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.
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.
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 isHistiostoma 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. palustredo 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. palustrewas 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.
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.
Bumble bee Bombus sp. in Berlin, copyrights Stefan F. Wirth 2021/2022Honey bee Apis mellifera in Berlin, copyrights Stefan F. Wirth 2021/2022Deutonymphs of the microscopically tiny mite Schwiebea cf. eurynymphae (Acaridae, Astigmata) formally attached to beetle Phosphuga atrata under the bark of felled tree trunk of Tilia platyphyllos in urban park Rehberge in Berlin, copyrights Stefan F. Wirth, 2021/2022
Larvae of beetle Oryctes nasicornis from Italy with associated gamasid mites under studio light conditions, copyrights Stefan F. Wirth, Berlin 2016/2022
Land crab Metasesarma obesum under studio conditions, copyrights Stefan F. Wirth, Berlin 2017/2022
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Was ist ein Ökosystem? Welche Ökosysteme sind gut untersucht, welche eher nicht? Wie gut kennt man die Artenvielfalt von Mikro-Lebensstätten in Deutschland, und was ist über deren biologische (ökologische) Zusammenhänge bekannt? Was ist denn eigentlich eine Art, was sind denn dann Zwillingsarten, und was versteht man gar unter einem Artenkomplex (kryptische Artengruppe)? Ist das Aussterben von Arten ein normaler Bestandteil der Evolution oder ist das Aussterben einer Art immer zwingend ein alamierender Hinweis auf eine (evtl. menschengemachte) Naturkatastrophe? Wieviele Arten aus allen Organismengruppen weltweit kennen wir, und wieviele in etwa kennen wir noch nicht? Warum kennen wir viele Arten, sogar in Deutschland, noch immer nicht? Wie erkennt man neue Arten, und wie ist eine sogenannte Artbeschreibung aufgebaut? Ist der Mensch eine Tierart, und wo im Stammbaum der Tiere ist er dann anzusiedeln?
Warum sind ein Wald, ein Teich oder eine Wiese Orte für interessante Entdeckungen, und zwar insbesondere auch für Kinder? Was lebt denn da, und wie ist es an seinen Lebensraum angepasst? Was haben unterschiedliche Arten in solchen Lebensräumen eigentlich miteinander zu tun? Und wie beobachtet man Tierverhalten am besten? Wie dokumentiert man es aussagekräftig, um sein Wissen später mit Freunden oder über soziale Netzwerke teilen zu können?
Wie kommt es zum sogenannten Global Warming, der globalen Klimaerwärmung? Wie können wir sie nachweisen? Warum ist sie zu einem beträchtlichen Teil menschengemacht? Und welche Auswirkungen haben Klimaerwärmung und die Ausbeutung natürlicher Ressourcen (Energiespeicher, Rohstoffe, wie zum Beispiel Tropenholz) für die Zukunft der Menschheit und die Artenvielfalt auf unserer Erde. Welche Auswege erhofft man sich? Woran wird derzeit gearbeitet?
Was benötigt man zur Naturfotografie, was, wenn man zusätzlich oder alternativ auch noch auf gutem Niveau filmen möchte? Was ist grundsätzlich wichtiger: Das Equipment oder das Bild, das zuvor im Kopf des Fotografen oder Filmers entsteht? Muss taugliches Equipment immer ultra-teuer sein? Welche Software eignet sich am besten zum Editieren? Was genügt dabei den Ansprüchen von Anfängern, was benötigen Fortgeschrittene und Profis? Wie filme oder fotografiere ich draußen in der Natur? Wie hole ich stattdessen die Natur in mein Fotostudio und inszeniere sie dort so, dass es aussieht, als habe man im Freien gearbeitet?
Dies sind alles mögliche Themen, die in meinem Unterricht, meinen Kursen oder Vorträgen vertieft werden können. Beliebige weitere Fragestellungen aus den Bereichen Naturkunde, Biologie, Ökologie und Evolution arbeite ich gerne für Sie aus.
I offer lessons, remedial courses, lectures and advanced training courses on the subjects of natural history, nature conservation, biodiversity, ecology, climate protection and evolution, as well as lessons or lectures on nature photography or nature filming. All this either on a fee basis or via employment. Please see my menu item on the subject of teaching for further information. Of course, I have proofs of qualifications for my various previous teaching activities as well as my professional competence. Please also note my menu item Curriculum Vitae.
But what are my topics? In the following you will find interesting questions from my areas of competence:
What is an ecosystem? Which ecosystems have been well studied and which not? How well do you know the biodiversity of micro habitats in Germany and what is known about their biological (ecological) relationships? What is actually a species, what are sibling species, and what is meant by a species complex (cryptic species group)? Is the extinction of species a normal part of evolution or is the extinction of a species always an alarming indicator of a (possibly human-made) natural disaster? How many species from all groups of organisms worldwide do we know, and roughly how many do we not yet know? Why do we still not know many species, even in Germany? How do we recognize new species and how is a so-called species description structured? Are humans an animal species, and if so, where do they belong in the animal tree?
Why are a forest, a pond or a meadow places for interesting discoveries, especially for children? What lives there and how is it adapted to its habitat? What do different species actually have to do with each other in such habitats? And what is the best way to observe animal behavior? How can you document it meaningfully so that you can later share your knowledge with friends or via social networks?
How does the global warming come about? How can we prove its existence? Why is it largely human-made? And what are the effects of global warming and the exploitation of natural resources (energy stores, raw materials such as tropical wood) on the future of humanity and biodiversity on our planet? What exits to avoid emergency situations are we hoping for? What are scientists currently working on to ensure a healthy human future?
What do we need for nature photography, what if we also want to film at a good level in addition or as an alternative? What is fundamentally more important: the equipment or the image that is created in the head of the photographer or filmmaker? Does suitable equipment always have to be ultra-expensive? Which software is best for editing? What meets the requirements of beginners, what do advanced and professionals need? How do we film or take photos outdoors in nature? Instead, how do we bring nature into our photo studio and stage it there in such a way that it looks as if we were working outdoors?
These are all possible topics that can be deepened in my teaching, courses or lectures. I would be happy to work out any other questions from the fields of natural history, biology, ecology and evolution for you.
all copyrights Stefan F. Wirth Berlin 2022
More Specialists are needed to study our biodiversity: recognizing and describing new species, redescribing known ones, mapping their distribution and understanding their ecological role in an ecosystem.Thus we have to support our children and students to become fascinated by nature.
