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Article

Morphological Ontogeny and Ecology of a Common Peatland Mite, Nanhermannia coronata (Acari, Oribatida, Nanhermanniidae)

by
Stanisław Seniczak
1 and
Anna Seniczak
2,*
1
Department of Evolutionary Biology, Faculty of Biological Sciences, Kazimierz Wielki University, 85-093 Bydgoszcz, Poland
2
Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Inland Norway University of Applied Sciences, 2318 Elverum, Norway
*
Author to whom correspondence should be addressed.
Animals 2023, 13(22), 3590; https://doi.org/10.3390/ani13223590
Submission received: 27 September 2023 / Revised: 8 November 2023 / Accepted: 10 November 2023 / Published: 20 November 2023
(This article belongs to the Special Issue The Ecology, Evolution, Systematics and Behaviour of Mites)
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Abstract

:

Simple Summary

Nanhermannia coronata is a common and abundant oribatid species in peatlands, but it can be easily mistaken for N. sellnicki as an adult. The identity of adults of N. coronata investigated herein from several sites in Norway and Ireland was supported by the COI sequence data. Based on this material, the morphological ontogeny of N. coronata was investigated, and some characters were found that clearly differentiate N. coronata from N. sellnicki, like the number of setae on femora of adults and tritonymphs, the shape of insertions of prodorsal seta in and all gastronotal and adanal setae of juveniles. Our ecological observations confirm a common occurrence of N. coronata in raised bogs, a high percentage of juvenile stages in populations and a preference of this species for humid microhabitats, whereas N. sellnicki is less common than N. coronata and occurs in drier habitats.

Abstract

Nanhermannia coronata Berlese, 1913, is a common and abundant oribatid species in peatlands but can be easily mistaken for N. sellnicki Forsslund, 1958, as an adult. Therefore, the identity of adults of N. coronata from several sites in Norway and Ireland was supported by the COI sequence data, and based on this material, the morphological ontogeny of this species is described and illustrated to highlight the differences between N. coronata and N. sellnicki. In all juvenile stages of N. coronata, the bothridial seta is absent, but two pairs of exobothridial setae are present, including short exp and exa reduced to its alveolus. In the larva, seta f1 is setiform, but in the nymphs, it is reduced to its alveolus. Most prodorsal and gastronotal setae of larva are short, and of nymphs they are long. In all instars, the leg segments are oval in cross section and relatively thick, and many setae on tarsi are relatively short, thick and conical, except for longer apical setae. Seta d accompanies solenidion σ on all genua, φ1 on tibia I and φ on other tibiae. We found some morphological characters that clearly differentiate N. coronata from N. sellnicki, like the number of setae on femora of adults and tritonymphs, the shape of insertions of prodorsal seta in and all gastronotal and adanal setae of juveniles; in N. sellnicki, these setae are inserted in small individual depressions, whereas in N. coronata, these depressions are absent. Our ecological observations confirm a common occurrence of N. coronata in raised bogs, a high percentage of juvenile stages in its populations and a preference of this species for humid microhabitats, whereas N. sellnicki is less common than N. coronata and occurs in drier habitats.

1. Introduction

Nanhermannia Berlese, 1913, sensu stricto (N. nanus Nicolet, 1855, as the type species) comprises 31 nominative species [1]; all are of medium size as adults. The diagnosis of adults of this genus was given by Seniczak et al. [2] as follows: Adults are medium-sized (450–660 µm), elongated, brown to dark brown and narrow, with bothridia and bothridial setae situated on top of the prodorsum; there are protuberances on the posterior part of the prodorsum with sclerotized tubercles, often extending above the anterior part of the notogaster. The notogaster is cylindrical, with a coarse structure of pits and 15 pairs of long setae, curved and appressed to body, extended ventrally and connected ventro-medially; an arched suture delimits the genital and aggenital area laterally (diagastry). The aggenital plate is fused with the epimere, and the adanal plate is fused with the notogaster, but still recognizable; the formula of the epimeral setae is usually 3-1-3-3 or 3-1-3-4, but in some species, hypertrichy on some epimeres occurs; 7–10 pairs of genital setae, two pairs of aggenital setae, two pairs of anal setae and three pairs of adanal setae are present. Legs have one claw.
Identification of Nanhermannia species is not easy because they are relatively similar to each other by having a similar body shape, structure of pits and shape of notogastral setae. The species differ from one another by the shape of protuberances on the posterior margin of the prodorsum and the number of sclerotized tubercles, which are considered diagnostic in Nanhermannia, but these characters vary in some species [3,4,5,6,7,8,9,10], including N. coronata Berlese, 1913, being a source of confusion. For example, during the revision of the twenty-year-old oribatid mite collection of the Institute of Biology, University of Latvia, a high discrepancy in the identification of N. coronata was detected, and ca. 50% of specimens (out of 40 studied) were wrongly identified [11]. Therefore, more investigations on the morphology of Nanhermannia species are required, including the juvenile stages and molecular investigations to improve the diagnosis of species of this genus, and in some cases to support the identity of individuals within species.
Systematic problems in Nanhermannia also occur. For example, in the past, N. coronata was confused with N. nana (Nicolet, 1855) sensu Willmann [12], which was clarified by Forsslund [13], and it was confirmed by Solhøy [14,15] that in Norwegian oligotrophic bogs, N. coronata was present. Another example is considering N. coronata by Subías [1] as a junior synonym of N. dorsalis (Banks, 1896), whereas Weigmann [8] and Norton and Ermilov [16] treated it as a separate species, and we agree with the latter opinion.
Identification of Nanhermannia species can also be problematic in ecological investigations. For example, N. coronata can be mistaken for N. sellnicki Forsslund, 1958. An identification of N. coronata is commonly based on the shape of protuberances on the posterior margin of the prodorsum and the number of sclerotized tubercles, which vary within this species, so the ecology of N. coronata given in some papers can be imprecise and needs improving. These species have undoubtedly different ecological preferences; N. sellnicki is less common than N. coronata and occurs in drier habitats [13], like birch forests, especially with understory formed by Vaccinium and Empetrum, while N. coronata is found in moist habitats, especially in raised bogs [15,17,18,19,20,21,22]. The latter species can be very abundant and dominant among the Oribatida [15,19,20,23] and among the mites [20]. For example, in western Norway, among nearly 60,000 mites collected from different peatland microhabitats, and represented by 154 species from all mite orders (Mesostigmata, Trombidiformes and Sarcoptiformes), N. coronata was the most abundant species (it made up 18% of all mite specimens) and occurred in about 90% of samples [20,22]. Such an abundant and common occurrence of N. coronata in peatlands requires better knowledge of the morphology of adults and juveniles, which justifies the need for the current study. The more so that in populations of this species, the juveniles are often very abundant, e.g., in peatlands in Norway, they constituted about 40% of individuals [20,22], so it is very important to include them in ecological analyses.
The morphology of juveniles of Nanhermannia is insufficiently known. According to Norton and Ermilov [15] and Seniczak et al. [2], the full morphological ontogeny of N. comitalis Berlese, 1916, N. cf. coronata, N. nana and N. sellnicki is known, which constitutes nearly 13% of all species. The morphological ontogeny of N. cf. coronata was investigated by Ermilov and Łochyńska [9], but these authors treated the leg setation generally, without labelling of leg setae, which we consider species-specific. Moreover, we found some differences in the morphology of juveniles of this species investigated by these authors and those studied herein, which probably illustrates the morphological variability of the species.
The aim of this paper is to describe the morphological ontogeny of N. coronata and compare it with that of congeners. The identity of specimens from several sites in Norway and Ireland is supported by the COI sequence data. We also add some data on the ecology of this species.

