In biodiversity research, knowledge of species numbers is the basis for planning environmental protection and climate research. However, the taxonomic work is made more difficult by cryptic species complexes in the world of organisms. Careless determinations of similar species must be prevented. For a beter understanding, examples from different animal groups are given. Using two species complexes of the mite taxon Histiostomatidae (Astigmata), two different forms of cryptic species complexes are presented in detail. Based on three species from a group associated with bark beetles, an example of a species complex is presented in detail, in which all stages of development look confusingly similar to one another. On the other hand, four species of mites from the bank mud of standing waters can only be confused with one another on the basis of their phoretic dispersal stage (deutonymph), while the adults differ distinctly. The meaning of such species complexes is discussed in the evolutionary and applied context. It is critically pointed out that too few specialists are funded worldwide and few taxonomists have to work too quickly, so that there is a risk of cryptic groups of species not being taken into account in surveys.
Biodiversity research is an essential fundament for disciplines like climate research and climate changes and thus contributes to an understanding about, how we humans need to treat our own environments. A main aspect of biodiversity research besides species monitoring is the evaluation of how many species we have. Specialists need to recognize and scientifically describe new species, especially, when it for example comes out that a complex of very similiar species contains more species than expected before (e.g. Laska et al. 2018). In tendency researchers in the field of biodiversity focus most on vertebrates in temperate regions and generally less in invertebrates (Titley et al. (2017).
The number of recently existing species in numerous cases is still unknown, especially in taxa of small organisms, such as mites. Due to a lack of specialists and due to a lack of fundamental research fundings, relatively much is known about direct pests of human sources, such as Varroa or Tetranychidae mites. But within the major clade Acariformes, ecological contexts and numbers and distribution of species of some free living taxa of Prostigmata and Oribatida/Astigmata are still an open field, even in Central Europe, e. g. Germany (Wirth, 2004).
This is despite the fact that for example phoretic mites, which use other arthropods as carriers for dispersal, can have highly complex relationships with their phoretic hosts, thus being of interest from the evolutionary, the ecological and even an applied point of view. The latter is discussed for example in context with different bark beetles, which their mites might affect by acting as vectors for fungus spores (Klimov & Khaustov, 2018).
Cryptic species complexes are a topic that is currently being widely dealt with in science. Such species complexes are characterized by the fact that they are difficult or impossible to distinguish morphologically. However, they can be clearly differentiated from one another using barcoding (e.g. Kameda et al, 2007), behavioral or ecological studies. Crossing experiments are a frequently used ecological method. Because according to the biological species concept, individuals of different species either cannot be crossed with one another or the offspring of such a hybridization is not fertile (e.g. Sudhaus & Kiontke, 2007).
Crossing experiments are particularly suitable for the investigation of cryptic species complexes in species that have a rapid life cycle and, due to their small size, can be accommodated well in standardized conditions. Such organisms are, for example, free-living nematodes of the Rhabditidae (e. g. Sudhaus & Kiontke, 2007) or mites of the Histiostomatidae (e.g. Wirth, 2004).
The cryptospecies phenomenon, which means that closer investigations show that animals once attributed to the same species actually represent several species, can in principle occur in the entire animal kingdom and in plants and fungi too (Shneyer & Kotseruba, 2015). Previously known subspecies are often given their own species status as a result. One example are the two monitor lizard species Varanus niloticus and V. ornatus (e. g. Böhme & Ziegler, 2004).
In this monitor lizard research mainly ecological differences to V. niloticus have been studied. As one of the results, V. ornatus does not have a diapause in summer, which is a distinct difference to V. niloticus (Böhme & Ziegler, 2004). As an unusual phenomenon, a case of parthenogenesis was even observed in V. ornatus, but not in V. niloticus (Hennessy, 2010) so far. However, morphological differences between these two monitor lizards were known even before, for example relating to aspects of the dorsal drawing. But the authors named above were able to provide evidence that these morphological differences do not occur gradually, as orgininally assumed, but rather distinctly.