Also interested laypeople, hobby researchers and nature lovers can contribute to species preservation and nature conservation (and thus climate protection) and encourage their children or relatives to study biology or a similar subject.
Es werden mehr Spezialisten benötigt, um unsere Biodiversität zu studieren: neue Arten zu erkennen und zu beschreiben, schon bekannte Arten neu zu beschreiben, ihre Verbreitung zu kartieren und ihre ökologische Rolle in einem Ökosystem zu verstehen. Daher müssen wir unsere Kinder und Schüler dabei unterstützen, sich für die Natur zu begeistern.
Auch interessierte Laien, Hobbyforscher und Naturfreunde können einen Beitrag zum Arten- und Naturschutz (und damit zum Klimaschutz) leisten und ihre Kinder oder Angehörigen für ein Studium der Biologie oder eines ähnlichen Faches animieren.
I provide advices and information about the topics mites (in general, in your house or your company and in a hygiennic context), biodiversity, correlation biodiversity research and climate change, speciation processes, describing new species, species extinction, taxonomy for private people, educational institutions, e.g. schools or university students. Please see my menue item „Angebot biologische Beratung…“
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The mite Histiostoma piceae Scheucher, 1957 is a member of the mite family Histiostomatidae (Astigmata, Acariformes). Scheucher discovered the mite based on all instars from spruce, infected by the bark beetle Ips typographus. She collected her samples in Regensburg, Höbing (bei Roth) and Harz. Scheucher reared her specimens on potatoes and bran, but describes that her cultures did grow well only to some degree.
According to her findings, phoretic carrier (hosts) is the bark beetle species Ips typographus, she also found deutonymphs rarely on some staphylinids. She discovered that free living non-deutonymphal stages develop on fresh detritus, while deutonymphs appear only on old detritus („after it was for a longer time removed from the trees“, „wenn der Mulm einige Zeit aus den Bäumen entfernt ist“). I could like Scheucher culture the mites on potato, but a bit better in their original gallery substrate. Under laboratory conditions, they indeed did not rear very well in both kinds of cultures.
I collected H. piceae between 2000 and 2004 once from a wooden log infested by I. typographus in Berlin, then got access to microscopic slides from Europe in the collection of John C. Moser (Louisiana, USA) in 2007 and 2009, then I collected samples from Ips typographus and I. cembrae in Central Croatia (publication Wirth, Weis and Pernek, 2016) and found out that H. piceae is not restricted to I. typographus, but also to its sibling species I. cembrae. I finally collected the mite from I. typographus galleries between 2015 and 2016 in Western-Siberia near the city Tyumen.
I repeatedly observed deutonymphs of H. piceae under natural conditions (bark samples directly after the excursions) to develop in very high numbers, then attaching to all available arthropods nearby, smaller bark beetle species and numerous bigger mites of different groups, such as for example oribatids.
Published recordings of H. piceae from other bark beetles than I. typographus and I. cembrae are doubtful and need to be named Histiostoma cf. piceae. In some cases with I. typographus additionally present, I interpret the mites to have switched from their regular carrier (host) to an adjacent gallery of e.g. another smaller bark beetle species. In other cases, the existence of similar looking species new to science needs to be tested. In cases of determinations by non specialists from bark beetles other than the above mentioned two beetle species, it needs to be assumed that these people could not differ between similar mite species, such as Histiostoma trichophorum Oudemans, 1912, Histiostoma ulmi Scheucher, 1957 or Histiostoma crypturgi Scheucher, 1957.
I never before published the full set of SEM and light microscopic photos from these times (except of my article about host specificity). In this explicite photo publication here on my homepage, I herewith publish SEM-photographs, objects sputtered with gold, which might be not unique to science, but very rare.
Any subsequent research on this mite in Europe is not happening (a few not too relevant findings are published by a former Russian colleague). Reason is that modern science does not understand, especially not in Germany, that fundamental research in applied fields is worth to be funded. It is for example known that deutonymphs of different mite species on bark beetles regularly carry fungus spores (different fungus species, just sticking on the mite’s cuticle), discovered by John C. Moser and confirmed by several of my own publications. This phenomenon is still not closer studied. Fungus transport into bark beetle galleries can influence the micro climate there.
Male and female of Histiostoma piceae, A venter of male, B dorsum of male, C mouthparts with Digitus fixus, D dorsum of female, E side-frontal view to female; Berlin 2002-2020, copyrights Stefan F. Wirth
Deutonymph of Histiostoma piceae in ventral view, collected in Western Siberia, 2015 – 2016, copyrights Stefan F. Wirth
Systematics: Histiostoma piceae is according to my phd thesis from 2004 and according to my more recent research findings a member of a clade (monophylum) within Histiostomatidae with most species associated with bark beetles (Scolytinae) or other bark inhabiting coleopterans; these phylogenetic findings are based on morphological characters.
The mite Histiostoma maritimum Oudemans 1914 is a member of the mite family Histiostomatidae (Astigmata, Acariformes). Oudemans discovered the mite based on its deutonymph only from a Dutch island. The German acarologist R. Scheucher found the species in 1957 in mud at the riverside of Regnitz and for the first time could rear H. maritimum and was able to redescribe it by its adult stages, especially females look morphologically conspicuous due to a sclerotized cuticula shield around its copulation opening. She reared her specimens on potatoes, mud and bran, but describes that her cultures did not grow well.
Phoretic carrieres (hosts) are beetles of genus Heterocerus, some carabids and according her findings also rarely some staphylinids.
I discovered H. maritimum between 2000 and 2004 repeatedly in sapropel around ponds in an old gravel pit area in Berlin, forest Grunewald, named „im Jagen 86“. They were mainly attached to the beetles Heterocerus fenestratus and Heterocerus fusculus, but could regularly also be found on the carabids Elaphrus cupreus and Bembidion sp.. I could several times rear the mites, like Scheucher almost unsuccessfully on potatoes, but well on cadavers of their carriers. I thus reconstructed a so called necromenic life-strategy for H. maritium. This means that a phoretic stage ascends a carrier, but never leaves, instead it awaits the carrier’s natural dead to develop on its cadaver (published in my phd thesis, online, 2004).
I will not publish my full set of SEM photos from earlier times here. Some photos will be saved for one of my upcoming paper submissions in scientific and peer-reviewed journals. In this photo publication here on my homepage, I at least publish some interesting SEM-photographs, based on objects sputtered with gold and a subsequent critical-point-drying procedure.