2. Material and Methods

2.1. Morphological and Biological Studies

The adults and juveniles of N. coronata used in morphological and biological studies were collected on 29 and 30 June 2020 (leg. A. Seniczak, K.I. Flatberg, K. Hassel and S. Roth) from an Atlantic raised bog located in Hitra (Hitra municipality, Trøndelag, Norway, 63°29′21.7” N, 8°52′25.1” E, 82 m a. s. l.). In total, 26 samples were collected. These samples were transported in plastic bags in cool boxes for four days to the laboratory of Bydgoszcz University of Science and Technology, Bydgoszcz, Poland, and extracted with Tullgren funnels for ten days into 90% ethanol. In these samples, N. coronata was the only member of Nanhermannia, and therefore, we considered the juveniles to belong to this species. The morphological ontogeny of N. coronata investigated herein is based on the abundant individuals from the transition zone between hummock and hollow in Hitra, but the morphological characters of instars were checked with those from other sites studied herein. We investigated the stage structure of mites, and based on 30 randomly selected adults, we determined the sex ratio, number of gravid females and carried eggs. We also measured the total body length (tip of the rostrum to the posterior edge of the notogaster) in the lateral aspect and the body width (widest part of the notogaster) in the dorsal aspect. In a similar way, we measured the morphological characters of juvenile and adult instars of N. coronata given in Table 1, as well as the size of anal and genital openings and setae perpendicularly to their length in μm. We used the microscopy Nikon Eclipse Ni.
The illustrations are limited to the body regions that show substantial differences between instars and were prepared from individuals mounted temporarily on slides in lactic acid. In the text and figures, we used the following abbreviations: rostral (ro), lamellar (le), interlamellar (in) and exobothridial (exa, exp) setae, bothridium (bo), bothridial seta (bs), notogastral or gastronotal setae (c-, d-, e-, f-, h-, p-series), cupules or lyrifissures (ia, im, ip, ih, ips, iad, ian), cheliceral seta (cha, chb), Trägårdh organ (Tg), palp setae (sup, l, cm, acm, vt, ul, su) and solenidion ω, epimeral setae (1a–c, 2a, 3a–c, 4a–d), genital setae (g), adanal and anal setae (ad-, an-series), leg solenidia (σ, φ, ω), famulus (ε) and setae (bv, ev, d, l, v, ft, tc, pv, a, s, p, u). The leg setae l on femora, and l and v on tarsi were labelled according to their appearance in the ontogeny. The terminology used follows that of Grandjean [24,25,26,27,28] and Norton and Behan-Pelletier [29]. The species nomenclature follows partly Subías [1], Weigmann [8] and Norton and Ermilov [14].
For scanning electron microscopy (SEM), the mites were air-dried and coated with Au/Pd in a Polaron SC502 sputter coater and placed on Al-stubs with double-sided sticky carbon tape. Observations and micrographs were made with a QUANTA FEG 450 scanning electron microscope.

2.2. DNA Barcoding

For molecular studies, we used the specimens of N. coronata collected in raised bogs in southern, central and northern Norway, and Ireland (Table 1). In all locations, samples of Sphagnum mosses of 500 cm3 each were collected and extracted into 90% ethanol in the same way as described above. Additionally, we used publicly available DNA sequences of N. coronata from Finland and some other species of Nanhermannia identified (N. nana from Germany and Slovakia) and N. comitalis from Germany. We used species of putatively close genera as outgroups, P. punctatus (L. Koch, 1879) and Camisia foveolata Hammer, 1955.
Specimens of N. coronata from different locations were sent for DNA barcoding to the Canadian Centre for DNA Barcoding (CCDB) in Guelph, Canada. Each specimen was photographed, and the photos are the vouchers that are available in the Barcode of Life Data System (BOLD, http://boldsystems.org, accessed on 1 October 2023). The specimens were subsequently placed in a well containing 50 mL of 90% ethanol in a 96-well microplate and sent to the CCDB. Mites were sequenced for the barcode region of the COI gene according to standard protocols at the CCDB (www.ccdb.ca, accessed on 1 October 2023, using either LepF1/LepR1 [30] or LCO1490/m HCO2198 [31] primer pairs. The DNA extracts were placed in archival storages at −80 °C, most at the CCDB, and some (with sequencing code starting with UMNFO) at the University Museum of Bergen (ZMBN). Fifteen COI sequences that met the criteria of animal barcodes (sequence length ≥ 500 bp) were obtained. These sequences were blasted against GenBank in order to detect and exclude possible contaminations and were further used in the analyses. The sequences are available in GenBank (accessions numbers in Table 1).
Sequence variation within N. coronata specimens and between species was calculated in BOLD using the Kimura 2 Parameter distance model, pairwise deletion and BOLD Aligner (Amino Acid based HMM). The sequences were aligned by eye and neighbor-joining trees were constructed using MEGA6 [32]. Joint neighborhood topologies were visualized in FigTree 1.4.2 (available at http://tree.bio.ed.ac.uk/software/figtree).

2.3. Ecological Studies

Ecological studies on N. coronata were carried out in Atlantic raised bogs located in Hitra (coordinates were given above) and Høstadmyra (Trondheim municipality, Trøndelag, Norway, 63°24′19.4” N, 10°07′13.5” E, 110 m a. s. l.). Hitra is an island and forms a separate municipality, which is characterized by the mild oceanic climate, with a mean annual temperature of 8 °C and an annual precipitation of 917.8 mm. In the coldest month (January), the average temperature is 1 °C, and in the warmest month (July), it is 16 °C. Høstadmyra is characterized by a slightly colder and drier climate than in Hitra. The mean annual temperature is 6 °C, and annual precipitation is 575.6 mm. The average temperature in the coldest month (January) is −1 °C, and in the warmest month (July), it is 15 °C (https://www.timeanddate.no, accessed on 1 October 2023). In total, 63 samples of Sphagnum mosses of 500 cm3 each were collected on 29 and 30 June 2020 (26 from Hitra and 37 from Høstadmyra) from the following microhabitats: hummocks (31 samples; 12 from Hitra and 19 from Høstadmyra), lawns (14 samples; 4 from Hitra and 10 from Høstadmyra), low part of hummocks (4 samples; 2 from Hitra and 2 from Høstadmyra) and hollows (14 samples; 8 from Hitra and 6 from Høstadmyra). The method of extraction of samples was described above.
Populations of N. coronata from Hitra and Høstadmyra, and from studied microhabitats (hummocks, lawns, low part of hummocks and hollows) were characterized by abundance (A in 500 cm3). The basic statistical descriptors included the mean values and standard deviation. Equality of variance was tested with the Levene test, and normality of the distribution was tested with the Kolmogorov–Smirnov test. As the assumptions of variance analysis were not met, non-parametric tests were employed. Kruskal–Wallis ANOVA by ranks was utilized to test for significant differences between means [33]. The significance level for all analysis was accepted α = 0.05. These calculations were carried out with STATISTICA 12.5 software.

