# A new species Zachvatkinibates svanhovdi sp. nov. (Acari: Oribatida: Punctoribatidae) from Norway with comments on Punctoribatidae in Fennoscandia

Anna Seniczak, Stanislaw Seniczak, Snorre B Hagen, Cornelya F C Klütsch

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Anna Seniczak, Stanislaw Seniczak, Snorre B Hagen, Cornelya F C Klütsch. A new species Zachvatkinibates svanhovdi sp. nov. (Acari: Oribatida: Punctoribatidae) from Norway with comments on Punctoribatidae in Fennoscandia. Acarologia, 2023, 63 (1), pp.41-57. 10.24349/skz7-19nt. hal-03943670

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# A new species *Zachvatkinibates svanhovdi* sp. nov. (Acari: Oribatida: Punctoribatidae) from Norway with comments on Punctoribatidae in Fennoscandia

Anna Seniczak IDª, Stanisław Seniczak IDᵇ, Snorre B. Hagenᶜ, Cornelya F. C. Klütschᶜ

## Original research

### ABSTRACT

The adults of the new species *Zachvatkinibates svanhovdi* A. Seniczak & S. Seniczak sp. nov. are described and illustrated from Norway, and this is the first finding of *Zachvatkinibates* Shaldybina, 1973 in Fennoscandia. This species is the most similar to *Z. quadrivertex* (Halbert, 1920), but differs from it mainly by the shape of notogastral setae, posterior tectum of notogaster and lack of postanal porose area *Ap*, which in *Z. quadrivertex* is present. In *Z. svanhovdi*, the prodorsal seta *in* is long, translamella is narrow, notogastral setae are short and distally pliable, notogastral porose areas are usually oval and of medium size, but *Aa* can be larger, especially in males. Dorsal crest on tarsus I is present. The cytochrome oxidase I (COI) barcodes (length: 658 bp) of five specimens of the new species are provided; the maximum variation within the species was 2.41% (p-dist). The morphology and ecology of the new species is compared with other *Zachvatkinibates* species. The knowledge on family Punctoribatidae in Fennoscandia is updated, and *Mycobates carli* (Schweizer, 1922) is first reported from Norway.

**Keywords** oribatid mites; sexual dimorphism; leg setation; dorsal crest; sex structure; DNA barcoding

**Zoobank** http://zoobank.org/9351C829-16CD-4084-A12E-1C98889982CD

### Introduction

*Zachvatkinibates* Shaldybina, 1973 sensu stricto, with the type species *Z. nemoralis* Shaldybina, 1973, is a medium sized genus that comprises 18 species, according to Subías (2004, 2022). However, this author included *Minunthozetes selgae* Pérez-Iñigo, 1976 in this genus, which is not consistent with diagnosis of Behan-Pelletier & Eamer (2008). In the later species, the posterior notogastral tectum is complete, whereas in *Zachvatkinibates* it is medially broken.

The diagnosis of the adult of *Zachvatkinibates* was proposed by Shaldybina (1973), but Behan-Pelletier (1988) and Behan-Pelletier & Eamer (2005) consecutively modified it, and in the latter paper it was as follows: notogaster with rounded, convex medial process on anterior tectum and pair of thickened bands bordering medial process, posterior notogastral tectum incomplete medially, indicated by wide margination or narrow slit bordered by overlapping tectal lobes; lenticulus absent, octotaxic system expressing sexually dimorphism in some

Received 12 August 2022
Accepted 09 December 2022
Published 13 January 2023

Corresponding author
Anna Seniczak ID:
Anna.Seniczak@uib.no

Academic editor
Baumann, Julia

https://doi.org/10.24349/skz7-19nt

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_6.png" /></figure>

Seniczak A. et al.

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote>ªDepartment of Natural History, University Museum of Bergen, University of Bergen, Bergen, Norway.</footnote>

<footnote>ᵇDepartment of Evolutionary Biology, Faculty of Biological Sciences, Kazimierz Wielki University, Bydgoszcz, Poland.</footnote>

<footnote>ᶜNorwegian Institute of Bioeconomy Research, NIBIO - Division of Environment and Natural Resources, Svanhovd, Norway.</footnote>
species, postanal porose area and axillary saccule of subcapitulum present, and mental tectum
of subcapitulum absent.

The main aim of this paper is to describe and illustrate the morphology and ecology of
*Z. svanhovdi*, and to compare it with congeners. Another aim is to update information on
Punctoribatidae in Fennoscandia, and add *Z. svanhovdi* and *Mycobates carli* (Schweizer, 1922)
to the list of species in this region.