Adults of Histiostoma maritimum: A left male, right female, B, C, copulation opening, D dorsal view to female with mouthparts and copulation opening
Systematics: H. maritimum shares morphological characters of deutonymph (setation, apodemes) and adults (mouthpart details, shape of Digitus fixus) with species like Histiostoma feroniarum, H. insulare, H. litorale, H. palustre, H. polypori, H. myrmicarum. This might indicate a separate clade, but according to the old findings in my phd thesis, also a paraphyletic grouping including these species is thinkable.
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
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
We recently read a lot about the pandemic consequences of infections with the new corona virus Sars-CoV-2, most are medical issues, hygienic advises and information about political reactions in different countries worldwide. But there is not much known about the biological host reservoir, putative intermediate hosts and how the human infections might be explained. It is a normal lack of information, because the scientific research about topics, being generally new to science, is time costing, especially, when life strategies and the population dynamics of organisms a concerned. Organisms? Viruses are per definitionem not considered organisms, because they lack important aspects, which characterize real life: they cannot reproduce on their own power, they do not have an own metabolism, no ingestion, no excretion. But they are organic and show traces of life by possessing a genome, which might indicate that they evolved from living cells. Viruses represent a diverse group of protein bodies containing nucleic acid, either DNA or RNA.
New corona virus Sars-CoV-2, Wikipedia: CDC/ Alissa Eckert, MS; Dan Higgins, MAM / Public domain
Viruses in general, host specificity, host increase, host change
For reproduction viruses depend on living host cells, which they reprogram by inserting their virus genome into the cell’s genome in order to stimulate the forming of a number of virus copies, all that happening on cost of the host cell’s life. Thus viruses need to be named parasites as they harm their hosts to their own advantage. Different groups of viruses attack different kinds of cells using in detail different methods to enslave their host cells. There are plant viruses, viruses associated with bacteria (named bacteriophages) and animalistic viruses. They all show characters, which are typical for parasite – host – relationships. Parasitic partners of any kind of host – parasite – relationship can be exclusively associated with one host species only (host specificity) or a limited group of systematically closely related hosts, while others can have a wider range of different host species. The latter generally might have evolved out of the former, although also the opposite direction is thinkable. When former host-specific parasites make themselves one or even several further hosts accessible, then this phenomenon is named host-increase (Wirtserweiterung). In case an new host was infested as permanent host, while the former host is given up, then a so called host change (Wirtswechsel) happened. The same term is also used in a different context, namely when a parasite requires in its development a change between different hosts.
Host specificity: A parasite (or an organism with similar life-strategy) is associated with one host only, which requires a specialization and a competition between host evolution and parasite evolution (coevolution). This strategy needs to be separated from generalism, which means that a parasite has a very wide range of not related regular main hosts. Host specificity is more common than generalism. But this also depends on definitions. I herewith define the association with one main host species only as host specificity. But I furthermore consider host specificity also given, when parasite-host relations are specific on a higher taxonomic level, for example, when certain closely related genera of parasites are specialized for certain closely related genera of hosts. This part of my definition has variable borders. In the chapter after next, I describe the parasitic case of the trematode Leucochloridium paradoxum, whose main hosts are represented by different systematically not closer related bird species. A host specificy on the level of birds in general (Aves), then present in only some species with similar food preferences might already need to be named a limited generalism.
Obligatory host change in ticks and lifstyle-change in water mites
Some parasites need several hosts to be enabled to finish their life-cycles. This is another context, in which the German term „Wirtswechsel“ (host change) is used. In that kind of parasite – host – association, the host change is often obligatory, meaning that the parasite cannot survive in the absence of one of the required hosts. The castor bean tick Ixodes ricinus represents a parasite, which needs a host change to successfully go through its full development until adulthood, but there is a wider range of suitable hosts, as intermediate host and as final host. Thus the tick is a generalist with obligatory host change. Water mites (Hydrachnidia) are parasitic as first nymphs (juvenile instar, usually named „larva“) and predators as older nymphs and adults. A host specificity of „larvae“ can appear, but a wider range of host species is common. These mites perform a life style change during their development.
Intermediate host, for example in the parasitic flatworm Leucochloridium paradoxum
An example for a parasite, obligatory requiring a specific intermediate host, is the flatworm Leucochloridium paradoxum („green-banded broodsac“, Trematoda, Platyhelmintes), whose larvae (miracidium) need to infest snails of the genus Succinea. This trematode parasite is host specific for a genus of snails, while there is no specificity for their main hosts. They parasite birds, but infest different bird species, which are not closer related to each other, such as finches, the crow family Corvidae or woodpeckers. Although there is a main host specificity on the very high taxonomic level of Aves, the use of the term (limited) generalism might in this case even be appropriate. Inside the smail’s midgut gland, miracidia (larvae) modify into another larva-form, named cercaria, which invade the liver, where they form so called sporocysts, sac-shaped muscular tubes, which grow through the entire snail host until they reach the snail’s tentacles, which they fill up with their tube-shaped bodies entirely. Lastly the snail is unable to retract her swollen organs. The snail tentacles are now well visible as conspicuous greenish stripes, pulsating permanently. The sporocysts as larval stage of this trematode parasite do even more than only increasing the visibility of the snail for bird predators, which represent the worm’s final host. They additionally manipulate the nervous system of the snail so far that the snail performs an unusual behavior and moves towards very well exposed elevated areas, such as leaves of adjacent plants. Thus the probability to be eaten by birds is remarkably increased.
Host specificity on humans with side-hosts and coevolution with the ancestor line of Homo sapiens: skin mite Sarcoptes scabiei
An interesting example of a host specificity with numerous side-hosts and even an additional host-increase is the skin parasitic mite Sarcoptes scabiei (also named the „seven-year itch“). It was originally exclusively specific for Homo sapiens and accompanied mankind over its entire evolution (e. g. J. R. H. Andrew’s Acarologia, 1983). Systematical relatives of that mite species can only be found within the Great Apes. Originating from the recent Homo sapiens, S. scabiei conquered the human’s domestic animals, such as dogs or bovine animals within long-term periods, in which humans and their domestic animals had shared the same buildings or even rooms. Domestic animals may transfer the mite-parasite subsequently to wild animals. In case main host (humans) and side hosts (domestic animals, wild animals) can supply everything, which the parasite needs for its development without the necessity to leave its host specimen, one might speak about real hosts. In case side hosts cannot supply the necessary basic equipment, they represent either intermediate hosts or dead-end hosts. It can for example be discussed, whether dogs might in fact be dead-end hosts, as the skin disease can harm them under certain conditions to dead.