3. Results

3.1. Morphological Ontogeny of Nanhermannia coronata Berlese, 1913 (Figure 1, Figure 2, Figure 3, Figure 4, Figure 5, Figure 6, Figure 7, Figure 8, Figure 9, Figure 10, Figure 11, Figure 12, Figure 13, Figure 14, Figure 15, Figure 16, Figure 17, Figure 18, Figure 19, Figure 20, Figure 21, Figure 22 and Figure 23)

3.1.1. Diagnosis

Adults are of medium size (length 450–660), with characters of Nanhermannia given by Seniczak et al. [2]. The prodorsal seta in is thin, longer than the bothridial seta; protuberances on the posterior part of the prodorsum are highly sclerotized, with 5–7 small posterior tubercles and five pairs of light spots between setal pair in. Seta exp is reduced to its alveolus. The ratio of body length/width is 2.3:1, and the notogastral setae are long, c1, c3, d2 and e1 reaching insertions of setae d1, d2, e1 and h1, respectively. The formula of the epimeral setae is 3-1-3-4. Seta d accompanying solenidion σ on all genua, φ1 on tibia I and φ on other tibiae are present.
Juveniles are elongated, the body unpigmented and with pits, the hysterosoma cylindrical and the central part of the prodorsum, epimeres and legs light brown. The bothridium is small, the bothridial seta absent, seta exp short and exa reduced to its alveolus. The larva has 12 pairs of short gastronotal setae, including f1 and h2; nymphs have 15 pair of long setae, excluding f1 reduced to its alveolus, all setae smooth. Leg segments are relatively thick and oval in cross section, and many setae on tarsi are relatively short, thick and conical, except for the longer apical setae. Seta d accompanying solenidion σ on all genua, φ1 on tibia I and φ on other tibiae are present.
The formula of the genital setae is 1-4-6-9 (protonymph to adult), and femora of deutonymph are 4-4-2-2 (leg I–IV), tritonymph 5-(5-6)-(2-3)-3 and adult 5-7-3-3.
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Figure 1. Nanhermannia coronata, adult, legs partially drawn, scale bars 50 μm. Dorsal aspect.
Figure 1. Nanhermannia coronata, adult, legs partially drawn, scale bars 50 μm. Dorsal aspect.
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Figure 2. Nanhermannia coronata, adult, legs partially drawn, scale bars 50 μm. Ventral aspect.
Figure 2. Nanhermannia coronata, adult, legs partially drawn, scale bars 50 μm. Ventral aspect.
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Figure 3. Nanhermannia coronata, adult. (a) Lateral aspect, legs partially drawn. (b) Posterior part of notogaster, posterior aspect. Mouthparts, right side, antiaxial aspect. (c) Chelicera. (d) Palp. Scale bars (a–c) 50 μm, (d) 20 μm.
Figure 3. Nanhermannia coronata, adult. (a) Lateral aspect, legs partially drawn. (b) Posterior part of notogaster, posterior aspect. Mouthparts, right side, antiaxial aspect. (c) Chelicera. (d) Palp. Scale bars (a–c) 50 μm, (d) 20 μm.
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Figure 4. Nanhermannia coronata, adult, SEM micrographs. (a) Dorsal view, (b) dorsolateral view, (c) lateral view, (d) ventral view.
Figure 4. Nanhermannia coronata, adult, SEM micrographs. (a) Dorsal view, (b) dorsolateral view, (c) lateral view, (d) ventral view.
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Figure 5. Nanhermannia coronata, adult, SEM micrographs. (a–c) Anterior part of body, dorsal view, (d) bothridial seta, lateral view.
Figure 5. Nanhermannia coronata, adult, SEM micrographs. (a–c) Anterior part of body, dorsal view, (d) bothridial seta, lateral view.
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Figure 6. Nanhermannia coronata, adult, SEM micrographs. (a–c) Mouthparts, lateral view, (d) anterior part of body, ventral view.
Figure 6. Nanhermannia coronata, adult, SEM micrographs. (a–c) Mouthparts, lateral view, (d) anterior part of body, ventral view.
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Figure 7. Nanhermannia coronata, adult, SEM micrographs. Ventral view, (a,b) anterior part of body, (c) genital plates, (d) anal plates.
Figure 7. Nanhermannia coronata, adult, SEM micrographs. Ventral view, (a,b) anterior part of body, (c) genital plates, (d) anal plates.
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Figure 8. Nanhermannia coronata, adult, SEM micrographs. (a) Bothridial seta, lateral view; (b) pattern of notogaster; lateral view, (c) adult with rejected ovipositor, (d) ovipositor.
Figure 8. Nanhermannia coronata, adult, SEM micrographs. (a) Bothridial seta, lateral view; (b) pattern of notogaster; lateral view, (c) adult with rejected ovipositor, (d) ovipositor.
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Figure 9. Nanhermannia coronata, 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, genu (l’, v’), tibia (l’, v’), tarsus (pl’, v1’, pv’); (b) leg II, femur (l1’), genu (l’, v’), tibia (l’, v’); (c) leg III, tibia (v’’); (d) leg IV, tibia (v’’); (e) seta d on leg IV; (f) seta l2’’ on femur II (e, f enlarged).
Figure 9. Nanhermannia coronata, 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, genu (l’, v’), tibia (l’, v’), tarsus (pl’, v1’, pv’); (b) leg II, femur (l1’), genu (l’, v’), tibia (l’, v’); (c) leg III, tibia (v’’); (d) leg IV, tibia (v’’); (e) seta d on leg IV; (f) seta l2’’ on femur II (e, f enlarged).
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Figure 10. Nanhermannia coronata, adult, SEM micrographs. Dorsal view, (a) leg I; (b–d) parts of leg I.
Figure 10. Nanhermannia coronata, adult, SEM micrographs. Dorsal view, (a) leg I; (b–d) parts of leg I.
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Figure 11. Nanhermannia coronata, leg I of adult, SEM micrographs. Dorsal view, (a) solenidion σ and seta d on genu I, (b) part of genu, tibia and tarsus I, (c) part of leg I; (d) part of tarsus I, lateral view.
Figure 11. Nanhermannia coronata, leg I of adult, SEM micrographs. Dorsal view, (a) solenidion σ and seta d on genu I, (b) part of genu, tibia and tarsus I, (c) part of leg I; (d) part of tarsus I, lateral view.
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Figure 12. Nanhermannia coronata. Larva, dorsal aspect, legs I and II partially drawn, scale bar 20 μm.
Figure 12. Nanhermannia coronata. Larva, dorsal aspect, legs I and II partially drawn, scale bar 20 μm.
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Figure 13. Nanhermannia coronata. Ventral part of hysterosoma, legs III and VI partially drawn, scale bar 50 μm, (a) larva, (b) protonymph.
Figure 13. Nanhermannia coronata. Ventral part of hysterosoma, legs III and VI partially drawn, scale bar 50 μm, (a) larva, (b) protonymph.
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Figure 14. Nanhermannia coronata, lateral aspect, legs partially drawn, scale bars 50 μm. (a) Larva, (b) tritonymph.
Figure 14. Nanhermannia coronata, lateral aspect, legs partially drawn, scale bars 50 μm. (a) Larva, (b) tritonymph.
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Figure 15. Nanhermannia coronata, leg segments of larva (part of femur to tarsus), right side, scale bar 20 μm. Antiaxial aspect, (a) leg I, tarsus (pl’ not illustrated); (b) leg II; (c) leg III; dorsal aspect, (d) leg I, genu, tibia and part of tarsus.
Figure 15. Nanhermannia coronata, leg segments of larva (part of femur to tarsus), right side, scale bar 20 μm. Antiaxial aspect, (a) leg I, tarsus (pl’ not illustrated); (b) leg II; (c) leg III; dorsal aspect, (d) leg I, genu, tibia and part of tarsus.
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Figure 16. Nanhermannia coronata, scale bars 50 μm. Ventral part of hysterosoma, legs IV partially drawn, (a) deutonymph, (b) tritonymph.
Figure 16. Nanhermannia coronata, scale bars 50 μm. Ventral part of hysterosoma, legs IV partially drawn, (a) deutonymph, (b) tritonymph.
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Figure 17. Nanhermannia coronata, scale bars 50 μm. Tritonymph, dorsal aspect, legs I and II partially drawn.
Figure 17. Nanhermannia coronata, scale bars 50 μm. Tritonymph, dorsal aspect, legs I and II partially drawn.
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Figure 18. Nanhermannia coronata, tritonymph, SEM micrographs. (a) Dorsal view, (b) lateral view, (c), ventral view, (d) anterior and medial part of body.
Figure 18. Nanhermannia coronata, tritonymph, SEM micrographs. (a) Dorsal view, (b) lateral view, (c), ventral view, (d) anterior and medial part of body.
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Figure 19. Nanhermannia coronata, tritonymph, SEM micrographs. Lateral view, (a) anterior part of body, (b) medial part of body, (c) seta h3, (d) leg I and II.
Figure 19. Nanhermannia coronata, tritonymph, SEM micrographs. Lateral view, (a) anterior part of body, (b) medial part of body, (c) seta h3, (d) leg I and II.
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Figure 20. Nanhermannia coronata, leg segments of tritonymph (part of femur to tarsus), right side, seta on the opposite side not illustrated are indicated in the legend, scale bar 10 μm. Antiaxial aspect, (a) leg I, femur (l’), genu (l’), tibia (l’’), tarsus (pl’, v’); (b) leg II, genu (l’), tibia (l’), tarsus (l1’, v1’); (c) leg III, tibia (v’’); (d) leg IV, tibia (v’’); dorsal aspect, (e) genu and tibia I, dorsal view.
Figure 20. Nanhermannia coronata, leg segments of tritonymph (part of femur to tarsus), right side, seta on the opposite side not illustrated are indicated in the legend, scale bar 10 μm. Antiaxial aspect, (a) leg I, femur (l’), genu (l’), tibia (l’’), tarsus (pl’, v’); (b) leg II, genu (l’), tibia (l’), tarsus (l1’, v1’); (c) leg III, tibia (v’’); (d) leg IV, tibia (v’’); dorsal aspect, (e) genu and tibia I, dorsal view.
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Figure 21. Nanhermannia coronata, tritonymph, SEM micrographs. Lateral view, (a–d) parts of leg I.
Figure 21. Nanhermannia coronata, tritonymph, SEM micrographs. Lateral view, (a–d) parts of leg I.
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Figure 22. Nanhermannia coronata, tritonymph, SEM micrographs. Parts of leg I, (a–c) dorsal view; (d) ventral view.
Figure 22. Nanhermannia coronata, tritonymph, SEM micrographs. Parts of leg I, (a–c) dorsal view; (d) ventral view.
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Figure 23. Neighbor-joining tree based on COI sequences of Nanhermannia coronata from southern, central and northern Norway (S Norway, C Norway and N Norway, respectively), Ireland, and Finland, and some other Nanhermannia species; Platynothrus punctatus and Camisia foveolata were used as outgroups. Details on sampling locations are given in Table 1.
Figure 23. Neighbor-joining tree based on COI sequences of Nanhermannia coronata from southern, central and northern Norway (S Norway, C Norway and N Norway, respectively), Ireland, and Finland, and some other Nanhermannia species; Platynothrus punctatus and Camisia foveolata were used as outgroups. Details on sampling locations are given in Table 1.
Animals 13 03590 g023

