Mite Histiostoma palustre (Histiostomatidae, Astigmata), a mite species with two different male types

Current state of knowledge

Habitat and summary of general biology

I discovered the mite Histiostoma palustre in 2000 (and reared it at least until 2001) in the Berlin gravelpit area „Im Jagen 86“ at about 52°29’18.7″N 13°14’28.2″E and published the description in 2002 (2003). It is phoretically associated with at least two Hydrophilidae beetle species in bank mud (sapropel) around (in times of the sampling several) ponds. The phoretic juvenile instar deutonymph attaches its host insect to be carried that way to a new suitable habitat.

SEM photograph of ventral side of the deutonymph of Histiostoma palustre. © Stefan F. Wirth

SEM photograph of ventral side of Histiostoma palustre deutonymph as negative version, emphasizing single sicking structures and shapes of hind legs, © Stefan F Wirth

Phoretic hosts and attachment site

I in those times determined two host beetles as belonging to the genera Coelostoma and Cercyon. In laboratory host choice tests, in which deutonymphs in a culture had the choice between three different potential host species, there was a significant preference for those Hydrophilidae beetles, while another offered beetle species sympatrically living the same habitat was almost not at all attractive for the phoresy instar. But the choice test was performed as a smaller side project only and offered not more than a host tendency, as for example not a representative variety of insect species from the same natural field habitat of the mite was involved in these choice tests. But indications were at least clear enough that a host specificy (on a higher taxonomic level) of H. palustre can be stated. Mites preferred according to these tests a specific area for attachment on their hosts, which was the dorsal head area.

Two different male types and the difference between polymorphism und polyphenism

The mite showed under laboratory conditions a male diphenism with highly modified gripping organs or normal legs2. While the modified leg can appear on both sides or only on the left or the right side.

„Polymorphism“ is used internationally as the overriding term for such a phenomenon. Here, following the teaching of my former research group at the FU Berlin, I distinguished between a purely genetically determined expression of distinctly deviating morphs and the determination of these morphs by the presence of certain environmental factors. In the latter case we are talking about polyphenism. Of course, genetics also play a certain role here additionally.

Studies with cultures grown at different temperatures suggested that males with modified legs 2 appeared significantly more frequently at higher temperatures than at lower temperatures.

Temperatures: 30-32 °C (measured in summer inside the culture dishes at room temperature) and 20°C (climate cabinet, temperature inside and outside the cultures almost equal). Experimental approaches and total numbers of males counted: 10 cultures with lower (don’t remember exactly) numbers of developing mite stages per each of the two different temperatures. n=237 males in total at low temperature, n=729 males in total at high temperature. The number of modified males grew significantly from 18% (20°C) to 45% (30°C), while normal males decreased significantly from 82% to 55%.

Therefore I called the mechanism for the formation of two morphs diphenism. These studies were also a besides project and the amount of data would need to be even bigger according to higher level statistical requirements.

SEM photos of a male morph with modified legs 2, © Stefan F Wirth

Detailed SEM view to a male with legs 2 on both sides modified into gripping organs, perspective almost from the front side. © Stefan F Wirth

About the function of the modified male legs

I could observe in the mite cultures in which H. palustre thrived very well on decomposing potato pieces at about 20°C room temperature that the modified legs of the heteromorphic males used as prehensile organs played a role in intraspecific competition for female mating partners. Here several males, with and without gripping organs, were usually wedged together in veritable clusters around female tritonymphs or adult females. This was because the fighting happened more or less in a slow motion speed with apparent longer lulls in between. During the fights, the legs of the opponents were seized with the prehensile organ and the competitor was slowly pulled away from the coveted female.

Less is known about which selective pressure would favor the morph with gripping organ

Males with normally developed legs were at least seemingly in my observations clearly at a disadvantage here, since their second legs in particular could be easily grasped and they were hardly capable of defense and counterattacks. Nevertheless, random counts at that time did not provide any clear indication that heteromorphic males were possibly found significantly more frequently in the final mating position. The same applies to expectations regarding an unequal sex ratio in favor of males. Instead, males and females appeared to be almost evenly distributed in random evaluations. Additionally males with modified legs on both sides and with only one modified leg on the right and the left were in randomly counted cultures seemingly equally distributed.