**Material and methods**

**Study site**

The sample was collected at the river outlet (Figure 1) of the Jacob's River in north-eastern
Finnmark, Sør-Varanger municipality, Norway. The region is characterized by an oceanic-
continental climate with a mean annual temperature of 0.1 °C and a mean annual precipitation
of approximately 620 mm (Bjørklund et al. 2015). The Jacob's River (Grense Jakobselv;
area 237 km²) is a river that runs along the Norwegian-Russian border and discharges into
the Varanger Fjord, a bay located in the Barents Sea. The river is part of the Norwegian
Protection Plan II3 and hence cannot be used for hydropower purposes. Generally, the
vegetation can be characterized as northern boreal in a transition zone between oceanic and
continental vegetation sections (Moen 1998, Bjørklund et al. 2015) in north-eastern Finnmark.
The landscape along the river consists of hills with sparse heath vegetation near the coast,
hilly landscapes and plateaus further inland. Along the river, the vegetation is commonly
characterized by different vegetation types: closest to the river is often a well-developed sandy
beach with key plants like the oyster plant (Mertensia maritima L.), followed by a transition
zone with oyster plant and sea pea (Lathyrus japonicus Willd.), white sandy dunes with sand
ryegrass [Leymus arenarius (L.) Hochst.] and sea pea, grey sandy dunes with sand ryegrass
and red fescue (Festuca rubra L.), and dry, species-poor dune grass heath with strong hints of
fringed pink (Dianthus superbus L.; https://elvedelta.miljodirektoratet.no/delta-322.htm,
https://www.biodiversity.no/NiN2.0/T21-C-2). At the river outlet, where the sample was
taken, there is also a semi-limestone beach meadow (Elven & Johansen 1983), which is
characterized by freshwater-affected salt weed (Salicornia sp.) communities (P.E. Aspholm,
NIBIO, Svanhovd; pers. comm.). The sampling location was in an area that is considered
brackish water area (https://elvedelta.miljodirektoratet.no/delta-322.htm).

**Sampling and studying of mites**

The adults of *Z. svanhovdi* A. Seniczak *et* S. Seniczak **sp. nov.** used in this study were
collected by A. Fjellberg on 26.08.2021 from Troms and Finnmark county, Northern Norway,
Sør-Varanger community and Gr. Jakobselv locality, at river outlet (69.71258° N, 30.87123° E,
0.5 m a. s. l.). One collective sample was taken on the shore of Jacob's River, in the transect
from the river edge to more distant part from it (about 10 m long, Figure 1). This sample was
extracted in Berlese funnel for one week. We investigated the density, sex ratio, number of
gravid females and carried eggs, and body length and width, based on 30 randomly selected
specimens. We measured a total body length (tip of rostrum to posterior edge of notogaster) in
lateral aspect and body width (widest part of notogaster) in dorsal aspect, and size of anal and
genital openings and setae perpendicularly to their length in µm.

The illustrations are limited to the dorsal, lateral and ventral aspect and some leg segments of
adult, and palp and chelicera. Illustrations were prepared from individuals mounted temporarily
in lactic acid. In the text and figures we used the following abbreviations: rostral (ro), lamellar
(le), interlamellar (in) and exobothridial (ex) setae, lamella (La), translamella (Tr), bothridium
(bo), bothridial seta (bs), tutorium (Tut), pteromorph (Ptm), notogastral setae ($c_2$, l-, h-, p-
series), porose areas (Aa, A1, A2, A3), lymphassures (ia, im, ih, ip, ips, iad), circumpedal carina
cp), opisthonotal gland opening (gla), subcapitular setae (a, m, h), pedotectum (Pd), custodium
<figure><img src="image_7.png" /><figcaption>**Figure 1** General view of the river shore, where *Zachvatkinibates svanhovdi* A. Seniczak *et S.* Seniczak *sp. nov.* was collected (photo: Arne Fjellberg).</figcaption></figure>

(cus), discidium (Dis), genal tooth (gt), cheliceral setae (cha, chb), Trägårdh organ (Tg), palp setae (sup, inf, l, d, cm, acm, lt, vt, ul, su) and solenidion ω, epimeral setae (1a-c, 2a, 3a-c, 4a-c), genital (g) and aggenital (ag) setae, adanal and anal setae (ad-, an-series), leg solenidia (σ, φ, ω), famulus (ε) and setae (bv, ev, d, l, ft, tc, it, p, u, a, s, pv, pl, v). Terminology used follows that of Grandjean (1949, 1953) and Norton and Behan-Pelletier (2009). The species nomenclature follows Subías (2004).

For scanning electron microscopy (SEM), four mites were air-dried and coated with Au/Pd in a Polaron SC502 sputter coater and placed on Al-stubs with double-sided sticky carbontape. Observations and micrographs were made with a ZEISS Supra 55VP scanning electron microscope.