3.1.2. Morphology of Adult

The adult is elongated (Figure 1, Figure 2, Figure 3a and Figure 4), similar to that described by Berlese [34], but see Remarks. The mean length (and range) of females is 541.7 ± 14.0 (520–556, n = 30); the mean width (and range) is 252.0 ± 6.3 (228–260); males absent. The prodorsal seta in is long and thin, longer than the bothridial seta; protuberances on the posterior part of the prodorsum are highly sclerotized, with 5–7 small posterior tubercles (Figure 1, Figure 3a, Figure 4a–c and Figure 5 and Table 2), and six pairs of light spots present between setal pair in. The ratio of body length/width is 2.3:1. Seta exp short, exa is reduced to its alveolus. The prodorsum and hysterosoma are characterized by pits, observed in SEM figures as small holes (Figure 4, Figure 5, Figure 7c,d, Figure 8b,c and Figure 10a). The notogastral setae are long and smooth, c1, c3, d2 and e1 reaching insertions of setae d1, d2, e1 and h1, respectively (Figure 1, Figure 3a,b, Figure 4a–c, Figure 5a–c and Figure 8c and Table 2). Lyrifissure ia is posterior to seta c3, im anterior to seta e2, ip anterior to seta h2, ian anterior to seta an2, iad anteromedial to seta ad3, and ips and ih pushed anterior and anterolateral to seta ad3, respectively (Figure 2 and Figure 3a). Opisthonotal gland opening gla is not observed among pits. The chelicera is chelate-dentate, and seta cha is located anterior to chb and clearly longer than chb, both smooth (Figure 3c and Figure 6a–c). Most palpal setae are relatively short and smooth (Figure 3d and Figure 6a–c), solenidium ω and eupathidia are short, and the formula of setae (trochanter to tarsus + solenidion ω) is 1-0-2-7(1). Hypostomal and epimeral setae are short and smooth (Figure 2, Figure 4d, Figure 6c,d and Figure 7a,b). Aggenital setae (two pairs) are short, and genital setae (nine pairs) are slightly longer, all smooth (Figure 2, Figure 4d, Figure 7c,d and Figure 8c,d). Adanal setae (three pairs) are long, and anal setae are short, all smooth (Figure 2, Figure 3a,b, Figure 4d, Figure 7d and Figure 8c). The ovipositor is relatively thick, with relatively thick setae (Figure 8c,d). The legs are relatively thick, cuticle with ornamentation (Figure 4, Figure 5a–c, Figure 6c, Figure 7a, Figure 8c, Figure 9, Figure 10 and Figure 11) and all femora oval in cross section. Seta l on trochanter II and d on all femora are barbed, while other leg setae are smooth or finely barbed. Solenidion ω1 on tarsus I is located medial to seta ft’’, whereas solenidia ω2 and ω3 are located anterior to seta a’’. Solenidia ω1 and ω2 on tarsus II are located medial and lateral to seta ft’, respectively. Seta d accompanying solenidion σ on all genua, φ1 on tibia I and φ on other tibiae are present (Figure 9, Figure 10 and Figure 11 and Table 3). In all tarsi, hypertrichy occurs; setae on the basal and medial part of tarsi are conical, those on the distal part normal. The formulae of leg setae (and solenidia), trochanter to tarsus, are I–1-5-5(1)-5(2)-24(3), II–1-7-5(1)-5(1)-23(2), III–4-3-3(1)-4(1)-17 and IV–1-3-3(1)-4(1)-(14-15). Leg tarsi are monodactylous.
Remarks. The mean body length and width of individuals studied herein are larger than those described by Berlese [34]—length 490, width 220—but smaller than those investigated by Sitnikova [5]—length 575, width 250—and Weigmann [8]—length 480–570. Some authors [5,6,9,10] observed on protuberance of N. coronata, 4–5 posterior tubercles of different shape, whereas our adults have 5–7 small tubercles.