Still many questions unanswered about the biology of H. palustre

It was unfortunately not tested at all, whether females mated by heteromorphic males and those mated by homomorphic ones had different reproduction successes. In connection with this biologically highly interesting mite species, many questions about its life strategies could not be examined deep enough during times of my PhD thesis. There was no time for this, since my research at the time was geared towards systematic comparative studies of numerous species. Unfortunately, the species could so far not be found again in its habitat „Im Jagen 86“ – at least not by the means of substrate samplings with only accidental Hydrophilidae inside – until today so that further studies were not yet carried out. This seeming lack of the species‚ presence in its former habitat is probably due to ecological changes in this area, which originally had several ponds, from which now only one remained.

A normal leg 2 of H. palustre compared with a modified leg indicating homolog leg articles and corresponding setation.

What makes this male diphenism interesting from the evolutionary point of view

Two distinct morphs within the same sex of a species is the result of evolution. It does at first not matter, whether it is a dimorphism or a diphenism. A fundamental question, which cannot be answered at this time, is: does the trait of these two morphs indicate that the species is on the way to finally having exclusively only males with prehensile organs, and is the species even on the verge of becoming the to cross the species boundary into a new species? Or is the characteristic of two male types a permanently stabled character, because exactly this has strong selection advantage?

In this context, fossil finds of closely related species (in amber) would be of interest, but unfortunately these are not yet known. It would also be interesting to compare different populations of this species, which are also not yet available. It furthermore would be of interest to understand, what exactly the advantage of modified males versus non modified males might be. Is it possible that an advantage of the modified-legged males is simply that they copulate more frequently, although smaller datasets have not yet confirmed this? And/or do females mated by modified males produce significantly more offspring?

And in the special context of the knowledge at that time that it is a temperature-dependent diphenism, one could raise the question of whether the species „evolutionally prepared itself“ by developing two male forms to cope better with temperature fluctuations. In view of increasing global warming, this may be of general biological interest.

A similar species from Egypt

A publication about a species with modified legs 2 as in H. palustre

In a conference proceedings contribution Bishlawy, S. E. M. O., and S. F. M. Allam published an article containing the description of a species of Histiostomatidae, which the authors have named Histiostoma egypti ( Proc.2nd Inter.Conf.Ent.Soc.Egypt, vol. 1, pp. 407-420, 2007). This is worth of being mentioned here more detailed in context of a possible close relationship of these species or a convergent evolution. But due to the lack of systematic facts, a decision hypothesis is not yet made. I can just emphasize that one showed me photos of that species on a conference in Cairo (Egypt) in 2003, and my first and superficial interpretation was that it even is Histiostoma palustre.

I have not checked whether the species Histiostoma egypti has been accepted internationally as a new species and is therefore also listed in the Zoological Record, as this is not relevant to the considerations here. It is interesting that a similar species was discovered in Egypt, about which there are some biological observations.

However, I only have a text version without illustrations, which is online published like that and which is why I cannot assess this species systematically/taxonomically on a deeper level. The authors refer to a similarity of their species with Histiostoma palustre including a similar diphenism of the males. I, as I already mentioned above, remember confirming this resemblance at an acarological conference in Cairo in 2003, where the co-author showed me photos of the later H. egypti.

Association with laboratory cultures of entomopathogenic nematodes

According to the authors, the species entered the laboratory with entomopathogenic nematode substrate and is doing well in the nematode cultures. The mite species is called semiaquatic, which quickly dies off at lower humidity. According to the authors, the thriving of the mites is to the detriment of the nematodes, which can die as a result. In this context, the authors mention having observed that the heteromorphic male form appears more frequently when live nematodes are present, while the homomorphic form and also the phoretic deutonymphs predominate when previously present nematodes have died.