Host increase due to the globalisation and human economic interests: example honey bee parasite Varroa destructor (mite)
Another example of a former host specificity on a species‘ level with host increase is the mite Varroa destructor (Parasitiformes, Mesostigmata). It was originally specific for the Eastern honey bee Apis cerana. The mite could only switch over to the Western honey bee Apis mellifera due to a human influence: Men transferred A. mellifera for economic reasons to the natural habitats of A. cerana in Eastern Asia, were it got infected by the mite V. destructor. A subsequent transfer of the Western honeybee back home established the mite parasite in Western countries. As A. mellifera colonies are much more harmed by V. destructor than its original host, our honey bee must be considered as an intermediate case between a new host and a dead-end host. Human international traffic enabled this host-increase primarily, although there are areas between Afghanistan and Iraq, where both bee species coexist due to natural distribution. But there is an almost insurmountable (allopatric) desert border between the population of both species of about 360 to 600 kilometers, although there are evidences for bees rarely surmounting this border. Thus a natural mite transfer between closely related bee species might have happened additionally. Species of animals, plants, fungi or bacteria and even viruses, which successfully established new (additional) living spaces are named neobiota or alien species.
Mite Varroa destructor, Wikipedia: The original uploader was Tullius at German Wikipedia. / Public domain
Can viruses as non-living genome possessing lumps be subject of evolution and complex host – parasite relationships?
Can this high complexity of modes of parasite – host – relationships in living organisms also be found in virus – host – relationships, although viruses do not represent living organisms at all according to biological definitions? The answer is yes, because viruses do not only share a genome with living cells, but based on this genome even are subject to the mechanisms of evolution. And evolution was the most important factor in all the mentioned complex parasite – host – interactions.
Parasitism versus mutualism or harming the host or not harming the host
Two different life-strategies with similar mechanisms as organism – to – organism associations
Are there other organism – to – organism relationships, being subject to a similar complexity than found in parasites with their hosts? Yes, a superordinate term for other close associations between different organism species is mutualism. While parasites need to harm their hosts by using them as final living-sources, mutualists are considered to practice a more neutral host contact, which per theoretic definition means that nobody harms anybody. But the assumption of a neutrality is in fact an artificial construct, as in detail it can come out that some of these organism associations represent unrecognized parasite-relationships, while in other cases a benefit for both partners (symbiosis) or for one partner only might be discovered in future studies. At least so called mutualists share as a feature that harmfulness or benefit are not easily noticeable.
Phoresy: taking a ride on a taxi-host as example of mutualistic relationships
An example for a more neutral organism, at least not harming association is called phoresy. It is often performed by nematodes and mites. These tiny organisms take a ride on bigger animals in order to become carried from one habitat to another. This „taxi-association“ is considered being of advantage for the phoretic part and harmless for the carrier (in English also often named host). But there are seeming phoretic interactions known, which based on developing technical scientific standards could be identified as unusual cases of parasitism. An example is a phoretic instar of an astigmatid mite (Astigmata, Acariformes), which as all phoretic instars within this big mite clade has no functional mouth, but sucking structures to fix itself to its host. This specific mite species had evolved a mechanism for opening the host cuticle in order to incorporate blood of its host using these sucking organs. This is unlike the common use of homologous suckers in related mite taxa, where they (as far as known so far) only support the adherence.
Another interesting example of a phoretic mite is Histiostoma blomquisti (Histiostomatidae, Astigmata), which is specifically associated with the red imported fire ant (sometimes referred as RIFA) Solenopsis invicta, which worldwide appears as troublesome neozoon, again a result of human global traffic. I am the scientific describer of that mite, and my research about it’s biology and abundance in ant nests refers to populations in Louisiana (USA). An interesting aspect is that the ant is originally native to Southern America. We lack studies, whether the mite appears in the native habitats of the ant also as its specific cohabitant or whether it originally deals with a wider range of phoretic hosts. We do not even know, whether the mite is at all native to the same area, in which S. invicta had its natural distribution. On one hand, we hypothesise that, but there is also a theoretical option that the mite performed a subsequent host change in areas, for example in the Southern USA, where the ant was accidentally established via sandy ballast substrate of ships as neozoon. It is further more not known, whether the mite – ant – relationship is indeed neutral, at least with no noticeable harming features. I discovered (S. Wirth & J. C. Moser, Acarologia 2010) that mite deutonymphs (= phoretic instar) can attach to active nest queens in such extraordinary high numbers (hundreds of mite specimens) that mobility restrictions for the concerned queens were sometimes visible. On the other hand, my video documentations showed that even completely overcrowded queens could still freely move and, much more important: stayed reproductive. The purpose of the mites inside the fire ant nests is unknown. But generally, mites of the Histiostomatidae can appear as beneficial animals in ant nests. At least according to my findings about the mite Histiostoma bakeri, which is a phoretic associate of the leafcutter ant Atta texana in Southern USA. I discovered these mites improving the hygienic conditions inside specific nest chambers (detritus chambers) due to their fungi and bacteria feeding activities (Wirth & Moser, European Association of Acarologists proceedings, 2008).
I will in different chapters of this article repeatedly refer to examples with phoretic mites of the family Histiostomatidae (Astigmata, Acariformes). As mutualism and parasitism follow similar organism-host association patterns, I will in those chapters not each time mention again that examples with these mites do not concern parasitism, but mutualism. It is by the way no accident that both life-strategies share common features, as there are examples known, which indicate that one strategy can evolve out of the other.
Mite Histiostoma blomquisti Wirth & Moser, 2010 (Histiostomatidae, Astigmata, Acariformes) on queens of ant Solenopsis invicta, Pineville/ Louisiana, copyrights Stefan F. Wirth
Mutualism often used as neutral term for organism associations with unknown effect of both partners to each other.