3.1.3. Description of Juvenile Stages

The larva is elongated (Figure 12, Figure 13a and Figure 14a), the body unpigmented and with pits and the central part of the prodorsum, epimeres and legs light brown. The prodorsum is subtriangular, the central part punctate and with small pits. Prodorsal setae ro, le, in and exp are short and smooth (Figure 12 and Figure 14a and Table 2), and seta exa is reduced to its alveolus. The mutual distance between setal pair le is slightly shorter than that between setal pair ro, and the mutual distance between setal pair in is about two times longer than that between setal pair ro. The opening of the bothridium is small and rounded, and the bothridial seta is absent. The prodorsum and hysterosoma of the larva have small pits.
The hysterosoma of the larva is cylindrical, and the gastronotum has 12 pairs of setae, including dorsal f1 and ventral h2, inserted lateral to the posterior part of the anal valves (Figure 13a and Figure 14a); most are short and smooth, except for slightly longer f2 and h1 (Table 2). Cupule ia is posterior to seta c3, cupule im anterolateral to seta e2, cupule ip posterolateral to seta f2 and cupule ih lateral to the anterior part of the anal valves (Figure 13a and Figure 14a). The opisthosomal gland opening is medial to seta f2. The anal valves of the larva (segment P) are glabrous. The leg segments are relatively thick, and most leg setae are short, thick and conical, except for the longer apical setae on tarsi (Figure 15). Seta d accompanying solenidion σ on all genua, φ1 on tibia I and φ on other tibiae are present.
The shape and color of nymphs and prodorsal setae are as in the larva, but seta in is clearly longer, and pits in central part of prodorsum denser than in the larva. The bothridium is weakly developed and bothridial seta absent. The gastronotum of the protonymph has small pits and 15 pairs of setae because seta f1 has been lost and only the alveolus of this seta remains, and setae h3 and p-series appear and remain in the deutonymph and tritonymph (Figure 13b, Figure 14b, Figure 16, Figure 17 and Figure 18a–c). The prodorsum and hysterosoma of nymphs have pits, observed in SEM figures as small holes (Figure 18 and Figure 19). All gastronotal setae are long (Table 2) and smooth. In the protonymph, one pair of seta appears on the genital valves, and three pairs are added in the deutonymph and two pairs in the tritonymph (Figure 13b, Figure 14b and Figure 16), all short and smooth. In the deutonymph, one pair of aggenital setae appears, and one pair is added in the tritonymph, all short and smooth (Figure 14b and Figure 16a,b). Anal valves of the protonymph and deutonymph (segments AD and AN, respectively) are glabrous, and those of the tritonymph have two pairs of short and smooth setae. In the tritonymph, the opisthonotal gland opening and cupules ia, im and ip are as in the larva, cupule iad lateral to the anal valves, cupules ips and ih pushed anterolateral to cupule iad (Figure 14b and Figure 16b). The leg segments of the tritonymph relatively thick, and most leg setae are short, thick or conical, except longer apical setae on tarsi (Figure 18, Figure 19a,d, Figure 20, Figure 21 and Figure 22). Seta d accompanying solenidion σ on all genua, φ1 on tibia I and φ on other tibiae are present (Table 3).

3.1.4. Summary of Ontogenetic Transformations

In the larva of N. coronata, the prodorsal setae ro, le, in and exp are short, and in the nymph, seta in is relatively longer, whereas in the adult, setae ro and le are short, in is long and exp is reduced to its alveolus. In all juveniles, the bothridium is weakly developed, and the bothridial seta is absent, whereas in the adult, the bothridium is well developed, with a small, rounded opening, and the bothridial seta is setiform, with a slightly thicker, barbed head. The larva has 12 pairs of gastronotal setae, including f1 and h2, whereas the nymphs and adult have 15 pairs (in the protonymph, f1 is reduced to its alveolus, and h3 and p-series are added). The formula of the gastronotal setae of N. coronata is 12-15-15-15-15 (larva to adult, excluding alveolar f1). The formulae of the epimeral setae are 3-1-2 (larva, including scaliform 1c), 3-1-3-2 (protonymph), 3-1-3-3 (deutonymph) and 3-1-3-4 (tritonymph and adult). The formula of the genital setae is 1-4-6-9 (protonymph to adult), the aggenital setae is 1-2-2 (deutonymph to adult) and the formula of setae of segments PS–AN is 03333-0333-022. The ontogeny of leg setae and solenidia is given in Table 3.

3.2. Results of DNA Barcoding

A neighbor-joining tree based on cytochrome oxidase I (COI) nucleotide sequences confirmed morphological observations that the adults from all included localities (southern, central and northern Norway; Ireland; and Finland) represented the same species (Figure 23). The maximum mitochondrial DNA variation within N. coronata was 0.31%, while the minimum distance to compared representatives of putatively close genera was 26.53%.

3.3. Ecology and Biology

Our data on the ecology of N. coronata indicate that this species is common in raised bogs; it was present in 70% of collected samples. This study also shows wider ecological tolerance of N. coronata towards moisture; this species was found in different bog microhabitats: hummocks, lawns, transition zone between hummocks and hollows, and in hollows (Figure 24). It seems to prefer intermediate moisture conditions; in hummocks, it occurred in 87% of samples and was particularly abundant in the lower zone between hummocks and hollows (Figure 24). Results of Kruskal–Wallis ANOVA by ranks show significant differences between microhabitats (H = 9.27, p = 0.03).
In Hitra, the average abundance of N. coronata was higher (on average 4.42 specimens per 500 cm3) than in Høstadmyra (on average 2.73 specimens per 500 cm3) (Figure 25), but significant differences were observed only in deutonymphs (Kruskal–Wallis ANOVA by ranks: H = 3.96, p = 0.04). In Hitra, the average abundance of deutonymphs was 0.73 specimens per 500 cm3, while in Høstadmyra, it was 0.19 specimens per 500 cm3. In Hitra, the percentage of juveniles was higher than in Høstadmyra (39% and 32% of all individuals of species, respectively).
The stage structure of N. coronata we investigated in both populations of this species from Høstadmyra and Hitra. In total, 213 individuals of N. coronata were found, including 7 larvae, 24 protonymphs, 24 deutonymphs, 21 tritonymphs and 137 adults. Among 30 individuals investigated, all were females, and 25% were gravid, carrying one or rarely two large eggs (215 × 119), constituting about 40% of the total body length of females.