Another example of two sibling species (the most simple form of cryptic groups) that have been identified as different species by molecular biological studies are Homo sapiens and H. neanderthalensis (e.g. Prüfer et al., 2014). Originally it was assumed that H. neanderthalensis was a subspecies of H. sapiens. This is for example supported by the proven cultural exchange between the two species and the great morphological similarity. In the meantime, however, morphological findings such as the morphology of the nasal duct of the Neanderthal man have also supported the genetic findings (Márquez et al., 2014). However, very recent studies show that Neanderthal genetics have entered the lines of H. sapiens (Hajdinjak er al., 2021). As a result, both forms have crossed and produced fertile offspring. It remains to be seen whether this will possibly dismiss the concept of two species again.
Since the aim of all studies of cryptic species complexes is to find distinctive differences in the areas of morphology, ecology or barcoding (or all approaches together) that distinguish one species from all others, ultimately clearly definable, very closely related species remain in case of successful studies.
If the cryptic organisms are members of an organism-socialization, such as parasites and their hosts, the idea that a proven host specificity can be an indicator for a certain species of a cryptic complex is obvious. In fact, Wirth et al. (2016) for example postulated a host specificity for the phoretic mite Histiostoma piceae and its hosts, the bark beetles Ips typographus and I. cembrae. Nevertheless, relationships between associated species are usually not studied extensively enough to be able to unequivocally identify certain species on the basis of for example their hosts (Wirth, 2004).
Since cryptic species represent nevertheless separate species despite their extraordinary similarity, they are subject to the species concepts. As a result, they form different niches and can therefore appear sympatric in the same living space (e. g. McBride et al., 2009). This makes it difficult for biodiversity researchers and systematics to investigate the real numbers of species in such habitats.
If, instead, cryptic species are not sympatric, but distributed in adjacent areas, this can for example indicate that an allopatric species formation has either not been completed for a long time or is even still in the process of speciation (e. g. Gollmann, 1984).
Animal species that have different developmental stages can appear cryptic, i.e. being morphologically confusingly similar, with regard to all these developmental stages, such as for example certain phoretic free-living nematodes, which then additionally have to be studied ecologically or genetically (e. g. Derycke et al. 2008).
Other species can hardly be distinguished morphologically with regard to a certain developmental stage, which is particularly common, but differ distinctly in other developmental stages, which are more difficult to find. Very similar looking lepidopteran caterpillars of sibling species (e. g. Scheffers et al. 2012) can be more commonly available than their adults, which might be easier to distinguish.
As a specialist for mites of the family Histiostomatidae (Astigmata, Acariformes) I will in my further argumentation refer to my biodiversity studies on these mites and explain the difficult situation for describers of new species based on several specific histiostomatid species, some being phoretically associated with bark beetles and others associated with different coleopterans from muddy sapropel-habitats around ponds in Berlin/Germany. In connection with these cryptic groups of species, reference should be made to the applied difficulties in connection with biodiversity research. I am referring to the fact that, for a variety of reasons, often only a certain juvenile stage (deutonymph) is used for species descriptions (e. g. Klimov & Khaustov, 2018 B), although cryptic species can occur sympatricly in the same habitat and in many cases not be sufficiently differentiated from one another on the basis of just this one stage.
In Histiostomatidae as in most Astigmata taxa, the deutonymph (in older publications hypopus) represents the phoront, being adapted morphologically and behaviorally in getting dispersed by insects or other arthropods. This instar has no functional mouth, possesses a ventral suckerplate to attach to its carriers and a thicker sclerotization against dehydration. The deutonymph is often collected together with its phoretic host. Bark beetle traps are for example a common source, where dead deutonymphs still on their hosts come from and are subsequently forwarded to acarologists, who then are of course unable to create a mite culture in order to have also adult instars available for species descriptions (e. g. Klimov & Khaustov, 2018 B) and other taxonomic purposes. This paper shall clarify, why it is instead necessary for a clear species determination to have the deutonymph and additionally at least adults available.
In this publication two cryptic species complexes from the taxon Histiostomatidae (Astigmata) are presented as result of my original scientific work. On the one hand morphologically very similar representatives of the Histiostoma piceae-group, which are originally associated with bark beetles (Scolytinae), on the other hand similar looking representatives, which are bound to insects in the area of the banks of ponds with digested sludge (sapropel). It needs to be emphasized in that context that those herewith introduced two cryptic clades are phylogenetically not closer related to each other.
The presented bark beetle mites (chapter 1 in results) can only be distinguished morphologically by very gradual characteristics, in terms of phoretic deutonymphs as well as in terms of adults. However, there is a tendency towards host specificity (e.g. Scheucher, 1957), which is why there could be a permanent spatial separation of the species despite common occurrence in the same region.