The copepod (Crustacea) Ommatokoita elongata on Greenland and sleeper sharks
So called mutualistic associations can sometimes represent interactions of unknown benefits or damage regarding both of the associated partners. Another interesting example of such an association with a not yet understood status is the copepod Ommatokoita elongata (Crustacea), which was discovered as specific cohabitant on the Greenland shark (Somniosus microcephalus) and the pacific sleeper shark (Somniosus pacificus). Larvae of the crustacean in their copepodit stadium and adult females attach to the ocular globes of the shark, where they can cause visible tissue damages. They are thus considered being parasites, although alternating hypotheses assume a more neutral mutualistic copepod – shark – association, based on the sometimes high abundance of the crustacean on one shark specimen (B. Berland, Nature, 1961). There are even assumptions about a benefit contributed by the copepode to the sharks: reasearchers say that it might improve the shark’s hunting success by attracting suitable prey with bioluminescence signals.
Shark Somniosus pacificus, Wikipedia: National Oceanic and Atmospheric Administration / Public domain
Greenland shark with copepod Ommatokoita elongata, hardly visible, when the shark turns to show his right eye, Youtube: copyrights The Canadian Press, video by Ben Singer, footage Brynn Devine, Marine institute of Memorial University of Newfoundland
Human parasites with mutualistic features: the mites Demodex folliculorum and D. brevis
Can viruses be compared with some mites, nematodes or copepodes by performing mutualistic virus – host – relationships? A priori it must be stated that they are unable for a neutral relationship with another organism, as they need the destruction of living cells for their own persistence. But indeed there are viruses known, causing no known diseases and thus being named passenger viruses. But first, an example of an organismic example of parasitism without harmfulness will be presented: the mites Demodex folliculorum and Demodex brevis (Trombidiformes, Prostigmata), which appear as so named „face mites“ inside hair follicles of humans, preferring eyebrows and eyelashes, but also other hairy body parts. The abundance in humans is high and grows with a human age. According to Schaller, M. (2004), new born children are free of Demodex, while over 70 years old people are at almost 100 percent infested with the mites. The mite in fact is a parasite and feeds on sebum from the sebaceous glands. Incorporating needed human gland secretions must be named parasitism. Nevertheless mites under normal conditions cause no visible damages nor do they seem to harm their host noticeably.
So called passenger viruses as mutualists with a more or less neutral affect to their human hosts
Such a parasitic relationship might be comparable with so called passenger viruses, which do not harm noticeably, although they destroy living tissue as all viruses do. They can accompany more harmful viruses and even might harm the pathological success of the diseases, caused by these harmful viruses, and for example might slow the disease’s progression. An example is the GB virus C (GBV-C), which was before known as Hepatitis G virus. The virus is common in humans and shows no pathogenic damaging effect. According to an US-study, about 13 percent of probands, whose blood was examined, possessed antibodies against the virus. GBV-C is considered to slow the effects of an HIV disease by negatively effecting the replication of the HI-virus.
Host increase towards systematically not closer related new hosts
Example for a transfer within related host taxa in mites is the bark-beetle-clade within Histiostomatidae (Astigmata), an example for non related side hosts is the mite Histiostoma maritimum (Histiostomatidae, Astigmata)
Do side-hosts or intermediate hosts as results of host increases commonly need to be systematically close relatives of the main host? The answer is no, although parasites are usually better pre-adapted in infesting a host, which shares a maximum of common characters with the main host. Within the mite family Histiostomatidae, there exists a clade of mites being associated with a clade of beetles. I named it bark beetle-clade (e.g. Wirth, phd thesis, 2004). Mites and bark beetles performed a parallel evolution, which required host increases and host changes towards related hosts and subsequent evolutionary adaptations to harmonize with these new hosts, either to become specific for a new host or to deal with a range of host species.
But the transfer of a parasite to new hosts can also happen towards not closely related host species, representing a scenery being based on a common ecological context between main hosts and side hosts. The phoretic mite Histiostoma maritimum for example is host specific for at least two closely related beetle-species of genus Heterocerus (Heteroceridae). But the mite regularly also appears on predatory beetles of genera Elaphrus and Bembidion (Elaphrus cupreus and Bembidion dentellum, Carabidae) (S. Wirth, phd thesis 2004 and subsequent studies). These beetles partly share the same habitats with Heterocerus: sapropel around ponds, being exposed to sunlight and warmth. In my research about the mite H. maritimum, I hypothesised that the phoretic mite instar might switch over to Elaphrus and Bembidion, for example when these predators feed on adult Heterocerus beetles, larvae or cadavers. Although I could regularly find mites in lower abundances over years on the side hosts (collected in the Heterocerus sampling sites), it is unknown, whether the „switch-over“-scenario was a starting event in an evolutionary past to establish the mite to new additional hosts, where they would today survive more or less independently from the original Heterocerus source, or whether the mites regularly need to switch over in the above mentioned situations, and in consequence side hosts with no Heterocerus-contact would thus lack the mite. A possible support for the latter hypothesis are my laboratory findings about the preferred developmental habitat of the mite, which was cadavers of died Heterocerus beetles. In my experiments the mite remained on its Heterocerus– carrier until this died. Mites subsequently developed on the beetle’s cadavers, feeding there on bacteria and fungi (the phenomenon is named necromeny). Mites under laboratory conditions developed also seemingly successfully on E. cupreus– and B.dentellum-cadavers. But I could so far never continue these studies and don’t know, whether or how well mite colonies with having only cadavers of these two side-hosts available would reproduce compared to mites being reared in Heterocerus settings. In case of a strict substrate specialization for Heterocerus cadavers, the side hosts would be dead-end hosts, and permanent reinfections from the original host source would be required to explain the regular mite abundance in Elaphrus and Bembidion.
Histiostoma maritimum, a adult female with conspicuous copulation opening, b both adult genders in dorsal view, c, d copulation opening in dorsal and sideview, SEM, Berlin 2020/ ca. 2002, copyrights Stefan F. Wirth
Assumed transfer of virus SARS-CoV-2 from bat main hosts via a pangolin as intermediate host towards humans:
There is an ecological context between bats and pangolins
The new corona virus SARS-CoV-2 is assumed to be host specific to a group of animals and from there infesting another animal as intermediatehost, from which presumably humans were opened up as new host source. There are researchers interpreting us humans as an dead-end hosts, as unlike in bats human people can be harmed remarkably with the lung disease COVID-19 (corona virus disease 2019), triggered by SARS-CoV-2. As at least from a general statistical point of view a high majority of infested people shows no or only slight symptoms, thus it can up-to-date not be excluded that Homo sapiens is in order to become a fully potential side host, because all a parasite needs in order to „survive“ before all other requirements is the (statistically) surviving of its host.