3.4. Comparison of Morphology of Nanhermannia coronata with Congeners

Seniczak et al. [2] compared the morphology of adults of Nanhermannia species, which differ from one another mainly in the shape of posterior prodorsal protuberances and tubercles (from prominent tubercles to small ones), and the length of some setae on the prodorsum and notogaster. Some species differ also from one another by the number of genital setae and formula of epimeral setae, but in most species, these characters are unknown. The shape of posterior prodorsal protuberance and the number of tubercles are considered diagnostic in Nanhermannia, but in some species, the number of tubercles varies [3,4,5,6,7,8,9,10], lowering their diagnostic values. The adult of N. coronata differs from that of N. sellnicki by its darker body color and larger number of posterior tubercles (6–7 tubercles) than that of N. sellnicki (3–4 tubercles), but some authors [5,8,9,10] observed on this protuberance 4–5 posterior tubercles, so the number of these tubercles in these species overlaps and has a small diagnostic value.
In Table 4, we compared selected morphological characters of adults, tritonymphs and larvae of N. coronata, N. comitalis, N. nana and N. sellnicki. The adult of N. coronata differs from that of N. sellnicki by the length of setae in and d1 and the formula of setae on femora ([2], Table 4). The juveniles of these species differ from one another by the length of some setae, the number of posterior tubercles and the formula of femora of the tritonymph. The juveniles of N. coronata differ from those of N. sellnicki by the formula of femora of tritonymph and the shape of insertion of prodorsal seta in and all gastronotal and adanal setae. In the latter species, these setae are in small individual depressions [2], whereas in N. coronata, these depressions are absent. The juveniles of N. nana have clearly longer gastronotal setae than other species, both in the larva and nymphs.

4. Discussion

Nanhermannia coronata is a common and abundant oribatid species in peatlands, but the adult of this species is often mistaken for N. sellnicki, which deteriorates our knowledge on the ecology of both species. According to our observations, the adult of N. coronata differs from that of N. sellnicki by having a darker body color [2], which is diagnostic only for darker adults of N. coronata. Freshly emergent adults of both species are light brown. In our studies, the adult of N. coronata differs from that of N. sellnicki by the length of seta in and the number of posterior tubercles on prodorsal protuberance (5–7 and 3–4 tubercles, respectively), but this character also has a small diagnostic value. Some authors [5,6,9,10] observed on prodorsal protuberance of N. coronata 4–5 posterior tubercles, which overlap with the diagnostic character of N. sellnicki. Diagnostic for these species is the formula of setae on femora, which clearly differentiates the adult and tritonymph of N. coronata studied herein from those instars of N. sellnicki ([2], Table 4). All juvenile stages of N. coronata differ from those of N. sellnicki by the shape of insertion of prodorsal seta in and all gastronotal and adanal setae. In the latter species, these setae are inserted in small individual depressions, whereas in N. coronata, these depressions are absent.
In the juveniles of N. coronata studied herein, some morphological differences were observed, comparing to those investigated by Ermilov and Łochyńska [9]. In the larva studied herein, seta f2 is slightly longer and seta h2 slightly shorter than in that studied by Ermilov and Łochyńska [9], whereas the nymphs have most gastronotal setae slightly longer than those investigated by these authors. The nymphs studied herein have clear posterior prodorsal protuberance, which was not observed by the mentioned authors [9]. These differences may reflect either the geographic variability of juveniles of N. coronata or different methods of preparation of figures.
In N. coronata, seta f1 is lost in the protonymph, as in N. sellnicki [2] and most genera of Crotonioidea, except for Hermannia Nicolet, 1855, Phyllhermannia Berlese, 1916, and Nothrus C.L. Koch, 1836, in which this seta is retained in all instars [36,37,38,39,40,41]. The larva of N. coronata has a pit on dorsal part of genu I, as that of N. sellnicki [2], which Grandjean [27] considered an alveolus of second solenidion σ. Two solenidia on genu I occur in some groups of lower Oribatida, for example in Lohmanniidae [42,43].
The leg segments, setae and solenidia of N. coronata are generally similar as in N. sellnicki except for the pattern of sculpture on femora, the shape of some setae and the number of setae on femur II and tarsi III and IV [2]. In both species, most leg setae are short, thick or conical, and most solenidia are blunt. In these species, an additional seta l occurs on femora II and III, and l and v on most tarsi, as in Platynothrus coulsoni A. and S. Seniczak, 2022, P. punctatus (L. Koch, 1879) and P. troendelagicus Seniczak et al., 2022 [21,44,45]. However, in N. coronata and N. sellnicki, setal pairs l1 and l2 on tarsus II are separated by solenidia ω1 and ω2 and setae ft, whereas in Platynothrus species, solenidia ω1 and ω2 and setae ft are located in anterior position, and all setae l are added posterior to them. In all species of Nanhermannia and Platynothrus, the number of setae on femora of the deutonymph, tritonymph and adult, and the number of setae on tarsi of the adult is species-specific.
The shape of the chelicera and palp of adults of N. coronata is similar to N. sellnicki and other Nanhermannia species discussed by Seniczak et al. [2]. The chelicera is chelate-dentate and has short and thick seta chb located posterior to longer seta cha, and the palpal setae are short and thick, and solenidion ω is separated from seta acm.
Nanhermannia coronata is known from the Holarctic region [46]. It is hygrophilous [47], with some range of tolerance towards moisture [48], but according to Rajski [23], with narrow tolerance towards pH and preferences of its low values. The acidity is considered the main factor for abundant occurrence of N. coronata, and this species has been mainly reported from raised bogs and swamps [15,17,18,19,20,22,49,50,51,52,53,54,55,56,57], less from other acidic forest soils and heaths [8], and it was absent from eutrophic mires [49,58]. Some studies show however that it has been abundant in beech forests [59], deciduous and birch forests [60], and in Scots pine forests [61]. In contrast, in broadleaf forests in Norway, it was few and rare [62,63,64]. In forests and open heath, it was strongly negatively affected by reduced soil moisture [65] and clearly associated with soil (not litter) [66]. It was also found in fruiting bodies of bracket fungi, and its abundance and frequency increased with the degree of decay of fungi [67]. Single individuals were also found in bird plumage and bird nests [68]. It was dominant in 14 Danish spring areas, which may reflect its tolerance to low levels of calcium [69]. In studied gradients in mires, its abundance was neither affected by the moisture nor by the pH, when it ranged between 3.68 and 5.75 [70].
Nanhermannia coronata is a secondary decomposer and fungivorous feeder (feeds partly on fungi and partly on litter) [71,72]. It reproduces parthenogenetically [46]. Ermilov and Łochyńska [9] cultured N. cf. coronata in laboratory conditions (22–23 °C, 100% air humidity) and fed it with algae (Protococcus sp.) and raw potatoes. The adults fed on both types of food, whereas the juveniles preferred algae. The total development of this species lasted 105–124 days, and each stage developed during the following days (+ immovable stage between instars): egg (6–8), larva [(13–20) + (4–8)], protonymph [(11–31) + (5–11)], deutonymph [(12–26) + (6–12)] and tritonymph [(9–26) + (10–13)] to obtain the adult. The maturation time of N. cf. coronata lasts 21 weeks at 20 °C and 16 weeks at 22.5 °C. Its tolerance to temperature was tested in another experiment, and it was similar in adults and juvenile specimens: 38 °C for 4 h and 36 °C for 12 h [73].
Our ecological observations confirmed a common occurrence of N. coronata in raised bogs, with a high percentage of juvenile stages in populations. In two studied locations, Hitra and Høstadmyra, we observed significant differences in the abundance of deutonymphs and proportion of the juvenile stages was higher in Hitra than in Høstadmyra, which can be explained by the warmer (average annual temperature higher by 2 °C) and milder, oceanic climate in Hitra compared to Høstadmyra. More advanced development of species and higher proportion of juveniles has also been observed in another oribatid species, Ceratozetes parvulus Sellnick, 1922, collected from the same bogs [74]. In wetter microhabitats, N. coronata was not abundant and occurred with a low constancy. This is consistent with the observations from inundated bog habitats—edges of water bodies—where N. coronata was absent or very few in number [75]. In degraded, drier bogs, it was more abundant [76,77], but in completely dry bogs, it was absent [78]. When the bog was dried, the abundance of N. coronata decreased in the hummocks but, at the same time, increased in the hollows [79].
Nanhermannia coronata prefers acid and humid microhabitats [23], whereas N. sellnicki is less common than N. coronata and occurs in drier habitats [13], like birch forests, especially with understory formed by Vaccinium and Empetrum. In investigated habitats, like hummocks, lawns, transition zone between hummocks and hollows, and in hollows, N. coronata was particularly abundant in between hummocks and hollows, with abundant juvenile stages.
According to the literature, this species can also be abundant in beech forests [59], deciduous and birch forests [60], Scots pine forests [61], bird plumage and bird nests [68], but these data need confirmation using the diagnostic characters of species given herein.