My general experiences with histiostomatid mites and free living (phoretic) nematodes

A connection between the development of different mite morphs and the presence or absence of living nematodes never became particularly obvious to me in my studies at the time on species with different male types. Therefore, I have never specifically investigated such a connection and can therefore judge the correctness of such a connection as neither improbable nor very probable.

I can confirm, however, that basically all Histiostomatidae species that I examined always appeared together with nematodes, which were usually free-living, microorganism-eating representatives of the Rhabditidae or Diplogastrinae, which are also phoretic and possibly even were spread by the same hosts as the mites.

In this context, I am aware of the phenomenon that nematodes and mites can either compete with each other for food or reduce each other, for example chemically. In any case, it can often be observed that after some time in a culture vessel in which both animal groups were initially numerous, only the mites or the nematodes thrived in large numbers.

General information about the feeding behavior in Histiostomatidae

According to my recent research, based on frame-by-frame videographic analysis and SEM studies, Histiostomatidae species use their filter-feeding mouthparts to feed on decomposing fungal material that is also bacteria-rich. It is highly probable that the fungi are brought by the mites themselves into their habitat, which seems to happen hyperphoretically via fungal spores. The mites probably also control the growth and partial death of the fungus themselves in the form of their fungicidal gland secretions. Specific studies about the feeding behavior of H. palustre do not exist on a more advanced level. Slides and numerous SEM shots might indicate the transport of spores, but I so far had no time to examine those specimen closer. Such results thus need to be presented at another occasion.

Information about nomenclature used in the 2003 publication and about the citation of this species description

Nomenclature for dorsal/ventral setation of juveniles and adults used in this old species description of mite H. palustre was my own invention and is no longer used by me for practical/systematic reasons of comparison.

The nomenclature I introduced at the time of that species description for the pattern of individual fields that cover the thickening of the cuticle (proterosoma shield) on the anterior dorsal side of the proterosoma is still retained by me to this day. I still consider this characteristic in adults to be systematically relevant.

Cuticular shield of H. palustre stabilizes muscle origins at the dorsal proterosoma, close to the mouthparts, © Stefan F. Wirth

The original species description is cited as a 2001 publication on the Acarologia abstract page and is cited as 2002 publication in the PDF of the article provided by Acarologia as open access publication. In fact it was accepted for publication in 2001 and finally published in 2003. Thus I sometimes cited it as 2003 publication, but 2002 would be a correcter citation too. At least the full species name must be Histiostoma palustre Wirth, 2002.

Link to the original species description:

https://www1.montpellier.inra.fr/CBGP/acarologia/article.php?id=1849

© Stefan F. Wirth 2023, Berlin

Copulation details of snail Cornu aspersum (4K)

I collected two specimens of the land snail Cornu aspersum from an old olive grove in the city Sorrento (Gulf of Naples, Italy) in April 2019. The land molluscs could be found in that spring season in and under flat stones and smaller rocks. They shared this habitat with bgger diplopodes of genus Julidae and the harvestman Trogulus tingiformis.

The snails are until today successfully kept in a terrarium with sufficient moisture and regular food consisting of vegetable pieces. They share the terrarium with some specimens of diplopods from the original location in Italy.

Cornu aspersum mating, film made in Berlin, all copyrights Stefan F. Wirth

The snails go in a strict diapause several times a year for weeks or months, when temperatures in Berlin grow over 20°C . After getting active again on a colder period, they often quite immediately begin to mate. My video only focuses on details of such mating procedures, especially the spermatheca transfer. I decided to cut as less as possible and to present longer sequences of copulation activities, as they have due to slow slime exchange movements and rhythmical motions a special aesthetics. I additionally intended to show that scientific behavioral studies generally require patience and time as well as interest and fascination for aspects of life.

Cornu aspersum mates reciprocally, which means that both partners transfer a sperm package and produce ovules. The mating of my footage was successful (not visible in my video), and about 20-30 young snails hatched after about 2.5 weeks after egg deposition.