The mites from the sapropel in the area of the pond banks (chapter 2 in results) are presented on the basis of a certain area in Berlin (Germany), where they appeared sympatric. Unlike the bark beetle mites, they are morphologically clearly distinguishable with regard to the adults, but have morphologically very similar deutonymphs, which essentially only differ from one another in degrees.
Based on the representatives of two different cryptic species groups presented in this work, it should be shown that a sufficient range of morphological features for systematic and taxonomic differentiation and characterization of species can only be available if at least two developmental stages of a population can be studied. It is also pointed out that high-resolution optical methods can uncover a possibly systematically relevant variety of morphological features that would otherwise remain hidden. It is suggested that a suspected host specificity cannot always be used to differentiate between very similar species and that cryptic species can be found sympatricly on the same host as well as in the same habitat. The main aim is to show that there is a risk of confusion and a risk of underestimating the existing biodiversity if only the deutonymph is used for taxonomic purposes, just because it is for example easily available, when the host is captured. Nevertheless species descriptions based only on the deutonymphs are unfortunately still surprisingly common.
Due to the lack of sufficient research fundings and a corresponding decrease of experienced specialists, trends to remarkably simplify determinations and species descriptions are about to manifest themselves. Non specialists or less experienced acarologists increasingly try to recognize or describe new species based on the availability of deutonymphs only, because these phoronts are often easily accessible as bycatch of entomological material. It is mistakenly assumed that faster procedures could accelerate the level of scientific knowledge about the biodiversity of astigmatid mites (Wirth, 2004).
Material and Methods
Chapter 1 is an illustration of the current state of my research about a cryptic bark beetle-associated group of species. Problems and questions are additionally shown both on the basis of existing, in part own, literature. Chapter 2 is about four species of Histiostomatidae that were recorded from an old gavelpit area in the urban Berlin forest Grunewald, named „Im Jagen 86“, located 52° 29′ N, 13° 14′ E. This chapter focuses specifically on Histiostoma maritimum, collected between 2002 and 2012 (and also between 1999 and 2000 during my diploma thesis). Besides H. maritimum three other species were found in the same area and habitat: Histiostoma palustre, collected once via deutonymphs from a beetle of Genus Cercyon in 2002 and reared in culture over about two years on moist decomposing potato pieces, Histiostoma litorale, isolated as adults from sapropel mud once in 2002 and Histiostoma n. sp., reared only one generation long from adults to adults in 2019, inside sapropel-mud samples with moss growth and moist decomposing potato pieces.
Mites of H. maritimum were collected as deutonymphs on the beetles Heterocerus fenestratus (rarer on Heterocerus fusculus) and Elaphrus cupreus from sapropel around two ponds in the named area. After different experiments, mites developed successfully on beetle cadavers on 1.5 % water agar in Petri dishes (diameter 5 cm) at room temperature (ca 20°C, summer 2002). Three cultures (one cadaver of C. elaphrus and twice each time two cadavers of H. fenestratus) were observed over a period of about three weeks (additionally small pieces of beef heart were added to all these cultures to maintain suitable food sources). Adult mites were stored in 80 % ethanol for about 5 days and then critical point dried for SEM studies. Photos were taken by an analogous medium size camera via a Philips SEM 515 and later developed. Still unpublished copies from 2002 were scanned in a high 600 dpi solution and as tiffs via a CanoScan Lide 2010 in 2021. Restauration and picture quality improvement were performed via Adobe Lightroom. The areal panorama of the former multiple pond area was captured in September 2018 via a Dji Mavic Pro drone at a height between 30 and 50 m and subsequently modified into black and white.
Setal nomenclature follows Griffiths et al. (1990).
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.
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
Pavel B. Klimov, Barry OConnor, Is Permanent Parasitism Reversible? (2013): —Critical Evidence from Early Evolution of House Dust Mites, Systematic Biology, Volume 62, Issue 3, Pages 411–423.