5. Conclusions

1. Nanhermannia coronata is an abundant and common oribatid mite in raised bogs, with a high percentage of juveniles, and has a preference for humid microhabitats, whereas N. sellnicki is less common than N. coronata and occurs in drier habitats.
2. Nanhermannia coronata differs clearly from N. sellnicki by the following morphological characters: the number of setae on femora I–IV of the adult (N. coronata 5-7-3-3, N. sellnicki 5-6-3-2) and tritonymph [N. coronata 5-(5-6)-(2-3)-3, N. sellnicki 5-6-3-2], and the shape of insertions of prodorsal seta in and all gastronotal and adanal setae of juveniles; in N. sellnicki, these setae are inserted in small individual depressions, whereas in N. coronata, these depressions are absent.

Author Contributions

Conceptualization, S.S. and A.S.; methodology, S.S. and A.S.; line drawings, S.S.; DNA barcoding and SEM imaging, A.S.; writing—original draft, S.S. and A.S.; writing—review and editing, S.S. and A.S. All authors have read and agreed to the published version of the manuscript.

Funding

Norwegian Taxonomy Initiative (Grant No. 6-20, 70184243). The sequencing was financed by Norwegian Barcode of Life (NorBOL).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Ecological data on N. coronata are stored at DataverseNO open archive (https://doi.org/10.18710/F7LO2Y).

Conflicts of Interest

The authors declare no conflict of interest.

