sensitive leg setation

 

Mites of the Astigmata communicate and find their general orientation due to chemo-sensitive setae, mostly on legs I and II, which are named solenidia. They are even on the magnification level of my footage well visible on the male’s legs. Although a direct body contact is not necessary for a innerspecific communication by chemically interpreting scents produced from mite glands, the observed male in my video repeatedly was seeking for intense body-contacts and obviously „observed“ his conspecific while doing so with its first two legs. This might have intensified the perception of pheromones.

It showed this behavior also, when passing by the „false copulation-pair“ described above. It additionally seemed to invest power in its leg movements as if it would try to remove the „competitor“ on the female, in this case even belonging to another species.

 

Competitive fights between males

 

That mites of the Histiostomatidae can use their strongly sclerotized first legs to fight under each other for an access to a female is known to me from my older observations about the species Histiostoma palustre and Histiostoma feroniarum.

 

Origin of the compost samples

 

The compost samples were collected in SW-Germany (Saarland in October 2018). The footage was recorded in December 2018 in Berlin.

 

Berlin 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