Aspects of mating and copulation in Cornu aspersum are quite well scientifically studied. For example: the variation in spermathecal morphology does not depend on the sperm competition intensity in populations (E. Koemtzopoulos & A. Staikou (Zoology 110 (2), 139-146, 2007); aspects of courtship and copulation were studied by S. A. Adamo and R. Chase (Canadian Journal of Zoology 66(6): 1446-1453, 2011). According to the latter authors the typical mating behavior has a duration of about 421.8 minutes and consists of three major parts: introductory behavior, dart shooting and copulation. My video concentrates only on part 3.

Berlin August 2019 – 10 June 2020, copyrights Stefan F. Wirth

Ant cricket and beetle Amphotis marginata in a nest of Lasius fuliginosus

The ant Lasius fuliginosus builts its nests into wooden environments, for example tree stumps. In the depth it is shaped by a carton-like substance, produced by the ants and with a „domesticated“ fungus involved. When ant workers leave the nests on trails, marked with pheromones, they might seek for food (mostly aphid secretions) in distances up to 30 meters. In the area around the nest, so called foraging trails are especially busy, as different kinds of foraging substances need to be carried in, in order to feed the fungus, in order to create new cartonage and in order to feed queen, nest mates and offspring.

Such a foraging trail is of course a very attractive place for invaders (non ant species) to either capture some food from the workers on their ways into the nest, or even to attach to these workers to get a ride inside the nest too, interesting for brood parasites for example, but also for all kinds of organisms, which prefer nest micro climatic conditions and want to be additionally secured or at least tolerated by the ants. All these organisms, such as insects, mites or nematodes, even pseudoscorpions, need to have specific adaptations in order to be not attacked by the ants.

Film about nest cohabitants of Lasius fuliginosus, Berlin 2020, copyrights Stefan F. Wirth

Three examples are presented in my video. The ant cricket Myrmecophilus acervorum is a common inhabitant of different ant species. Here I found it while „walking in row and order with the ants“. That unusual tiny cricket is assumed to be able to adopt the „smell“ of a nest, which is why ant workers accept it around them. I discovered the specimen of my footage in a later afternoon (around 18:00 in May 2020) directly on top of the tree stump, in which the nest is hidden (in the depth). There it directly followed ants within their foraging walk to the nest entrances. It was directly walking with them in a row and seemed to imitate additionally antennae movements of ants. It after a while left the row of ants (unharmed and without getting a special attention) and went into a hideaway on the side of the tree stump. Generally, there is not much known about the biology of this cricket. There is evidence that it feeds on food and even brood of the ants.

Another ant trail invader is the tiny beetle Amphiotis marginata (Nitidulidae), which performs behaviors, which make its stay inside foraging trails of ants (seemingly associated with Lasius fuliginosus only) even necessary: Hölldobler & Kwapich (2017) had studied this beetle and its behaviors in detail. According to their findings, the beetle shows a complex behavior to beg for food from passing-by antworkers. Movements of its antennae are an important part of such a contact and might in the optimal case lead to a response by the ant to antennate back to the beetle’s head, and subsequently the beetle might be fed as if it were an ant conspecific. The authors describe that a beetle is not always successful. In the best case, hectic ants on their way home might simply oversee the invader (kleptoparasite), in the worst case, they might detect it as a stranger and would then attack it. For protection, the beetle is able to closely adhere to the ground with its claws, while the side edges of its elytrae are shaped downward to the ground. This way, ants are unable to lift such a beetle up and would continue their ways after a while. Hölldobler and Kwapich also mention that they observed cases, in which ants were nevertheless able to lift detected beetles up and then cut their legs off, which means the end of the beetles adventurous life. The beetle specimen in my footage found a bad position aside to an ant path, which was such busy that it was overseen and even unable to approach single workers to beg for food. The authors above found some indications that the beetle’s larvae might develop inside ant nests.