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Animals 13 03590 g024
Figure 24. Average abundance of Nanhermannia coronata (individuals per 500 cm3) (bars) with standard deviation (whiskers) in selected microhabitats of bogs in Norway: Hu—hummock, La—lawn, Hu/Ho—transition zone between hummock and hollow, Ho—hollow; numbers above bars present number of samples collected from a certain microhabitat and percentage of samples where the species was present (constancy index).
Figure 24. Average abundance of Nanhermannia coronata (individuals per 500 cm3) (bars) with standard deviation (whiskers) in selected microhabitats of bogs in Norway: Hu—hummock, La—lawn, Hu/Ho—transition zone between hummock and hollow, Ho—hollow; numbers above bars present number of samples collected from a certain microhabitat and percentage of samples where the species was present (constancy index).
Animals 13 03590 g024
Animals 13 03590 g025
Figure 25. Average abundance of developmental stages of Nanhermannia coronata in two bogs in Norway.
Figure 25. Average abundance of developmental stages of Nanhermannia coronata in two bogs in Norway.
Animals 13 03590 g025
Table 1. Information about sequenced specimens of Nanhermannia coronata and other oribatid species used in this study; ad—adult, juv—juveniles.
Table 1. Information about sequenced specimens of Nanhermannia coronata and other oribatid species used in this study; ad—adult, juv—juveniles.
SpeciesSequence Code at BOLDStageGeneBank
Access No.		LocalityCoordinatesElevation
(m a.s.l.)		Collection Data
N. coronataUMNFO663-18adOR732229NO: Vestland, Lydehorn60.370, 5.244216.86 October 2018, Seniczak, A.
UMNFO664-18adOR732235NO: Vestland, Lydehorn60.370, 5.244216.86 October 2018, Seniczak, A.
UMNFO665-18adOR732231NO: Vestland, Lydehorn60.370, 5.244216.86 October 2018, Seniczak, A.
MARBN105-21adOR732225NO: Trondelag, Hitra63.489, 8.87448.229 July 2020, leg. A. Seniczak, K.I. Flatberg, K. Hassel, S. Roth
MARBN106-21adOR732221NO: Trondelag, Hitra63.489, 8.87448.229 July 2020, leg. A. Seniczak, K.I. Flatberg, K. Hassel, S. Roth
MARBN158-21adOR732223NO: Trondelag, Høstadmyra63.405, 10.12110.030 July 2020, leg. A. Seniczak, K.I. Flatberg, K. Hassel, S. Roth
MARBN159-21adOR732230NO: Trondelag, Høstadmyra63.405, 10.12110.030 July 2020, leg. A. Seniczak, K.I. Flatberg, K. Hassel, S. Roth
MARBN160-21adOR732234NO: Trondelag, Høstadmyra63.405, 10.12110.030 July 2020, leg. A. Seniczak, K.I. Flatberg, K. Hassel, S. Roth
MARBN516-22adOR732224NO: Nordland, Kummeren67.043, 14.21628.126 July 2021, Seniczak, A., Flatberg, K.I.
MARBN517-22adOR732218NO: Nordland, Kummeren67.043, 14.21628.126 July 2021, Seniczak, A., Flatberg, K.I.
MARBN519-22adOR732227NO: Nordland, Kummeren67.043, 14.21628.126 July 2021, Seniczak, A., Flatberg, K.I.
MARBN343-21adOR732226IR: Leinster, Lullymore53.270, −6.94977.49 December 2014, leg. A. Seniczak, T. Bolger
MARBN344-21adOR732233IR: Leinster, Lullymore53.270, −6.94977.49 December 2014, leg. A. Seniczak, T. Bolger
MARBN345-21adOR732232IR: Leinster, Lullymore53.270, −6.94977.49 December 2014, leg. A. Seniczak, T. Bolger
MARBN346-21adOR732228IR: Leinster, Lullymore53.270, −6.94977.49 December 2014, leg. A. Seniczak, T. Bolger
FINOR987-16adMZ608481FIN: Varsinais-Suomi: Paimio, Jaervessuo60.451, 22.77549.99 October 2014, leg. R. Penttinen
FINOR988-16adMZ609187FIN: Varsinais-Suomi: Paimio, Jaervessuo60.451, 22.77549.99 October 2014, leg. R. Penttinen
FINOR989-16adMZ611116FIN: Varsinais-Suomi: Paimio, Jaervessuo60.451, 22.77549.99 October 2014, leg. R. Penttinen
N. nanaGBMYR861-15adIn BOLDGE:
UZINS170-23adIn BOLDSL: Bratislava, Sitina48.171, 17.0656213.31 June 2022, leg. Mangova, B.
UZINS171-23adIn BOLDSL: Bratislava, Sitina48.171, 17.0656213.31 June 2022, leg. Mangova, B.
Camisia foveolataMARBN334-21adOR732222NO: Svalbard, Longyearbyen, Endalen78.209, 15.71122.65 June 2018, Roth, S.
MARBN335-21juvOR732220NO: Svalbard, Longyearbyen, Endalen78.209, 15.71122.65 June 2018, Roth, S.
MARBN341-21adOR773185NO: Vestland, Finse60.593, 7.4321352.07 September 2018, Seniczak, A.
Platynothrus punctatusMARBN097-21adOL671034NO: Vestland, Finse60.593, 7.4321352.022 September 2019, Seniczak, A.
MARBN098-21adOL671021NO: Vestland, Finse60.593, 7.4321352.022 September 2019, Seniczak, A.
MARBN365-21juvOL671024SP: Andalusia, Borreguil de la Virgen37.087, −3.3742500.718 August 2017, Seniczak, A., F. Ondoño, E.
Table 2. Measurements of some morphological characters of juvenile stages and adult of Nanhermannia coronata (mean measurements of 10 specimens in μm); nd—not developed.
Table 2. Measurements of some morphological characters of juvenile stages and adult of Nanhermannia coronata (mean measurements of 10 specimens in μm); nd—not developed.
Morphological CharactersLarvaProtonymphDeutonymphTritonymphAdult
Body length284351416572559
Body width125155172305251
Length of prodorsum104120152171215
Length of:
seta ro1617213239
seta le1517193035
seta in1923304575
seta bsndndndnd67
seta c112193057129
seta c314223182136
seta cp10203295142
seta d112213370125
seta d213223293135
seta e110203588130
seta e211213883134
seta f18lostlostlostlost
seta f224324396133
seta h120223080120
seta h3nd233195133
seta p1nd81972122
seta p3nd163083130
genital openingnd26334683
anal opening526577123114
Table 3. Ontogeny of leg setae (Roman letters) and solenidia (Greek letters) in Nanhermannia coronata.
Table 3. Ontogeny of leg setae (Roman letters) and solenidia (Greek letters) in Nanhermannia coronata.
LegTrochanterFemurGenuTibiaTarsus
Leg I
Larva–d, bv’’d, (l), σ(l), v’, d, φ1(ft), (tc), (pl), (p), (u), (a), s, (pv), ε, ω1
Protonymph––––ω2
Deutonymphv’(l)–φ2–
Tritonymph–v”v’, v”v’’(v1), (it), ω3
Adult––––(l), (v2)
Leg II
Larva–d, bv’’d, (l), σl’, v’, d, φ(ft), (tc), (p), (u), (a), s, (pv), ω1
Protonymph– –l’’–
Deutonymphv’(l1)––ω2
Tritonymph–l2”, v” 1v’, v”v’’l1′, (v1), (it)
Adult–l3′––l1”, (l2), (v2)
Leg III
Larva–d, ev’d, σv’, d, φ(ft), (tc), (p), (u), (a), s, (pv)
Protonymphv’––––
Deutonymphl’1––––
Tritonymphl’2l’ 1v’v’’(it), (v1)
Adultl’3––l’–
Leg IV
Protonymph –––ft’’, (pv), (p), (u)
Deutonymph–d, ev’l’, d, σv’, d, φ(tc), (a), s
Tritonymphv’l’v’v”(v1)
Adult–––l’ft’ 2
Note: structures are indicated where they are first added and are present through the rest of ontogeny; pairs of setae are in parentheses; dash indicates no additions; 1 is added in some individuals; if not, it is added in the next stage; 2 in some individuals is absent.
Table 4. Comparison of selected morphological characters of some instars of Nanhermannia coronata, N. comitalis, N. nana and N. sellnicki.
Table 4. Comparison of selected morphological characters of some instars of Nanhermannia coronata, N. comitalis, N. nana and N. sellnicki.
CharactersN. coronataN. comitalis 1N. nana 1N. sellnicki 2
Adult
Length of seta inLonger than bsLonger than bsAs long as bsAs long as bs
Posterior tubercles 35–7413–4
Length of seta d1 4NoYesYesNo
Formula of femora5-7-3-3unknownunknown5-6-3-2
Tritonymph
Length of seta inAs long as c1As long as c1Shorter than c1As long as c1
Posterior tubercles 36–7564–5
Length of seta c1 5NoYesYesNo
Length of seta d1 4NoYesYesNo
Length of seta e1 6NoNoYesNo
Formula of femora5-(5-6)-(2-3)-3unknownunknown5-6-3-2
Larva
Length of seta inShorter than c1Longer than c1Shorter than c1As long as c1
Posterior tubercles 3Absent2–32Absent
Length of seta c1 5NoYesYesNo
Length of seta d1 4NoYesYesNo
1 According to Seniczak [35], 2 Seniczak et al. [2], 3 on each posterior prodorsal protuberance, 4 reaches insertion of seta e1, 5 reaches insertion of seta d1, 6 reaches insertion of seta h1.
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Seniczak, S.; Seniczak, A. Morphological Ontogeny and Ecology of a Common Peatland Mite, Nanhermannia coronata (Acari, Oribatida, Nanhermanniidae). Animals 2023, 13, 3590. https://doi.org/10.3390/ani13223590

AMA Style

Seniczak S, Seniczak A. Morphological Ontogeny and Ecology of a Common Peatland Mite, Nanhermannia coronata (Acari, Oribatida, Nanhermanniidae). Animals. 2023; 13(22):3590. https://doi.org/10.3390/ani13223590

Chicago/Turabian Style

Seniczak, Stanisław, and Anna Seniczak. 2023. "Morphological Ontogeny and Ecology of a Common Peatland Mite, Nanhermannia coronata (Acari, Oribatida, Nanhermanniidae)" Animals 13, no. 22: 3590. https://doi.org/10.3390/ani13223590

APA Style

Seniczak, S., & Seniczak, A. (2023). Morphological Ontogeny and Ecology of a Common Peatland Mite, Nanhermannia coronata (Acari, Oribatida, Nanhermanniidae). Animals, 13(22), 3590. https://doi.org/10.3390/ani13223590

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