# A new water mite species of the genus Teutonia Koenike, 1889 from Corsica, France, based on morphological data and DNA barcodes (Acari, Hydrachnidia, Teutoniidae)

Vladimir Pešić, Harry Smit

► To cite this version:

Vladimir Pešić, Harry Smit. A new water mite species of the genus Teutonia Koenike, 1889 from Corsica, France, based on morphological data and DNA barcodes (Acari, Hydrachnidia, Teutoniidae). *Acarologia*, 2022, 62 (2), pp.465-473. 10.24349/8n5y-mw7l. hal-03667474

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# A new water mite species of the genus *Teutonia* Koenike, 1889 from Corsica, France, based on morphological data and DNA barcodes (Acari, *Hydrachnidia*, *Teutoniidae*)

Vladimir Pešićª, Harry Smitᵇ

## Original research

### ABSTRACT

We describe a new species, *Teutonia corsicana* sp. nov. from Corsica, France using morphological data and DNA barcodes. Morphologically the new species can be separated on the basis of the female pregenital sclerite partially covered by genital flaps, and in both sexes a broad-based, triangular ventral projection of P-2 and slender P-3 (L/H ratio 1.6-1.7). The average K2P-distance between DNA-barcode sequences of *T. corsicana* sp. nov. and widely distributed *T. cometes* (Koch, 1837) was estimated at 18.7±1.9 %. An updated key for the European species of *Teutonia* is provided.

**Keywords** water mites; new species; taxonomy; species delimitation; France

**Zoobank** http://zoobank.org/F83B5FED-0C63-41BC-A2F5-A331E03CD95E

## Introduction

The family Teutoniidae Koenike, 1910 comprises of two genera, the monotypic *Limnolegeria* Motaş, 1928, which was for a long time considered as a representative of the Anisitsiellidae (see Cook 1974), and the widely distributed *Teutonia* Koenike, 1889. The latter genus, recorded from the Holarctic, comprises of two subgenera, the nominate subgenus and *Subteutonia* Habeeb, 1958 known only from Canada (Smit 2020).

Currently, two species of the genus *Teutonia* are known from the Western Palaearctic (Di Sabatino *et al.* 2010), i.e., the widely distributed *T. cometes* (Koch, 1837), inhabiting cold montane lakes and pools of shaded streams (Di Sabatino *et al.* 2010), and *T. subalpina* Thor, 1897. The latter, little known species is restricted to the far north of Europe (alpine and subalpine coniferous forests of Norway and Sweden) (Di Sabatino *et al.* 2010).

In this paper we used morphological data and DNA barcode of the mitochondrial cytochrome *c* oxidase subunit I (COI) gene sequence to describe one new species of the genus *Teutonia* from Corsica.

## Material and methods

Water mites were collected by hand netting by the junior author, sorted live in the field, and immediately preserved in 96% ethanol for the purpose of the molecular analyses. A part of the material was also preserved in Koenike-fluid. After DNA extraction, two voucher specimens were dissected and slide mounted in Faure's medium. Holotype and paratypes of the new

Received 11 March 2022

Accepted 05 May 2022

Published 09 May 2022

Corresponding author

Vladimir Pešić:

vladopesic@gmail.com

Academic editor

Makol, Joanna

https://doi.org/
10.24349/8n5y-mw7l

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

<figure><img src="image_7.png" /></figure>

Pešić V. and Smit H.

Licensed under

Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote>ªDepartment of Biology, University of Montenegro, Cetinjski put b.b., 81000 Podgorica, Montenegro.</footnote>

<footnote>ᵇNaturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands.</footnote>
species are deposited in Naturalis Biodiversity Center in Leiden (RMNH). The DNA sequences prepared in the course of this study were deposited in the Barcode of Life Data System (BOLD) and GenBank (Table 1).