## DNA barcoding

Five specimens of *Z. svanhovdi* were used for DNA barcoding. Each specimen was photographed, and the photos are available at Barcode of Life Data System (BOLD, http://boldsystems.org). The specimens were subsequently placed in a well containing 50 ml of 90% ethanol in a 96-well microplate and send to the Canadian Centre for DNA Bar-coding (CCDB 2021). Mites were sequenced for the barcode region of the COI gene according to standard protocols at CCDB (http://www.ccdb.ca), using either LepF1/LepR1 (Hebert *et al.* 2003) or LCO1490/HCO2198 (Folmer *et al.* 1994) primer pairs. The DNA extracts were placed in archival storage at -80 °C at the Canadian Centre for DNA Barcoding. The sequences are available in GenBank (accession numbers in the section on the Type material). The voucher specimens are stored in the Entomological Collections at the Department of Natural History, University Museum of Bergen, University of Bergen, Bergen, Norway (ZMBN).

Obtained COI sequences were blasted against GenBank to detect and exclude possible contaminations and ensure high quality. Sequence variation within *Z. svanhovdi* specimens was calculated in BOLD using Kimura 2 Parameter distance model, pairwise deletion, and BOLD Aligner (Amino Acid based HMM).
## Results

**Zachvatkinibates svanhovdi A. Seniczak et S. Seniczak sp. nov.**

Zoobank: A6272084-12BA-4FAC-BA43-7582C1D30CEB

(Figures 2–11)

### Diagnosis

Adult of medium size (449–507), with generic characters of *Zachvatkinibates* given by Behan-Pelletier & Eamer (2005). Prodorsal seta *in* long, translamella narrow, notogastral setae short and distally pliable, notogastral porose areas usually oval and of medium size, but *Aa* can be larger, especially in males. Postanal porose area *Ap* absent, dorsal crest on tarsus I present.

### Description of adults

**Measurements** — Holotype (female) length 494, and width 299. Mean length (and range) of females 490.3±7.1 (481–507, n = 15) and males 468.9±12.2 (449–481, n = 15), mean maximum width (and range) of females 299.0±6.2 (293–319) and males 276.7±8.3 (254–293).

**Integument** — Brown and microporose.

<figure><img src="image_8.png" /><figcaption>**Figure 2** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* sp. nov., scale bars 50 µm. A – female, dorsal aspect, legs partially drawn; B – porose area *Aa* of another female (enlarged); C – notogaster of male; D – shape of porose area *Aa* of another male.</figcaption></figure>
<figure><img src="image_9.png" /><figcaption>**Figure 3** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* sp. nov., female. A – ventral aspect, legs partially drawn, scale bar 50 µm; B – lateral aspect, legs partially drawn, scale bar 50 µm, mouthparts, right side, scale bars 20 µm; C – chelicera (Trägårdh organ in transparent area); D – palp.</figcaption></figure>

**Prodorsum** — Sub-triangular, rostrum rounded. Seta *ro* relatively long (47–52), laterally barbed and in lateral position (Figures 2A, 3A, 3B, 5A, 5D, 6A–C, 8C, 8D, 9A, 10A), seta *le* longer (62–66), and seta *in* longest (77–80), both finely barbed; all acuminate; seta *ex* short (17) and smooth. Lamella long (77–110), narrow at base and wider in place of connection with translamella, translamella of medium size (29–31), narrow, lamellar cusp (length 15) rounded. Bothridium with well-developed medial scale, bothridial seta of medium size (43–46), clavate with smooth stalk and barbed head (Figures 2A, 3B, 5A, 5B, 5D, 6A, 6B, 6D, 7B, 7C, 9C).

**Notogaster** — Longer (females 380–398, males 357–378) than wide (females 293–319, males 254–293) with well-developed movable pteromorphs. Notogastral setae (10 pairs, including *c*₂) short (15–26), and smooth (Figures. 2–3B, 4, 5, 6A, 6D, 7A–C, 8A, 9C, 9D). Lyrifissure *ia* posterolateral to seta *c*₂, *im* anterior to seta *h*₃, *ip* anterolateral to seta *p*₁, *ips* anterior to seta *p*₃, *ih* posterior to seta *la*, and opisthonotal gland opening *gla* anterolateral to seta *h*₃ (Figures 2A, 8D). Shape of porose areas (4 pairs) differs between females and males. In females, porose areas oval, *Aa* slightly larger than other porose areas (Figures 2, 3B), in some individuals *Aa* can be elongated. In 15 males investigated, porose areas clearly larger (Figure 2C, 2D) than in females, and in some individuals *Aa* can be elongated, and of crescent shape or divided in two parts. Generally, in both sexual forms shape of other porose areas slightly varies within individuals, and postanal porose area is absent. Posterior tectum of notogaster is broken in middle and rounded ends overlap (Figure 4).