Kristian G. Andersen, Andrew Rambaut, W. Ian Lipkin, Edward C. Holmes, Robert F. Garry (2020): The proximal origin of SARS-CoV-2. Nature Medicine. Martin Schaller: Demodex-Follikulitis. In: Gerd Plewig, Peter Kaudewitz, Christian A. Sander (Hrsg.): Fortschritte der praktischen Dermatologie und Venerologie 2004. Vorträge und Dia-Klinik der 19. Fortbildungswoche 2004. Fortbildungswoche für Praktische Dermatologie und Venerologie e.V. c/o Klinik und Poliklinik für Dermatologie und Allergologie LMU München in Verbindung mit dem Berufsverband der Deutschen Dermatologen e.V. (= Fortschritte der praktischen Dermatologie und Venerologie. 19). Springer Berlin, Berlin 2005, ISBN 3-540-21055-5, S. 273–276.
Wirth S. (2004): Phylogeny, biology and character transformations of the Histiostomatidae (Acari, Astigmata). phd thesis. Internet Publikation FU Berlin, http://www.diss.fu-berlin.de/2004/312.
Wirth, S. & Moser, J.C. (2008): Interactions of histiostomatid mites (Astigmata) and leafcutting ants. In: M. Bertrand, S. Kreiter, K.D. McCoy, A. Migeon, M. Navajas, M.-S. Tixier, L. Vial (Eds.), Integrative Acarology. Proceedings of the 6th Congress of the European Association of Acarologists: 378-384; EURAAC 2008, Montpellier, France.
Wirth S. & Moser J. C. (2010): Histiostoma blomquisti N. SP. (Acari: Histiostomatidae) A phoretic mite of the Red Imported Fire Ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Acarologia 50(3): 357-371.
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I was part as acarologist and natural scientist in a 2011 scientific paper about a mite preserved as fossil in amber, which was analyzed using the X-ray computed tomography and determined systematically on a family level. In this time, this scientific publication had a remarkable impact in international scientific media, because it seemed, as if this mite was the smallest animal ever visualized via CT on a high quality level.
Strange behaviors of so called „colleagues“?
The technical work was performed by technical scientists in Manchester UK. The natural scientific analyses was performed by me as the only European specialist for the mite family Histiostomatidae. But I noticed already in the time period of this publication that there were strict tendencies by the so called „colleagues“ to mention my name as less as possible, this concerned the drafting of international media releases and also a poster presentation (my name was added days later) and an online abstract on a conference in Berlin. The corresponding poster was even awarded, but I got my award certification only after demanding explicitly for it. I much later, when I decided to complain officially at the Museum of Natural Sciences in Berlin, needed to learn that I was not even considered as one of the first authors. I didn’t notice that before, because the former „colleague“, Dr. Jason Dunlop, curator at this museum, was mentioned in the original citation with 1) after his name, me too. Thus I interpreted this as a double-first-author-ship. It then came out that the „1)“ only indicated the same scientific address, because I was in that time officially a volunteer at the MFN in Berlin.
Mite in an amber fossil, made visible by using the x-ray computed tomography, acarological work: Stefan F. Wirth
The work of a scientific specialist: here an acarologist
The question must be: Who is needed to scientifically interpret three dimensional photos of an amber fossil, in this case the deutonymph of a mite of the Histiostomatidae? A specialist for this taxon is needed, who is able to perform scientific drawings, based on the photos. He first needs even to decide, which of the photos are showing details of scientific relevance. While drawing, the specialist must distinctly recognize single microscopic structures, so that all these structures can be clearly separated from each other including all borders or gaps between single components. The scientific term is „homologisation“. Homologisation means: comparing single structures with (phylogenetically) equivalent structures of other (related) species. As there were not more fossils available, the homologisations needed to be based on recent mites. Thus the specialist must have a very competent knowledge of a high number of species from this family. To reach that level requires hard work over many years. I had the necessary level and found character details in the fossil, which were fitting to recent members of mites of the Histiostomatidae. But it’s of course not enough to discover such homologous structures. They must be made visible for every reader of the scientific paper. Thus the drawings need to be correctly labelled, which requires careful morphological studies. Then a detailed description needs to be written. But that is far not enough. Readers of a scientific paper are usually no specialists. That’s why they need a written introduction, in which the summary of the general recent knowledge of a mite group needs to be presented. And after all that they even expect you to discuss your results. It’s an own chapter, subsequent to the result descriptions.
The discussion chapter also requires a maximum of specialized competence. Some researchers even say that this is the first part of a paper that they read as it puts the results into a general scientific context based on arguments, mostly according to the principle of the most economical explication. Conclusions in the discussion part have usually the character of theories based on the facts, which the paper could contribute. Topics of a discussion part in such a paper as ours are systematic conclusions, the discussing of homologisation problems and also the formulation of a possible relevance for the recent scientific knowledge and also the future scientific importance of these new findings.