There is evidence that bats (Chiroptera) represent the main host, thus representing the „natural virus reservoir“, while pangolins (Pholidota) presumably act as intermediate hosts. This main-host-to-intermediate host context is for example reported as putative scenario by Ye Z.-W et al. (Int Biol Sci, 2020), who stated that based on molecular features the bat Rhinolophus affinis (Rhinolophidae, Chiroptera) is hosting a virus most similar to SARS-CoV-2 differing from all other known corona viruses (Similarity 96.2 %, nucleotide homology). The pangolin species Manis javanica was identified to carry formerly unknown CoV genomes, being according to the same authors with 85-92 % similar to SARS-CoV-2 (nucleotide sequence homology).
Bat Rhinolophus affinis as known reservoir of a virus most similar to Sars-CoV-2. Wiki commons: Naturalis Biodiversity Center
Pangolins and Chiroptera (bats and megabats, this taxon subsequently sometimes refereed as „bats“) are systematically not closer related to each other. Pangolins (Pholidota) are considered to represent the sister taxon of the clade Carnivora. Chiroptera were reconstructed as sister taxon to the clade Euungulata (containing animals such as horses, cattle or whales). But both, Chiroptera and Pholidota, can be connected by an ecological context. Pangolins (Pholidota) are species, which are either adapted to live preferably on the ground, or to spent most of their time on trees. Both types are specialised ant and termite feeders, which use cavities on the ground or inside trees as hideaways. They additionally give birth to their offspring inside these burrows and subsequently use to stay there with their young for a while. Such cavities can accidentally be the same time aggregation and resting places for bats, excluding megabats, which use to rest during daytime on exposed areas on trees. Manis javanica has a semi-arboricol life-style, spending time in trees and on the ground. This pangolin uses different resting cavities, either subterranean burrows or tree cavities.
Chinese pangolin Manis pentadactyla, a ground living species, Wikipedia: nachbarnebenan / Public domain, Zoo Leipzig, Tou Feng
Pangolin Manis javanica as known host of a virus similar to virus SARS -CoV-2. Wikipedia: creative commons Piekfrosch / CC BY-SA
Chiroptera and Pangolins are in South Eastern counties often subject to hunting, as both for example play a role in the traditional Chinese medicine. Thus a virus transfer to humans via main host or via the putative intermediate host is assumed to have happened on animal markets (in the province Wuhan in China).
Which indications point to animal hosts as original source of virus SARS -CoV-2 ?
The scientists Andersen et. al (2020) explain there was no virus-engineering instead of a natural evolution
But which proofs exist that animal hosts sources such as Chiroptera and pangolins are involved in the transfer of the virus SARS -CoV-2 to humans? The lack of general knowledge is still fundament for conspiracy theories, such as an artificial creation of the new corona virus in laboratories with biological warfare purposes.
K.G. Andersen et al. („The proximal origin of SARS-CoV-2“, Nature Medicine, 2020) concluded based on their molecular research that the genetic template for specific spike proteins forming structures, which the virus body possesses on its outside for holding on and penetrating into the host cells, showed evidence for a natural evolution and not for an engineering. They argue with the strong efficiency of the spikes at binding human cells, which makes an engineering implausible and evolution based on natural selection highly probable. The authors additionally examined the overall molecular structure of the backbone of SARS-CoV-2. Backbone can be explained as the „skeleton spine“ of a macromolecule as a continuous row of covalent bond atoms. This overall backbone structure of the new corona virus is according to the authors similar to viruses, which were isolated from Chiroptera and pangolins and dissimilar to other corona viruses, which are already known to science.
Spikes (here in red) in Sars-CoV-2 hold on and penetrate into host cells, Wikipedia: CDC/ Alissa Eckert, MS; Dan Higgins, MAM / Public domain
Can a host increase happen more or less spontaneously with a subsequent enormous success (as for example in virus SARS-CoV-2)?
And: Can the complexity of adaptations to a main host decide for the option of a host increase?
An example for a tendency to spontaneous temporary host changes is mite Histiostoma piceae (Histiostomatidae, Astigmata)
Is it imaginable that a host change or a host increase happens spontaneously and subsequently having such a remarkable impact to the new host, as it is recently ongoing with the SARS-CoV-2 pandemic? Host specificity, host changes and parasitism or mutualism in general are result of evolution. The most common case of evolutionary changes in organisms or viruses is a slow process of stepwise modifications being based on mutations and natural selection.
But it needs also to be stated that as more complex the pattern of characters is (genome, morphology, behavior, function-morphology, reproduction biology etc.), which binds a parasite or mutualist to a specific host, as more evolutionary steps are necessary to perform a host change and as longer an exposure to mutation and selection would need to take place. However it is alternatively possible that a host specificity is only based on a few, but important features. Slighter ecological pressures focusing towards these features might then theoretically allow rather fast host changes.
As an example with a putatively reduced complexity of host adaptations I herewith introduce the phoretic mite Histiostoms piceae (Astigmata, Histiostomatidae), which I repeatedly studied and reared under laboratory conditions. The scientific describer of this species (Scheucher, 1957) discovered a strict host specificity to the bark beetle Ips typographus. According to my and her research, the mite has along the geographic distribution of that bark beetle a high abundance, beetles without the mite are rare. In 2016 I discovered H. piceae being additionally associated with Ips cembrae as a second regular host. I cembrae represents the sibling species of I. typographus (Wirth, Weis, Pernek, Sumarski List, 2016). Exceptions are smaller bark beetle species, which regularly burrow their galleries into those of I. typographus or I. cembrae. It is unknown, whether these small bark beetles as cohabitants of I. typographus carry the mite temporarily or regularly. But the former might be confirmed by the following interesting phenomenon in the mite H. piceae:
In case of very high numbers of mites inside bark beetle galleries and a relatively low numbers of corresponding Ips species, the phoretic instar of the mite attaches under natural field conditions all available arthropods inside or adjacent to the galleries of the main hosts, including bigger mite species, different beetle species or – as already mentioned – smaller bark beetle species (for example my studies in the area of the city Tyumen, Siberia, Russia, 2015-2016). This indiscriminateness for specific hosts under certain conditions might indicate that the substrate specificity of the mite H. piceae is more developed than the phoretic specificity for the host insect itself as a carrier . In such a case, I would generally expect that a host change or a host increase might faster happen in future evolutionary steps than in mite species, which are strictly choosy for their specific host carrier. In H. piceae the tolerance for a variety of carriers (unlike the specificity for substrate conditions) might in a future evolution even succeed as pre-adaptation, which under suitable circumstances might spontaneously allow a regular transfer to new hosts. A second important step towards a real host increase would require that the mite becomes able to stay permanently on its new host. In the H. piceae context the evolution of a tolerance for different substrate conditions might once become an important selective factor in may be opening up new permanent host-associations.