**Gnathosoma** — Subcapitular setae *h*, *m* and *a* short (18–24) and smooth (Figures 3A, 3B,
<figure><img src="image_10.png" /><figcaption>**Figure 4** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* **sp. nov.**, female, posterior aspect, scale bar 50 µm.</figcaption></figure>

5C, 7B-D, 8C, 8D). Palp short (75), setae relatively short and smooth (Figures 3D, 5D, 9A, 10A), formula of palp setae [trochanter to tarsus (+ solenidion ω)]: 0-2-1-3-9(1). Chelicera longer (96), seta *cha* longer than *chb*, both barbed (Figures 3C, 9A, 10A). Axillary saccule short (12).

**Ventral and lateral regions** — Most epimeral setae short and smooth (10–13, Figures 3A, 5C, 8B, 8C). Genital setae (six pairs) as short as epimeral setae, $g_{1-3}$ in anterior position. Aggenital setae (one pair) also short, adanal setae longer (three pairs, 22–29) and anal setae longest (two pairs, 29–32), all smooth (Figures 3A, 3B, 4, 5B, 5C, 8A, 8B). Lyrifissure *iad* short, located lateral to anterior part of anal plates. Discidium large, triangular; custodium long (40–42), narrow and pointed; tutrium long (122–126), narrow at base and distally wider and pointed (Figures 3B, 7B-D, 9A, 9B, 10, 11). Pedotectum I large (73 x 30), narrow at base and concave dorsally (Figure 3B), and with longitudinal striae, genal tooth of medium size (30–22). Circumpedal carina well formed. Humerosejugal porose areas *Am* and *Ah* oval.

**Legs** — Trochanters III and IV and all femora flattened with ventral carina. Porose areas on paraxial side of trochanters III and IV and femora I–IV. Femur I with longitudinal striae, most leg setae barbed (Figures 5, 6, 7, 8B-D, 9A-C). Seta *ft"* on tarsus I relatively long, solenidia ω₁ and ω₂ of similar length, famulus short; solenidia ω₁ and ω₂ and famulus ε located on small dorsal crest. Solenidia ω₁ and ω₂ on tarsus II relatively long and of similar size. Formulae of leg setae [trochanter to tarsus (+ solenidia)]: I – 1-4-3(1)-4(2)-20(2); II – 1-4-3(1)-4(1)-15(2); III – 2-3-1(1)-3(1)-15; IV – 1-2-2-3(1)-12. Tarsi heterotridactlylous. Homology of leg setae and solenidia given in Table 1.
<figure><img src="image_11.png" /><figcaption>**Figure 5** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* **sp. nov.** adult, SEM micrographs. A – dorsal view; B – lateral view; C – ventral view; D – frontolateral view.</figcaption></figure>

## Results of DNA barcoding

The barcode based on the cytochrome oxidase I (COI) mitochondrial DNA region (length: 658 bp) of five specimens of the new species is provided under GeneBank accession numbers: OP824977 (SUB2896631 SOITS201-22), OP824978 (SUB2896631 SOITS202-22), OP824979 (SUB2896631 SOITS203-22), OP824980 (SUB2896631 SOITS205-22), OP824981

<table><caption>**Table 1** Leg setae (Roman letters) and solenidia (Greek letters) in *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* **sp. nov.**</caption><thead><tr><th>Legs</th><th>Trochanter</th><th>Femur</th><th>Genu</th><th>Tibia</th><th>Tarsus</th></tr></thead><tbody><tr><td>I</td><td>v'</td><td>d, bv'', (l)</td><td>(l), σ, v'</td><td>(l), (v), φ₁, φ₂</td><td>l'', (ft), (tc), (it), (pl), (p), v', (pv), (a), s, (u), ε, ω₁, ω₂</td></tr><tr><td>II</td><td>v'</td><td>d, bv'', (l)</td><td>(l), σ, v'</td><td>(l), (v), φ</td><td>(ft), (tc), (it), (p), (pv), s, (a), (u), ω₁, ω₂</td></tr><tr><td>III</td><td>v', l'</td><td>d, ev', l'</td><td>l', σ</td><td>(v), l', φ</td><td>(ft), (tc), (it), (p), (pv), s, (a), (u)</td></tr><tr><td>IV</td><td>v'</td><td>d, ev'</td><td>d, l'</td><td>(v), l', φ</td><td>ft'', (tc), (u), (pv), s, (a), (p)</td></tr></tbody></table>

Note: pairs of setae in parentheses.
<figure><img src="image_12.png" /><figcaption>**Figure 6** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* sp. nov. adult, SEM micrographs. Dorsal view. A – anterior and medial part; B, C – anterior part; D – medial and posterior part.</figcaption></figure>

(SUB2896631 SOITS204-22). The maximum variation within the species was 2.41% (p-dist) that is expected intraspecific variation of COI in oribatids. (e.g., Pfingstl *et al.* 2019, Seniczak *et al.* 2019). These are the first public COI sequences to be reported for the genus *Zachvatkinibates* and therefore, genetic distances to other species could not be calculated.