This all is, what I as a specialist needed to do. I additionally contributed one of my photos of a recent mite for comparative reasons and captured a stereomicroscopic photo of the mite fossil to demonstrate, how much the CT could improve the visible details of the amber fossil. I guess I did quite a lot, the other part was overtaken by the technical colleagues in Manchester. They needed to explain their technical situation and also needed to discuss their ideas about the meaning of their CT-technology for the future of science, focussed also on work with amber fossils.
Contributions of different authors to a scientific paper
To be honest I don’t remember, where there was still space left for content issue contributions by Dr. Dunlop. But he did some organizational stuff, he collected the contributions from the UK colleagues and me, he arranged the photo table via a graphic software based on the photos, which I had determined as scientifically relevant, and he was the so called corresponding author (I allowed him, because he is an English native speaker). That means, he submitted the final paper to the journal and communicated with the editors. Of course reviewers always ask for revisions. That was then mine and the technicians job again.
It is common that corresponding authors represent automatically the first authors of a paper. But it is not mandatory. I for example once was the corresponding author of a paper, which was based on a bachelor thesis that I (in major parts) supervised. I despite of my in fact major authorship regarding the scientific paper itself and my additional corresponding activities let her (the student) the first authorship. That even means that this paper can be easier found, when searching for her instead of my name. I just wanted to support a younger scientist.
And of course also a double first-authorship might be possible, especially representing an adequate solution, in case another author even contributed more concerning the scientific content itself. In case of objections by the editors, the one, who contributed more, should to be the first author.
„B-word“?
But to come back to the amber paper of this article, it is surely not fair to reduce the scientist, who had the major scientific work on a paper secretly to a second author. It is highly unfair to leave him out in the international press release information. And I don’t trust to say here, what it is, when deleting his name entirely from a poster and an online abstract presentation and even impeding him to get a certification of a poster award in time for his work. Should one use the „b-word“? Generally bullying would be an act against the good scientific practice, but there would be clear proofs for malevolence against specifically somebody needed to get corresponding behaviors sanctioned. But when „only“ the elbow mentality is obvious, which means that people leave somebody out for their own better recognition, then the distinct malevolence against the victim is not clearly proven. Thus the interesting question arises: when is elbow behavior equal to bullying and when not?
Warning to young scientists
What I can say for sure is, even when the original bullying assumption is still a kind of questionable: after you complained, you might need to expect a real merciless and long lasting bullying. That’s why I intend to warn all young scientists: be careful and double check, with whom you cooperate. The wrong choice can be a failure as long as you do not agree being a bullying victim. The consequences can last over years and can destroy your whole career. I even once was told by a bullying victim that the accused institution did not even deny its bullying activities, but stated that depending of the kind of position, somebody has in an institute, an equality right would not be automatically existent. I go further and say: don’t become a natural scientist at all, except you are in a love relationship with an internationally highly influential professor.
In these days there are alternatives for possible natural scientists. Earlier I was a harsh critic of the modern gender sciences (sometimes also named genderism). But they have much financial capacities. Nobody there needs to sharpen his elbows, a good basis for fair careers, and based on that after a while surely also the most important basis for a good quality work!
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
Gravel pit area „Im Jagen 86“ in Berlin as biotope
„Im Jagen 86“ is a former gravel pit area in the Berlin urban forest Grunewald. It today represents a dynamic biotope, consisting of different types of habitats: mud around ponds, sand dunes, dry grassland and forest. Since the early 2000th, its habitat composition partly changed remarkably. Out of several (smaller) ponds, only one bigger pond remained. All ponds originally were surrounded by sapropel, a habitat for different interesting organisms, such as beetles of Heterocerus, Elaphrus and Bembidion. The mite Histiostoma maritimum was commonly found phoreticaly on Heterocerus and Elaphrus. I additionally in those early 2000th described the new mite Histiostoma palustre from Hydrophilidae of Cercyon and Coelostoma, living inside the saporopel as well. Today only a few small areas with open sapropel exist. I so far did not look for Histiostoma maritimum again and don’t know, how common it still is. At least Heterocerus beetles are harder to find than in earlier years. I so far did not found Histiostoma palustre again.