Temporary side hosts, as described in the above explained observations, would represent nothing then dead-end hosts, as they are unable to carry the phoretic mite to suitable habitats for its development. But under favorable circumstances, a former dead-end host might even become a new permanent host.
Histiostoma piceae, a adult female in side view, b in dorsal view, c mouthparts and digitis fixus, d adult male in dorsal view, e in ventral view, Berlin 2020/ ca. 2002, copyrights Stefan F. Wirth
Phoretic instar of Histiostoma piceae, ventral view, lightmicroscope with dig contrast, Tyumen (Siberia, Russia), 2016, copyrights Stefan F. Wirth
Two possible ways of virus transfer from bats to humans according to Andersen et al. (Nature Medicine, 2020)
Did the virus evolution towards the recent state happened prior to a first human infection, namely inside animal main host populations, or did it happen afterwards inside human populations?
As there is not yet much known about the presumed host specificity of the virus SARS-CoV-2, Andersen et al. (Nature Medicine, 2020) reconstructed based on their up-to-date knowledge two possible ways of a virus transfer from bats to humans and finally to the recent pandemic situation in the world:
The virus might have evolved its recent human-pathogenic features within the main host populations of bats. Natural selection must have been the corresponding major driving force. The relevant adaptations are represented by the above mentioned two molecular characters of the spike proteins in SARS-CoV-2 (receptor-binding domain for host cell binding and cleavage sites for an opening up of the virus). Under such circumstances the authors expect that the infection of humans could have happened with an immediate effect, leading at once into the pandemic situation of today. An intermediate host would in this option be not obligatory. A direct transfer from bats to humans might be imaginable.
The second option is based on findings that corona viruses in pangolins possess similar receptor-binding domains (RBD) as in the human SARS-CoV-2 version. Thus the authors reconstruct a version according to which a non or less pathogenic form of the new corona virus was via pangolins transferred to humans and circulated there for an unknown period of time. Even further possible intermediate hosts, such as ferrets or civets, are considered to have been involved in that scenario. During its time inside human populations the virus would have developed its recent features due to evolution and finally was able to be spread explosively between human populations on a pandemic level.
A higher probability for one of the two scenarios can according to the up-to-date knowledge not be assumed
I am not sure, whether the authors take under consideration with their second option that pangolins might even represent a main host and whether bats would not necessarily be involved in the animal-human transfer of the virus. But according to Ye Z.-W. et al. (Int Biol Sci, 2020) the context between bats, pangolins and humans was stated: „We cannot exclude the possibility that pangolin is one of the intermediate animal hosts of SARS-CoV-2“. But whether the pangolin is intermediate host or main host would at this point not effect the general conclusion of each of the two scenarios. The virus was either pre-adapted regarding efficient spike protein characters and then infested human populations rapidly or was transferred to humans via an animal host and subsequently evolved its key-features for a pandemic „success“ within human populations. Although the authors have up-to-date no indications allowing a preference for one of the scenarios, they point out that the potential of new SARS-CoV-2 outbreaks after the extinction of the recent human pandemic would be much higher in case of the scenario one, as the pathogenic virus would under these conditions survive in the animal main host populations.
I would as addition to scenario two suggest to test a modified hypothetic scenario, in which the non pathogenic ancestral version of the virus did not only circulate between human populations until it reached its pandemic key-features, but even circulated between humans and animal hosts forth and back for a longer time. This would according to my understanding of evolution improve the probability of a stepwise evolution of important key-features.
Special and unusual features of main hosts can improve the diversity within their parasites, important conditions for subsequent host changes: a very efficient immune system in bats pushes the evolution of their viruses
Chiroptera (bats and megabats) are not only known as putative main hosts for SARS-CoV-2, but also for Mers, Sars, Marburg and ebola viruses. Scientists did a research about the question, whether there are specific features existing, which explain, why Chiroptera are favorable hosts for viruses with a seemingly potential for epidemic and pandemic effects in human populations.
C. E. Brook et al. (eLife, 2020) discovered an unusual efficient immune system in Chiroptera, which they think protects these hosts from harmful diseases by their virus parasites. This bat immune system is considered being the evolutionary driving force for the variety of viruses and their relatively fast modifications, as they would need to compete with immune system responses by regularly evolving new adaptive features.
The authors discovered that the antiviral messenger substance interferone-alpha is released in most mammals as a response to the detection of viral genetic material inside body cells. Whereas they found Chiroptera releasing this messenger substance permanently. This would according to the scientists enhance the virus defense in bats and might explain that the above named viruses do not trigger noticeable diseases in their main host recervoir.
I would resume that such conditions might support the scenario one of Andersen et al. (Nature Medicine, 2020), according to which viral key features to infest humans had evolved prior inside the animal host populations. Regular new virus modifications as result of the competition between these viruses and their bat-host immune responses might support the randomness of the development of features, which as pre-adaptations could support a relatively fast host change. Even when I generally prefer scenarios of stepwise adaptations of organisms to new conditions, a higher probability of the availability of suitable pre-adaptations might at least accelerate evolutionary proceedings.
Longtime parasite – host – relationships, a dead-end for the parasite?
Are relationships between organisms over longer time periods of advantage or disadvantage for parasitic or mutualistic passengers? A longtime host specificity of a parasite (or mutualist) requires a strict specialisation, which means complex morphological, ecological and behavioral adaptations.
According to the acarologists P. B. Klimov & B. Oconnor (Systematic Biology, 2013) long-term specialisations could impede the flexibility of such organisms to react to environmental changes via evolutionary adjustments. Thus parasites with long-term relationships to the same hosts might be endangered to reach a dead-end. They would die out. A possible way out from such a disastrous end can be a re-evolution of the parasite back to its ancestral free living conditions, a situation prior to the evolution of its parasitic host specificity. But Dollo’s law states that a complex trait cannot re-evolve again. Thus long-term parasitism could according to the law not other than leading into a dead end. Nevertheless the authors could present an impressive example as proof to the contrary: based on their complex research about house dust mites, the acarologists reconstructed that these mites were originally parasites of warm blooded animals and subsequently evolved into free living associates of mammals, as which they are of medical relevance due to the remarkable allergic reactions in humans.