**Distribution, ecology and biology**

*Zachvatkinibates svanhovdi* was found in Troms and Finnmark county, Northern Norway, in a collective sample, taken in the transect on the shore of the river, so it is not possible to give precisely the species microhabitat. Among the 30 individuals investigated, collected at the end of August, the sex ratio (females to males) was 1:1, and no females were gravid and carried eggs.

**Type material**

The holotype (female) and four paratypes (2 females and 2 males) are deposited in the University Museum of Bergen, University of Bergen, Bergen, Norway (ZMBN).
<figure><img src="image_13.png" /><figcaption>**Figure 7** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* sp. nov. adult, SEM micrographs. Lateral view. A – posterior and medial part; B – anterior and medial part; C, D – anterior part.</figcaption></figure>

**Etymology**

The species is named in honour of the biological station NIBIO Svanhovd, which is located in eastern Finnmark in the Pasvik valley, where the project leading to the collection and description of this new species was carried out.

**Comparison of morphology of *Zachvatkinibates svanhovdi* with congeners and remarks**

Seniczak *et al.* (2020d) compared selected morphological characters of adults of 15 *Zachvatkinibates* species and stated that the largest among them is *Z. nortoni* Behan-Pelletier *et Eamer, 2005*, and smallest is *Z. anoporosus* (Mahunka, 2006), and the body length of several species overlaps. These authors also compared the length of prodorsal seta *in* and notogastral setae, shape of translamella and some porose areas. To this list *Z. erimo* Shimano *et Aoki, 2019* should be added. Based on this list of species, *Z. svanhovdi* A. Seniczak *et S. Seniczak* sp. nov. is the most similar to *Z. quadrivertex* (Halbert, 1920), but differs from it mainly by the shape of notogastral setae, posterior tectum of notogaster and lack of postanal porose area *Ap*, which in *Z. svanhovdi*, most notogastral setae are distally pliable,
<figure><img src="image_14.png" /><figcaption>**Figure 8** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* **sp. nov.** adult, SEM micrographs. Ventral view. A – posterior part; B – medial part; C, D – anterior part.</figcaption></figure>

whereas in *Z. quadrivertex* they are not. In both species, the posterior tectum of notogaster
is broken in the middle, the ends are rounded, but in *Z. svanhovdi* they overlap, whereas in
*Z. quadrivertex* they are separated by a certain distance. In *Z. svanhovdi*, *lyrifissures ips* and
*ih* are placed at larger distances from seta *p* than in *Z. quadrivertex*. Moreover, in female of
*Z. svanhovdi* porose area *Aa* is located between setae *la* and *lm* (*vs*. anteromedial to seta *la* in
*Z. quadrivertex*), and porose areas *A2* and *A3* are as large as *A1*, whereas in *Z. quadrivertex*
they are smaller than *A1*. In some males of *Z. svanhovdi*, the porose areas *Aa* are larger than
in *Z. quadrivertex*. In the diagnosis for *Zachvatkinibates* by Behan-Pelletier & Eamer (2005),
postanal porose area *Ap* is present, but *Z. svanhovdi* has no porose area, so the diagnosis for
this genus should be changed to “postanal porose area *Ap* present or absent”.

In *Z. svanhovdi*, sexual dimorphism is observed in the body size and octotaxic system (Table 2). Males are slightly smaller than females, have larger porose areas, and *Aa* can be divided in two parts (*vs*. in females it is uniform). In six other species of *Zachvatkinibates*, sexual dimorphism is more distinct than in *Z. svanhovdi* (Table 2). Interestingly, the species listed in Table 2 seem to be correlated with littoral habitats of cold climate zones (Behan-Pelletier & Eamer 2010, Behan-Pelletier 2015).
<figure><img src="image_15.png" /><figcaption>**Figure 9** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak sp. nov.* adult, SEM micrographs. A, B – anterior part; frontolateral view; C – bothridial setae; D – gla opening.</figcaption></figure>

The adult of *Z. svanhovdi* has the dorsal crest on tarsus I, as other species of *Zachvatkinibates*
(Shaldybina & Grishina 1987, Behan-Pelletier 1988, Bayartogtokh & Aoki 1998, Behan-
Pelletier & Eamer 2005, Seniczak *et al.* 2020d). In *Minunthozetes* Hull, 1916, the leg crests
are better developed than in *Zachvatkinibates* and occur on part of tibiae and tarsi I, II and
IV (Seniczak & Seniczak 2018, Seniczak *et al.* 2018) and may suggest close relationship of
*Zachvatkinibates* with *Minunthozetes*. In other genera of Punctoribatidae such as *Mycobates*
Hull, 1916 and *Punctoribates* Berlese, 1908, the leg crests on tibiae and tarsi are absent
(Seniczak & Seniczak 2008, Seniczak *et al.* 2015, 2020c). The latter two genera have dorsal
projection on tibia II, which in *Zachvatkinibates* and *Minunthozetes* is absent.