Rearing conditions of a putatively new mite species
I collected new mud samples in March 2019 at different areas, but found developing histiostomatid mites in a sample from the edge between mud (sapropel) and mosses. It is a species I never found before there and which might represent a new species. Only females could be morphologically studied. Nymyphal stages (not deutonymphs) are only available as video footage. No males were found. I had added bigger potato pieces to stimulate microorganism growth as mite food into the soil sample (room temperature). After about one month, a few mites (females and proto/tritonymphs) developed on only one of these potato pieces and quickly died out shortly after my filming activities and after I could prepare a few females. I actually try to get them reared again. Due to the low temperatures in March, it is considered that these mites hibernate independently from insects in the substrate. No bigger insects could be found in the substrate, which might be the corresponding carriers. But different dipterans (e.g. Ceratopogonidae) developed, they had no mite deutonymphs after hatching in my sample.
Morphological reconstruction of females and important characters as well as behavioral observations
The females of Histiostoma sp. differ from other females, which I know, by the mosaic of the following characters: body conspicuously elongated with a distinctly big distance between hind ringorgans and anus, digitus fixus almost simple shaped, fringes or ridges on palparmembrane, 6 dorsal humps, unusually big copulation opening. Leg setation not yet studied. One pair of ventral setae hardly visible (not in the drawing). Nymphs were observed during burrowing activities (footage), females are may be also able to. Deutonymphs or males would be useful to decide, whether the species is new. Some species are only described by deutonymphs.
Berlin, March/ June 2019 All copyrights Stefan F. Wirth
Some animals live in environments, where there is (almost) no light available. It makes no sense to see in the dark, but it is important for a specimen to know, where it actually is, where it is going to, whether there is enough food and what the conspecifics are doing. Predators need to be recognized in time, and a sexual partner must be found. There is also need for an efficient communication between specimens of a species. How can all this be performed by mites of the Astigmata, which usually live inside decomposing soil habitats in a more or less permanent darkness?
Olfactory sense organs in mites of the Histiostomatidae
Histiostoma sachsi (Histiostomatidae, Astigmata) is such a mite, living inside cow dung or compost. It might have a rudimentary ability for a light perception, but has not visible or functional eyes. It cannot produce any sounds. It can only feel and smell. Seemingly very limited abilities, but the contrary is fact: Due to evolution this mite is perfectly adapted to its life-style. It can feel objects by touching on them using its body setation (= body hairs). And it smells by means of very specialized body hairs, which are called solenidia and appear in different types, shapes and functions. These mites don’t smell on the level of us humans, which would be very insufficient. If at all, it should be compared with a dog. I am always fascinated when seeing blind dogs and how perfectly they can interact with their environment, despite their handicap. That’s may be how the efficiency of olfactory perception abilities of such a mite must be imagined. They do not only perceive scent particles from other animals, plants and soil components. Even olfactory signals from their conspecifics will be correctly and differentiatedly interpreted. And that not only marginally. Olfactory signals represent indeed the major mode of their intraspecific communication.
Chemical communication of mites of the Histiostomatidae
Communication always requires contributions from both sides, a signal and an answer. These mites smell the signal of a conspecific using their solenidia, and they answer by the secretion of biochemical components. For these purposes, they possess a huge and complex gland system located on the upperside of their backs. Volatile excretions aggregate inside a big and rounded reservoir and finally leak to the outside via a pore, called oilgland opening. These gland systems are located symmetrically on both sides, each with one reservoir and one pore.
The meaning of the sent volatile message simply depends on the composition of the correspondingbiochemical components. Even diffferent stereochemical configurations of the same molecule can have different meanings. Citral for instance is a major component and has in different stereoisomers different functions. Such cummunicative volatile signals are usually named pheromones. And mites of the Histiostomatidae can indeed produce different kinds of pheromnes via the same gland system. Aggregation pheromones inform specimens about a suitable place to stay together with their conspecifics, for example due to a sufficient amount of food resources. Alarm pheromones solicit mites nearby to flee from an unpleasant situation. Sexual pheromones attract adult partners to each other in order to perform the mating procedure. But the gland secretions can even more. As allomones, they communicate with specimens of other species. They function as defenses against predators or other dangerous cohabitants.