As an acarologist, I am of course interested in mites, which are associated with ant nests. I in detail was involved in research about non-native ants: in the USA (Lousiana) I did research about the leafcutter ant Atta texana and the red imported fire ant Solenopsis invicta, all in cooperation with John C. Moser. I even described a new species of astigmatid mites from S. invicta. I also did some unpublished research on native ants and thus know that also Lasius fuliginosus possesses greater numbers of mite-associates (Parasitiformes and Acariformes). As an example given in this video, we see a rather big mite of the Mesostigmata (Parasitiformes), which I could not determine closer based on my footage. Mesostigmata generally can appear as phoretic organisms (feeding for example on nematodes or mites inside the ant nests, but being carried by ant workers there), they can also invade by themselves and might appear as brood or kleptoparasites. The mite in my footage walked directly on the ant trail without being harmed. It might be like the ant cricket able to adopt ant nest scents to be protected.

Berlin, Plötzensee/ Rehberge, May 2020, copyrights Stefan F. Wirth

Wild bee Andrena flavipes and nesting behaviors

The bee Andrena flavipes is also known as the common sand bee, as this species represents the most common of several regularly present sand bee species in Central Europe.

 

Aggregations at suitable nesting sites

 

Bee females create solitary nests, which is unlike to social hymenopterans such as the honey bee Apis mellifera. However huge and from a distance well visible aggregations of nesting A. flavipes specimens can appear. It is said that these aggregations are due to mated females being attracted to similar suitable nesting sites. In fact also a tolerance for conspecifics very close by is required to allow conditions, in which the whole ground seems to consist of bees, flying around and preparing their nests or importing pollen or nectar to feed their larvae. By the way: One nest contains contains about 2-3 brood cells only.

 

Specific conditions, in which specimens of my footage were found

 

The bees of my video were filmed between 4-6 April 2020 in the urban park around lake Plötzensee in Berlin. The site for my recordings was an area with forest edge character, interrupted by dry meadows, all at least in the afternoon exposed by the sun (temperatures between 15-20 °C).

 

 

Females of Andrena flavipes cleaning their nests, youtube: copyrights Stefan F. Wirth, April 2020

 

 

Orientation and nest cleaning behaviors of A. flavipes females, hindlegs as multifunctional organs

 

Contents of my behavioral documentation is the cleaning of nest hole entrance areas and behavior patterns, which seemingly support the orientation and finding their own nests again in midst of a sandy forest ground covered by fallen leaves.

To be enabled to recognize the entrance of the own nest again, bees perform regularly smaller walking tours around their nests to memorize soil structure and other details, being suitable to characterize this specific nesting site.

The bee’s hindlegs represent important multi-functional organs. They walk on them, collect pollen, which adhere to specific structures on legs III, and they are used to clean the areas in front of the nest openings from dirt, such as smaller stones or wooden particles. As nest entrance areas stay opened during the day, a proper cleaning of the soil around is regularly necessary. The bee performs that work mostly while backward-walking using its hindlegs like shovels to sweep dirt some centimeters away. This behavior is well visible in my footage.

 

General and short  information about mite associations

 

Andrena flavipes and other soil breeding wild bees are generally also of acarological interest. As presumably all hymenopterans, they have for example associations with phoretic mites, for example mites of the Scutacaridae (such as Imparipes apicola). I so far did not study mites on these bees, but phoresy means that mite instars use the insect as carrier to reach their final sites, in which they develop and reproduce. In case of Imparipes, adults feed on fungi and waste inside the bees brood cells.

 

 

Copyrights Stefan F. Wirth, Berlin april 2020, as always: all rights reserved

Diving, feather cleaning and water bathing of the Inca tern Larosterna inca

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Inca terns live along the South American Pacific coast and breed along rocky coastlines of Peru and North Chile. They can be easily identified by their grey plumages, their distinctly red beaks and feeds as well by their conspicious white feather curls on the bases of their beaks.

 

Geographic distribution and life-strategy aspects of Larosterna inca

 

Larosterna inca breeds inside rocky walls of coastlines either inside rocky cavities or in old nesting holes of other seabird species. Its hunting ground is the  Humboldt Current, which is famous for its cold temperature, but also its enormous fish wealth. To increase its chances for fishing success, the Inca tern might follow sea lions, cormorants and whales  and is then hunting fishes, which were flushed up by these bigger sea animals. They also follow fisher boats to catch some rests of their fishery.