Based on the checklists of Oribatida of Norway (Mehl 1979), Sweden (Lundqvist 1987), and Finland (Niemi *et al.* 1997), and other publications listed in Seniczak and Seniczak (2020) and Seniczak *et al.* (2020a,b, 2021), *Z. svanhovdi* represents the first finding of *Zachvatkinibates* in Norway and Fennoscandia. Species of *Zachvatkinibates* are known from Central and Eastern Asia, Northern America, and only few from Europe (Table 4). Their ecology varies a lot. For example, six species inhabit littoral zone and beach debris, other were found in bogs, forests, including forest canopy, and even in the debris in the alpine zone (Table 4).
<figure><img src="image_16.png" /><figcaption>**Figure 10** *Zachvatkinibates svanhovdi* A. Seniczak *et S. Seniczak* sp. nov. adult, SEM micrographs. Leg I. A – frontolateral view; B – lateral view; C – legs I–III; D – leg III and VI.</figcaption></figure>

**Comments on Punctoribatidae in Fennoscandia and new record of Mycobates carli**

The family Punctoribatidae to which *Zachvatkinibates* belongs to is represented in Fennoscandia by four genera and ten nominative species, including *Mycobates carli* (Schweizer, 1922) which is reported here as new to Norway and Scandinavian part of Fennoscandia (Table 3). It was found in alpine tundra in Finse (Vestland, Western Norway, 60°35′14.0″N 7°30′50.2″E, 1228 m a. s. l., 06.08.2022, leg. Anna Seniczak), on forest floor with mosses and lichens. The description of climatic conditions in Finse was given earlier (Seniczak & Seniczak 2021). This species was represented in the sample only by few adults, (density 10 individuals per 500 cm³), and co-occurred with more abundant *M. sarekensis* (Trägårdh, 1910).

*Mycobates carli* has been mostly considered a southern European (Subías 2004, 2022), European (Weigmann 2006) or Eurosiberian species (Honciuc 1993), known especially from the Central, Southern, Eastern, and South-Eastern parts of Europe. It has been reported from numerous localities in the Alps (Fischer & Schatz 2020, Schatz & Bruckner 2021), the Giant Mountains (Materna 2000, Starý 2006, Miko 2013), Carpathian Mts. (Nae & Băncilă 2017, Skubała & Maslak 2009), and is also known from Caucasus Mts. (Schatz & Bruckner 2021).
<figure><img src="image_17.png" /><figcaption>**Figure 11** *Zachvatkinibates svanhovdi* A. Seniczak *et S*. Seniczak *sp. nov.*, leg segments of adult (part of femur to tarsus), right side, antiaxial aspect, setae on the opposite side not illustrated are indicated in the legend, scale bar 20 µm. A – leg I, tarsus (*pl*'); B – leg II; C – part of tibia and tarsus I; D – leg III, tibia (*v*'); E – leg IV.</figcaption></figure>
<table><caption>**Table 2** Selected morphological characters of *Zachvatkinibates* species with distinct dimorphism of notogastral porose areas; ♀ – females, ♂ – males.</caption><thead><tr><th rowspan="2">Species</th><th rowspan="2">Body length (µm)</th><th rowspan="2">Seta in reaching rostrum</th><th rowspan="2">Shape of Aa</th><th colspan="3">Size of porose areas</th><th rowspan="2">Setae ad-, an-series</th></tr><tr><th>A1</th><th>A2</th><th>A3</th></tr></thead><tbody><tr><td>Z. erimo Shimano et Aoki 2019</td><td>♀440–449<br>♂450</td><td>yes</td><td>rounded, as large as in male</td><td>as large as Aa</td><td>as large as A1<br>A2 and A3 fused</td><td>smaller than A2</td><td>as long as ag</td></tr><tr><td>Z. nortonii Behan-Pelletier et Eamer, 2005</td><td>♀525–594<br>♂525–574</td><td>yes</td><td>rounded, as large as in male</td><td>smaller than Aa</td><td>as large as A1</td><td>as large as A2</td><td>as long as ag</td></tr><tr><td>Z. quadrivertex (Halbert, 1920)</td><td>♀450–500<br>♂415–480</td><td>yes</td><td>oval, smaller than in male<br>elongated</td><td>as large as Aa<br>smaller than Aa</td><td>smaller than A1<br>as large as A1</td><td>larger than A2<br>as large as A2</td><td>?</td></tr><tr><td>Z. shaldybinae Behan-Pelletier et Eamer, 2005</td><td>♀499–528<br>♂470–508</td><td>no</td><td>rounded, smaller than in male<br>elongated or divided</td><td>smaller than Aa<br>smaller than Aa</td><td>smaller than A1<br>smaller than A1</td><td>larger than A2<br>larger than A2</td><td>?</td></tr><tr><td>Z. schatzi Behan-Pelletier et Eamer, 2005</td><td>♀528–554<br>♂505–544</td><td>no</td><td>elongated, smaller than in male</td><td>smaller than Aa<br>smaller than Aa</td><td>smaller than A1<br>A1, A2 and A3 fused</td><td>larger than A2<br>as large as A2</td><td>?</td></tr><tr><td>Z. maritimus Shaldybina, 1973</td><td>♀473<br>♂?</td><td>yes</td><td>irregular, smaller than in male<br>elongated</td><td>smaller than Aa<br>smaller than Aa</td><td>smaller than A1<br>as large as A1</td><td>larger than A2<br>as large as A2</td><td>as long as ag</td></tr><tr><td>Z. svanhovdi A. et S. Seniczak sp. nov.</td><td>♀481–507<br>♂449–481</td><td>yes</td><td>oval, smaller than in male<br>elongated or divided</td><td>smaller than Aa<br>smaller than Aa</td><td>smaller than A1<br>as large as A1</td><td>larger than A2<br>as large as A2</td><td>longer than ag</td></tr></tbody></table>