Deutonymphs need to find a carrier for dispersal
Another form of communicative interspecific interactions is performed by a specific juvenile instar, the deutonymph. It looks morphologically quite different from all other instars (heteromorphic situation), does not need or possess a functional mouth, has a thicker cuticle as protection against drying out and a complex sucker organ on its underside in order to attach itself to an insect or another bigger arthropod. Deutonymphs of the astigmatid mites search for bigger carrier-arthropods to get carried from one habitat to another (dispersal strategy is calledphoresy). While doing so, they again use their specifically modified leg setation (hairs) on the first pairs of legs to perceive scents for the detection of a suitable and passing by carrier. Basically it is still unknown, whether the term „communication“ is indeed appropriate in this context as we don’t know yet about a mutual interaction between deutonymphs and their carriers, before the phoretic ride begins.
Olfactory orientation of the deutonymph of Histiostoma sachsi, copyrights Stefan F. Wirth, February 2019.
Specific way of walking in deutonymphs
In detail, different kinds of behaviors can be observed in deutonymphs, when searching a carrier. The detailed behavioral patterns in this context can slightly differ between even closer related species. Deutonymphs of Histiostoma sachsi as all deutonymphs show a characteristic mode of walking, in which especially the first pair of legs plays an important role. During each step, performed by four pairs of legs, the first legs are lifted up much higher than all other hind legs. While doing so, they slightly tremble up and down. A behavior that mostly supports a better basic orientation inside a „jungle-„micro-landscape, being filled up with soil particles and decomposing plant tissues. But what H. sachsi deutonymphs additionally need in order to find their carriers is repeatedly to rest between the walking activities. Thus the first legs, which normally are still walking legs, are made free and that way available for the perception of carrier-scent-components only. These namely are the legs that bear the highest densiy of solenidia.
Two different behavioral modes for an efficient orientation towards a carrier
Two different modes of resting with olfactory searching activities could be observed: In periodic intervals the deutonymph attached to the ground by using its sucking structures. They were then more or less laying on their entire undersides with only their forebodies slightly lifted up. By alternating moving the first legs up and down, olfactory information could be perceived from all directions without having the own body as a barrier to backwards. To improve its orientation situation, the deutonymph additionally turned on its own axis around, being stabilized by its sucking structures, which are flexible enough to follow these movements. When the deutonymph intended to continue its walk, it first needed to detach from the ground, which happened via muscle contractions that caused an abrupt detachment of the corresponding suckers. But main aim of the deutonymph is to find an elevated place, where the probability of a passing by carrier is especially high and from where a bigger insect (or other arthropod) can easier be ascended. There the second behavioral mode was performed. The deutonymph only fixed the edge of its hind body to the ground, again using the suckers on its underside, which are located close to this edge. This time the entire mite body stood in an upright position. The first legs again „waved“ alternating up and down and could under these especially elevated conditions even perceive scents from bigger distances. By occasionally slightly and alternating turning their upright bodies to both sides, olfactory information could be easier detected from all directions.
Carrier of H. sachsi still unknown
The frequency of such movements in mites increases typically as closer a suitable carrier approaches. But this was not yet observed or documented for Histiostoma sachsi. Its carrier inside the compost substrate is still unknown, which is why I so far could’t perform corresponding experiments. The species‘ describer, Scheucher (1957), found her mite specimens in cow dung and also didn’t identify the corresponding carriers there.
The observations presented in my video are part of my research project about morphologies and behaviors of deutonymphs in the Histiostomatidae.
Berlin, February 2019. All copyrights Stefan F. Wirth.
The city of Berlin geomorphologically consists of witnesses of the Weichselian glacier. The modern city itself and adjacent federal states represented end moraine areas with fluvio-glacial debris accumulations, even well visible today due to a very sandy soil composition and a corresponding vegetation, creating landscapes, which partly almost look like from around the Mediterranean Sea.
Sands carried by the glaciers towards their end positions remained in partly huge layers with a thickness of up to 20 meters or more.
Gravelpit zone and its history
Also the area of the old gravelpit zone, called „Sandgrube im Jagen 86“, in the Berlin forest Grunewald is located inside such an end moraine zone, which was represented by plates belonging to the geological Teltow-plateau. In the time period between 1966 and 1983, gravel was excavated for industrial purposes. After 1983 a part renaturation was supported by nature conservationists. In 1992 in total 13 hectares of the former gravelpit area were allocated as nature conservation areas.