#Inca #tern #Larosterna #inca #pictus in #freeflying #enclosure in #summer at #Berlin #zoologischergarten #July #2019 Copyrights #Stefan #F. #Wirth #light #bird #Inkaseeschwalbe #birdportrait #photography #Telelens #Laridae #close-up #sandy #photographer #photographylovers pic.twitter.com/0Ye1WYxi9b

— Stefan F. Wirth (@wirthstef) July 20, 2019

 

Phylogenetic (systematic) relationships

 

According to phylogenetic reconstructions L. inca, which represents the only recent species of its genus, branches off in the Animalia tree within the monophyletic clade of terns. Based on DNA sequences E. S. Bridge, A. W. Jones and A. J. Baker reconstructed in their 2005 paper (Molecular phylogenetics and evolution) a sister-clade relationship between Larosterna and species of the taxa Sterna, Thalasseus and Chlidonias (mitochondrial DNA was used to reconstruct the tern phylogeny).

Terns themselves seem representing an own clade (Sternidae), being for example based on characters of behavioral pattern, and are considered as a sister taxon of gulls (Laridae).

 

Filming conditions and filming locality

 

My footage was recorded in the Zoo Berlin, where terns together with other sea birds inhabit a for tourists accessible free-flight enclosure. There I captured scenes about the diving and „fishing“ behavior (specimens fished repeatedly wooden sticks) as well as their plumage cleaning activities on shore and their conspicuous plumage cleaning behaviors via extended bathing trips inside areas of low water. Size of my entire video is 4K. But parts of the scenes were originally recorded in Full HD to enable a better slow motion effect based on 100 frames per second. Such footage was subsequently digitally magnified into the 4K size to fit in the entire video project.

All behavioral activities are at first presented in a slow motion (ca. three to four times slowlier than  original speed), then in the much faster original speed.

 

Plumage cleaning

 

Plumage cleaning is part of the hygienic behaviors of birds. Feathers can only stay in full function, thermoregulation and flying, when dirt and parasites are removed regularly. Typical plumage parasites are represented by feather mites (no phylogenetic clade), which consist of taxa of the Astigmata (Acariformes) and of taxa of the Dermanyssoidea (Parasitiformes). Feather lice represent  a subclade of the (Phthiraptera = lice), named Mallophaga. The monophyletic situation of Mallophaga is seemingly doubtful.

 

Plumage cleaning and hunting behavior of Larosterna inca, video (4K9, copyrights Stefan F. Wirth. Please like my video on youtube too.

 

Putative reasons for plumage cleaning behaviors

 

I couldn’t research sufficient information about specific plumage parasites of Larosterna inca. There is indication that terns generally are relatively free of predators and parasites. Seemingly, plumage parasites of this particular species are still a more or less open research field. But the existence of a regular and visibly careful plumage cleaning might indicate a sensitiveness for corresponding parasites. L. inca can be according to literature (e.g. W. Pieters et al., Avian Diseases, 2014) fatally infested with the trematode Ichthyocotylurus erraticus.

 

Copyrights Stefan F. Wirth, Zoo Berlin July/ September 2019

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).

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

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 Histiostoma cf feroniarum feeding in its original substrate, fixed with hexamethydisilazane, SEM

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.

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

Oribatida mites: Fast runners and slow crawlers

Microhabitats often consist of a complexity of organism species. Under suitable conditions, samples can be kept „alive“ for months and even for years by regularly adding moisture and organic tissue, in case of my sample of this footage: patato pieces.

 

 

Mites of the Oribatida and their different ways of locomotion. Copyrights: Stefan F. Wirth, Berlin April 2019. Please give the video a like on youtube too.

 

Soil samples from island Norderney

 

This soil sample was collected in summer 2018 on the North Sea island Usedom during my participation at the „Geo Tag der Natur“. It contained several specimens of the predatory chilopode Lithobius sp. and pieces of rotting wood, moss and forestground, everything collected under rotting treetrunks and tree branches. The samples additionally contained the carabid beetle Pterosticus cf. niger and ants of genus Lasius. Samples were collected in a small forest area with wetland aspects. The soil quality was rather moist.