In the north of Europe it has only been reported from plain tundra of the Kola Peninsula (Leonov & Rakhleeva 2015). It is considered subalpine and alpine (Weigmann 2006, Fischer & Schatz 2007), predominantly lichenocolous (Materna 2000), and muscicolous (Schatz & Bruckner 2021). It has been found in a peatland, 870 m a. s. l. (Starý 2006), at dry limestone plateau, 550 m a. s. l. (Lazarus & Krisper 2014), and in soils polluted with heavy metals in Romania, where it was associated with Cu, As and Mn (Manu et al. 2019). Most often it has been collected from soil or ground vegetation, but has been also reported from the unique microhabitat of the intermediate layer between the base of the soil and the bedrock, called “mesovoid shallow substratum” (Nae & Băncilă 2017) and from decaying wood (Skubała & Maslak 2009).

## Acknowledgements

We thank the anonymous reviewers for helpful comments and suggestions that improved the quality of this paper. We also express our gratitude to Dr. Arne Fjellberg for collection of the sample that contained the specimens. Further, we would like to thank Juho Vuolteenaho for

<table><caption>**Table 3** Occurrence of Punctoribatidae in Fennoscandia, N – Norway, S – Sweden, F – Finland; * – new to Norway and Fennoscandia.</caption><thead><tr><th>Genus</th><th>Species</th><th>Country</th></tr></thead><tbody><tr><td rowspan="3">Minunthozetes Hull, 1916</td><td><i>M. pseudofusiger</i> (Schweizer, 1922)</td><td>F, N</td></tr><tr><td><i>M. semirufus</i> (C.L. Koch, 1841)</td><td>F, S, N</td></tr><tr><td><i>M. carli</i> (Schweizer, 1922)*</td><td>N</td></tr><tr><td rowspan="4">Mycobates Hull, 1916</td><td><i>M. parmeliae</i> (Michael, 1884)</td><td>F, S, N</td></tr><tr><td><i>M. sarekensis</i> (Trägårdh, 1910)</td><td>F, S, N</td></tr><tr><td><i>M. tridactylus</i> Willmann, 1929</td><td>N</td></tr><tr><td><i>P. hexagonus</i> Berlese, 1908</td><td>F</td></tr><tr><td rowspan="2">Punctoribates Berlese, 1908</td><td><i>P. punctum</i> (C.L. Koch, 1839)</td><td>F, S, N</td></tr><tr><td><i>P. sellnicki</i> Willmann, 1928</td><td>F, S, N</td></tr><tr><td>Zachvatkinibates Shaldybina, 1973</td><td><i>Z. svanhovdi</i> A. et S. Seniczak sp. nov.*</td><td>N</td></tr></tbody></table>
<table><caption>Table 4 Distribution and ecology of *Zachvatkinibates* species.</caption><thead><tr><th>Species</th><th>Distribution</th><th>Habitat</th><th>Sexual dimorphism</th><th>Reference</th></tr></thead><tbody><tr><td>Z. anoporousus (Mahunka, 2006)</td><td>Romania (Poiana Brazilor, Ocna Şugatag, Muntii Piatra, 900 m a. s. l.)</td><td>Bog, decaying wood, some Sphagnum</td><td>Unknown</td><td>Mahunka (2006)</td></tr><tr><td>Z. conjunctus Shaldybina, 1987</td><td>Russia (Stolby Nature Reserve, Krasnoyarsky Krai)</td><td>Moss cushion at the base of a pine tree</td><td>Unknown</td><td>Shaldybina & Grishina (1987)</td></tr><tr><td>Z. eoeryi (Mahunka, 1972)</td><td>South Europe: Croacia (near Split), Portugal (Lagoon of Faro)</td><td>Predominantly marine littoral debris, e.g., algae, detritus</td><td>Unknown</td><td>Mahunka (1972), Weigmann (2009)</td></tr><tr><td>Z. epiphytos Behan-Pelletier, Eamer et Clayton, 2001</td><td>Canada (Montane Alternative Silvicultural Systems, Vancouver Island), USA (Willamette National Forest, Oregon)</td><td>Epiphytes (mainly lichens) in coniferous forest canopy</td><td>No</td><td>Behan-Pelletier et al. (2001), Winchester et al. (2008)</td></tr><tr><td>Z. erimo Shimano et Aoki, 2019</td><td>Japan (Hyakuninhama, Shono, Erimo City, Horoizumi)</td><td>Roots of Poaceae, dead parts of plants, soil on a sandy beach</td><td>Yes</td><td>Shimano & Aoki (2019)</td></tr><tr><td>Z. latilamellatus Bayartogtokh et Aoki, 1998</td><td>Mongolia</td><td>Siberian larch (Larix sibirica Ledeb.) forest</td><td>Unknown</td><td>Bayartogtokh & Aoki (1998), Seniczak et al. (2020b)</td></tr><tr><td>Z. lobatus (Hammer, 1977)</td><td>Pakistan (Lowari Pass, 3400 m a. s. l.)</td><td>Thin moss on the soil, grass, Potentilla, Epilobium, Fragaria, Geranium, pine cones</td><td>Unknown</td><td>Hammer (1977)</td></tr><tr><td>Z. maritimus Shaldybina, 1973</td><td>Russia (Primorsky Krai), USA (Alaska), Canada (Yukon, Vancouver Island, British Columbia)</td><td>Coastal debris, supratidal meadows, coastal and lentic littoral habitats</td><td>Yes</td><td>Shaldybina (1973, 1987), Behan-Pelletier (1988), Ryabinin (2015)</td></tr><tr><td>Z. mongolicus Bayartogtokh, 2003</td><td>Mongolia (Gobi Gurvan Saykhan Nature Reserve, southern Mongolia)</td><td>Semidesert, soil under Artemisia santolinifolia (Pamp.) Turcz. ex Krasch and organic debris accumulated</td><td>No</td><td>Bayartogtokh (2003)</td></tr><tr><td>Z. nemoralis Shaldybina, 1973</td><td>Kyrgyzstan and Mongolia</td><td>Litter between roots, the alpine region</td><td>Unknown</td><td>Shaldybina (1973, 1987)</td></tr><tr><td>Z. nortoni Behan-Pelletier et Eamer, 2005</td><td>Canada (British Columbia)</td><td>Beach debris</td><td>Yes</td><td>Behan-Pelletier & Eamer (2005)</td></tr><tr><td>Z. quadrivertex (Halbert, 1920)</td><td>Atlantic coastal regions of Europe, from Denmark to South Portugal, Russia (Islands of White Sea, Far East: Kunashir Island in Kuril Islands)</td><td>Lower tidal level of marine saltmarshes, fir forest</td><td>Yes</td><td>Shaldybina (1987), Weigmann (2009), Ryabinin (2015)</td></tr><tr><td>Z. schatzi Behan-Pelletier et Eamer, 2005</td><td>Canada (British Columbia)</td><td>Beach debris</td><td>Yes</td><td>Behan-Pelletier & Eamer (2005)</td></tr><tr><td>Z. shaldybinae Behan-Pelletier et Eamer, 2005</td><td>Canada (eastern and northeastern), Greenland</td><td>Littoral</td><td>Yes</td><td>Behan-Pelletier & Eamer (2005), Makarova & Behan-Pelletier (2015)</td></tr><tr><td>Z. tetrasklerosis Behan-Pelletier, 1988</td><td>USA (California)</td><td>Litter of different species of conifers</td><td>No</td><td>Behan-Pelletier (1988)</td></tr><tr><td>Z. volgini Shaldybina, 1973</td><td>Russia (Far East: Shikotan and Yuri islands in Kuril Islands, Sakhalin)</td><td>Gull's nest on the coast, quackgrass [Elymus repens (L.) Gould] and brier meadow</td><td>Unknown</td><td>Shaldybina (1973, 1987), Ryabinin (2015)</td></tr></tbody></table>

assistance with field work. The study was supported by the grant to NIBIO from the Norwegian Taxonomy Initiative (Grant No. 9-21, 70184244). The DNA sequencing was financed by Norwegian Barcode of Life (NorBOL).

ORCID

Anna Seniczak https://orcid.org/0000-0003-0224-5397

Stanisław Seniczak https://orcid.org/0000-0001-7393-5606

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