Other parts of this unique landscape remained accessible for the public. They represent today popular places for leisure and experiences of nature. Especially the huge sand dune is a popular destination for families with children.
Aerial videography of the gravelpit area in January 2019, copyrights Stefan F. Wirth. Please like my video also on Youtube, in case you like it.
Gravelpit zone and its ecology and biodiversity
The whole area – nature protection and accessible zones – show a complex mosaic of different landscape types, offering numerous animal and plant species a well suitable refuge. Neglected grasslands and dry meadows are surrounded by sandy areas free of any vegetation („dunes“) and moist osier beds and wetlands with ponds. The wetlands represent breeding grounds for numerous amphids. Lizards such as the sand lizard Lacerta agilis and snakes such as the grass snake Natrix natrix can regularly be observed. Sandy habitats offer space and specific ecological conditions for a interstitial fauna, consisting for example of different bee and sand wasp species.
In total the area bears more than 300 ferns and flowering plants, 16 breeding bird species, 7 amphibian species and 188 butterfly species.
My own scientific mite research in the gravelpit area
I was performing scientific research in that gravel pit landscape during the work on my phd-thesis between 2000 and 2005. My interest was (and one of my interests is still) focussed on specific organisms living around the shoreline of ponds.
The whole area of the gravelpit landscape is a good example for ecological changes that happen naturally with the ongoing time or even being affected by climatic changes. Between 2005 and 2018, the landscape partly changed significantly. Neglected grasslands and dry meadows covered less space originally, and instead several smaller ponds existed and offered amphibs and wetland inhabiting insects additional habitats. But some of the ponds already years ago dried out permanently. Their remnants are now covered by extended dry grasslands.
In former times of my phd thesis and even today, my research interests focus and focussed on the mite fauna in and around the muddy shorelines of ponds inside this former gravelpit area. The ponds are mostly surrounded by sapropel, a seemingly black and brownish mud, which is colored that way due to the incorporation metal sulfides. These muddy areas develop due to biochemical modifications of organic material in the absence of oxygen. Different insects, especially beetles live on top of these waterside habitats or even inside. Carabids of genera Elaphrus or Bembidion represent predators, while heterocerid beetles of genus Heterocerus are substrate feeders, presumanly with a preference for diatoms. Also water beetles of Dytiscidae and Hydrophilidae inhabit these habitats.
The mites Histiostoma maritimum and Histiostoma palustre
I discovered some of these beetles as dispersal carriers for specific mites. The dispersal strategy to take a ride on bigger animals to become carried from one habitat to another is called phoresy. Mites of the Astigmata represent typical phoretic organisms. I am scientifically specialized in a specific family of the Astigmata, which is named Histiostomatidae, and I discovered the mite species Histiostoma maritimum Oudemans, 1914 on Heterocerus fenestratus and H. fusculus as well as on Bembidion and Elaphrus species insside and on top of these muddy zones. I was the first acarologist, who ever studied the biology of this mite species. I furthermore discovered another mite species that was completely new to the scientific knowledge, and thus I scientifically described it as Histiostoma palustre („palustris“ = „muddy“) in 2002.
This species deserves particularly mention due to an unusual biological phenomenon: populations show a so called male dimorphism (better diphenism). Besides males with a „normal“ morphology, morphologically modified males appear. Their second legs differ from the typical shape of a mite and are modified into clasping organs. The function of these conspicuous organs could so far only be interpreted in the context of male to male competition conflicts for a female. In such situations, I observed the organs being used as arms against other males, against such ones with and such ones without clasping organs.
Right modified leg of a male of Histiostoma palustre. Copyrights Stefan F. Wirth, 2002/ 2019
Modified leg of a H. palustre male in closed position. Copyrights Stefan F. Wirth, Berlin 2002/ 2019
Underside of a H. palustre male with modified leg. Copyrights Stefan F. Wirth 2002/ 2019
Asymmetry: male of H. palustre with only the right leg modified. Copyrights Stefan F. Wirth 2002/ 2019
Asymmetry: male of H. palustre with only the left leg modified. Copyrights Stefan F. Wirth 2002/ 2019
Copulation of a Histiostoma palustre male with both-sided modified legs. Copyrights Stefan F. Wirth, Berlin 2002/ 2019
Details of a copulation with a modified male, copyrights Stefan F. Wirth, 2002/2019
biologe 