 

Astigmatid mites

 

I later added potato pieces and regularly some water droplets to the sample with still living big arthropods/ insects. After some weeks, specimens of the astigmatid mite Acodyledon cf. schmitzi developed on dryer areas of the potato pieces. These mites were presumably phoretic associates of the carabid beetles. They died out after several months, after the sample had dried out a little bit and may be due to changes of the room temperature during winter time.

 

Oribatida

 

Now, almost a year later, the micro habitat is inhabited by mites of the Oribatida in greater numbers of specimens of at least three species: Nothrus sp. (genus not yet clarified), Nothrus palustris (already found for the first time shortly after the sample collection) and a species of Phthiracarida.

 

Locomotion and biodiversity

 

Purpose of the short film is to show different organisms, cultured after about a year in this sample: mites, nematodes, collembolans and microorganisms, fungae and bacteria. Of the bigger arthropods/insects, only one Lithobius species survived until now.  Also the diversity of ways of locomotion in different oribatid species is emphasized: There are slow crawlers (Nothrus) and fast runners (Phthiracarida).

 

Berlin, April 2019, Copyrights Stefan F. Wirth

Eudicella colmanti – Mating behavior of a colorful beetle

Rose chafers represent a group of colorful beetles, which taxonomically belong to the Scarabaeidae and thus are relatives of famous beetles such as Scarabaeus sacer, well known for rolling dung into balls and for being an important symbol for creation and the rising sun in the ancient Egyptian world. Even the stag beetles are more distant relatives of rose chafers.

 

Colorful and active during daytime

 

Unlike some related beetle clades, rose chafers are usually active during the day. This is also indicated by their very colorful bodies. Colors in insects can have different functions, but they usually all are optical signals, which require a visibility in the sun light. Greenish colors are common in rose chafer species and might have optical inner specific signal functions, but also might support an optical camouflage. This would also make sense in the preferred habitats of the adult beetles, which usually feed on softer parts of blossoms and on their pollen. But they also feed on fruits, whereby mostly liquids are incorporated as the chewing mouthparts are not very well developed.

 

Tropical rose chafer Eudicella colmanti during its copulation behavior, 4K videography, copyrights Stefan F. Wirth.

 

Tropical rose chafers from African countries

 

About 3000 species of rose chafers are known, of which most inhabit the tropical zones. The about 20 species of the genus Eudicella are more or less restricted to the African continent.

Eudicella colmanti is native to Gabun, Kamerun and Kongo, thus a species with a main distribution in Central Africa. But E. colmanti is like other species of this genus worldwide often kept in terraria, although species like E. smithi are more common inhabitants of this kind of artificial habitats. They all can be more or less easily reared.

 

Specific flying mode and copulation behavior

 

This is why I was able to study behavioral characters in detail. And rose chafers indeed show interesting behaviors. They for example perform a unique way of flying. It is a specific character of rose chafers (a so called apomorphy) that they fly with closed fore wings, which cannot be opened as in other beetles.

I documented in my video the mating behavior of a beetle couple. Interestingly this was not too difficult, although both genders can, when separated from each other, react to disturbances with a high agility.

 

Almost permanent copulation activities

 

But in the copulatory position, they accepted to be removed from their terrarium to the filming set and even stayed in position, when they were enlighted from different positions with very bright light beams. Please note the the female, which I observed regularly actively searching for a position underneath the male (behavior not clearly visible in my footage). But it also conspicuously never stopped feeding (on an apple) during the copulatory process (very well visible in my footage), obviously to obtain enough nutrients for the production of eggs. A copulation in my couple is not a unique event, but is repeated regularly and can take hours.

 

Phoretic mites

 

Both genders carried bigger numbers of mites. These were phoretic deutonymphs of the taxon Astigmata (Acariformes, Acaridae). As never determined the mite species, as it was not clear, whether it represented a natural associate of these tropical beetles, or whether it was a species native to Germany, which for example was carried into the terrarium via Drosophila flies.

Copyrights Stefan F. Wirth, Berlin March 2017/ February 2019

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

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