I think that the access of this paper does contain enough general biological aspects to ask, whether the dead-end scenario of long-term parasite relationships might also concern viruses, which don’t have an option for a free living existence, as they don’t live at all and are unable to perform independent strategies. At least might this long-term scenario support the findings of C. E. Brook et al. (eLife, 2020) that only unusual and regularly changing features of a long-term host might trigger regular corresponding responses by the parasite, another option to prevent a parasite from a dead-end due to a long-term host relation. This might explain, why certain viruses often parasite bats and successfully persist there, while other suitable hosts lack the very efficient immune system of bats and thus cannot host a specialized virus permanently. Regarding SARS-CoV-2 such theories might indicate that the virus would finally move towards dead-ends in humans and other host species, but might permanently survive in chiropterans. It’s a statement only being worth of consideration, in case of scenario one of Andersen et al. (Nature Medicine, 2020). And only in case, it would come out that the virus adapts well to humans, which would require a much reduced harmfulness, as parasites cannot survive by killing their hosts. In case of a dead-end host due to high mortality rates instead of a normal host increase, aspects of a long-term relationship with such a host don’t need to be discussed, as a shorter temporary outbreak and no beginning of a long-term relationship at all would result out of it. One needs additionally to consider that viruses as non living organic bodies with genome and with an unusual ability for fast modifications might often not fit into biological models based on living organisms.
House dust mite Dermatophagoides pteronyssinus. Wikipedia creative commons: Gilles San Martin from Namur, Belgium / CC BY-SA
Summary
Host specificity must be differed from generalism. Known host-parasite specializations include a complexity of strategies. And even different kinds of hosts must be named, such as main host, side-host, intermediate host or dead-end host. Evolutionary steps such as host increase, host change or temporary hosts can appear. Parasitism and mutualism differ from each other as life-strategies, but share common features as association between different organisms: host specificity follows similar rules, an indication that both life-modes can evolve out of each other. The human globalization sometimes supports the spreading of parasites or their hosts over the world, host changes or host increases can thus be performed including organisms, which would under normal conditions get no contact to each other.
Viruses do not represent living organisms, but protein lumps with a genome and depend on living host cells for their reproduction and „survival“. like in living organisms, also viruses underlay the mechanisms of natural selection and evolution. Viral parasite – host – relationships show general similarities with features in living organisms, including options for a host change or host increase, the use of intermediate hosts or a kind of mutualism (passenger viruses). There is evidence that the main host reservoir of SARS-CoV-2 are Chiroptera, while pangolins (and other mammals) might represent intermediate hosts. Humans are either dead-end hosts (preferred by most authors) or result of a successful host increase. Researchers could not yet decide, whether features to infest humans in a pandemic context evolved prior to the transfer to humans inside animal main host populations or whether a harmless version changed to humans and in their populations evolved its pandemic potential. A major drive motor for a long-term successful relationship with bats is the unusual immune system in chiropterans.
Copyrights Dr. Stefan F. Wirth (phd), all rights reserved, excluding photos labeled as creative common content from Wikipedia sources. Berlin, 2 April 2020
References:
J. R. H. Andrew’s (1983): the origin and evolution of host associations of Sarcoptes scabiei and the subfamily Sarcoptinae Murray. Acarologia XXIV, fasc. 1.
B. Berland (1961): Copepod Ommatokoita elongata (Grant) in the eyes of the Greenland Shark – a possible cause of mutual dependence. In: Nature, 191, S. 829–830. Cara E. Brook, M. Boots, K. Chandran, A. P. Dobson, C. Drosten, A. L. Graham, B. T. Grenfell, M. A. Müller, M. Ng, L-F. Wang, A. v. Leeuwen (2020): Accelerated viral dynamics in bat cell lines, with implications for zoonotic ermergence,eLife; 9:e48401.g W
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They look without magnification more like very motile and fast running ants or very tiny grasshoppers than like arachnids. But they indeed represent relatives of the web spiders and scorpions: Schizomida, a clade of whip scorpions. They are the sister taxon of Thelyphonida, the rather well known „big whip scorpions“, which are often kept as pets in terraria around the world. Schizomida are only rarely filmed in a higher resolution quality, which is due to their small size and their almost invisibility due to their semi-transparent cuticle and their very fast way of walking or even jumping. They are additionally difficult to be filmed as they strictly avoid all lights and tend to dry out quickly, when they cannot hide themselves by time in a slightly moist substrate.
Closeups of behaviors of a Schizomid species from a greenhouse in Germany. Copyrights Stefan F. Wirth
Schizomida in Greenhouses
Schizomids represent mostly tropical or subtropical organisms. But some species are regularly dispersed into greenhouses around the world. The filmed species might be Stenochrus portoricensis, but was not systematically studied in detail so far. As all known species, which appear in greenhouses, also S. portoricensis reproduces (apart from their original habitats) parthenogenetically with females producing females without mating procedures (thelytoky). I never found males so far.
S. portoricensis: native to subtropical Zones
The specimens, which I kept since months in a small terrarium, were collected in autumn 2016 at the famous fun and wellness bath „Tropical Islands“ South of Berlin. There they are a natural part of the world’s biggest indoor rainforest. The species S. portoricensis is originally native to Florida, Mexico, Cuba, Nicaragua, Porto Rico and other localities in similar tropical zones. These microscopical tiny organisms are predators and do not harm human beings at all. According to the available organisms in a suitable size in my terrarium, they might feed on the numerous collembolans and/or mites. Especially mites of the Gamasina appear in greater numbers in my substrate, which represents the original substrate from the greenhouse. I enriched this substrate regularly by smaller pieces of fruits or vegetables to stimulate the growth of microorganisms. I keep them at room temperature (about 20°C) and with not too much moisture. I do not know, whether they reproduced within these months, but the specimens of my recent video footage represent all sub-adults.
Film set and topic locomotion
Focus of my film is to present the different ways of locomotion, cleaning behaviors and burrowing activities of these fascinating animals. During the filming procedure, I used two cold-light-lamps for a suitable illumination and an ILCE-6300 (internal 4K mode), connected to a stereomicroscope and a lightmicroscope (with uplight).
Berlin December 2019/ March 2017, Copyrights Stefan F. Wirth
biologe 