biologe

Blog and online journal with editorial content about science, art and nature.

Tag: Histiostoma palustre

Two different forms of cryptic species-complexes in mites of the Histiostomatidae (Astigmata) from bank mud and bark beetle-galleries and their significance for applied biodiversity research

Biologe ISSN 2750-4158

Stefan F. Wirth, acarologist, freelancer, Berlin, Germany

Citation: WIRTH S. F. (2021): Two different forms of cryptic species-complexes in mites of the Histiostomatidae (Astigmata) from bank mud and bark beetle-galleries and their significance for applied biodiversity research. Biologe (ed. Stefan F. Wirth), category : original scientific papers volume 1 (2021; 2022) , 1-7. URL: https://biologe.wordpress.com/2021/12/31/two-different-forms-of-cryptic-species-complexes-in-mites-of-the-histiostomatidae-astigmata-from-bank-mud-and-bark-beetle-galleries-and-their-significance-for-applied-biodiversity-research

Abstract

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.


Keywords: cryptic species groups, evolution, biodiversity research, Acariformes, Histiostomatidae, Astigmata, phoresy, Histiostoma piceae, Histiostoma Scheucherae, Histiostoma piceae, Histiostoma ulmi, Histiostoma palustre, Histiostoma litorale, male morphology, SEM, Histiostoma maritimum, Scolytinae, Carabidae, sapropel


Introduction


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).



Results:

Seiten: 1 2 3 4 5 6 7

Mite Histiostoma sp., putatively new species, from mud around ponds (Berlin) and its morphology

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

Berlin forest Grunewald – former gravelpit area, type location for the mite Histiostoma palustre

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.

 

img_0015.jpgbest

Right modified leg of a male of Histiostoma palustre. Copyrights Stefan F. Wirth, 2002/ 2019

 

img_0016best

Modified leg of a H. palustre male in closed position. Copyrights Stefan F. Wirth, Berlin 2002/ 2019

 

img_0017.jpgbest

Underside of a H. palustre male with modified leg. Copyrights Stefan F. Wirth 2002/ 2019

 

img_0013.jpgbest

Asymmetry: male of H. palustre with only the right leg modified. Copyrights Stefan F. Wirth 2002/ 2019

 

img_0012.jpg best

Asymmetry: male of H. palustre with only the left leg modified. Copyrights Stefan F. Wirth 2002/ 2019

 

img_0014.jpgbest

Copulation of a Histiostoma palustre male with both-sided modified legs. Copyrights Stefan F. Wirth, Berlin 2002/ 2019

 

img_0010best

Details of a copulation with a modified male, copyrights Stefan F. Wirth, 2002/2019

 

 

Berlin, January 2019. Copyrights Stefan F. Wirth