All measurements are in µm. Morphological nomenclature follows Gerecke *et al.* (2016). The length of genital flaps and acetabula were measured on both sides, and therefore their dimensions are given as a range. The photograph of ejaculatory complex was made using a camera on a Samsung Galaxy smartphone. The following abbreviations are used: Ac – acetabula; a.s.l. – above sea level; Cx-I-IV – first-fourth coxeae; dL – dorsal length; H – height; I-IV -L-1-6 – first-sixth segments of first- fourth legs; L – length; P-1-P-5 – palp segment 1-5; RMNH – Naturalis Biodiversity Center, Leiden; vL – ventral length; W – width.

Molecular analyses were conducted at the Canadian Centre for DNA Barcoding (Guelph, Ontario, Canada (CCDB; http://ccdb.ca/)), where the specimens were sequenced for the barcode region of COI using standard invertebrate DNA extraction, amplification and sequencing protocols (Ivanova *et al.* 2007; Ivanova and Grainger 2007a, b). In CCDB the chromatograms were assembled into consensus sequences for each specimen and uploaded to the barcode of life database (BOLD; https://www.boldsystems.org/). The sample identifiers in BOLD are given for each barcoded specimen.

The final dataset for the phylogenetic analysis includes 17 COI sequences representing *Teutonia cometes* (n=7), and *T. corsicana* sp. nov. (n=10). *Limnolegeria longiseta* Motaş, 1928 (BOLD/GenBank SEPTA083-21/OM321062) was used as outgroup (Table 1).

Sequence comparisons were performed using MUSCLE alignment (Edgar 2004). Intra- and interspecific genetic distances were calculated based on the Kimura 2-parameter model (K2P; Kimura 1980), using MEGA-X (Kumar *et al.* 2018). MEGA-X software was used to calculate Neighbour-Joining (NJ) trees based on K2P distances (standard for barcoding studies) and pairwise deletion of missing data. The support for tree branches was calculated by the non-parametric bootstrap method (Felsenstein 1985) with 1000 replicates. Codon positions included were 1st+2nd+3rd+Noncoding. All ambiguous positions were removed for each sequence pair.

<table><caption>**Table 1** List of sequenced specimens used in this study.</caption><thead><tr><th>Locality (country, name)</th><th>Lat/Long</th><th>BOLD/GenBank Acc. Nos</th><th>Voucher code</th></tr></thead><tbody><tr><td colspan="4"><strong>Teutonia cometes (Koch, 1837)</strong></td></tr><tr><td>Norway, Trondheim, Bymarka, Lauglovatnet, st. 2</td><td>63.3617° N, 10.2410° E</td><td>MMHYD003-19</td><td>NTNU-VM 228621</td></tr><tr><td>Norway, Trondheim, Bymarka, Store Leirsjoe, st. 3</td><td>63.3862° N, 10.2872° E</td><td>MMHYD012-19</td><td>NTNU-VM 228623</td></tr><tr><td>Norway, Stjoerdal, Stjoerdalsvassdraget, Fugla, st. 1</td><td>63.4430° N, 10.9235° E</td><td>MMHYD060-19</td><td>NTNU-VM 228625</td></tr><tr><td>Norway, Orkdal, Orklavassdraget, Gjoesjoeen, st. 1</td><td>63.2349° N, 9.6626° E</td><td>MMHYD069-19</td><td>NTNU-VM 228627</td></tr><tr><td>Montenegro, Podgorica, Mareza canal</td><td>42.4790° N, 19.1813° E</td><td>DNAEC068-20/ OL870217</td><td>33. M19_20_3_F11</td></tr><tr><td>Montenegro, Danilovgrad, spring Svinjška vrela</td><td>42.6384° N, 19.0074° E</td><td>DNAEC079-20/ OL870180</td><td>Hyd_MN_VP5</td></tr><tr><td>Germany, Baden-Württemberg, Schönbuch, Goldersbach near Teufelsbrücke</td><td>-</td><td>JN018321</td><td>MNHN-JAC69</td></tr><tr><td colspan="4"><strong>Teutonia corsicana sp. nov.</strong></td></tr><tr><td>France, Corsica, Porto, Ruisseau d'Enova at crossing with road D124</td><td>42.2659° N, 8.7141° E</td><td>NLACA1050-17</td><td>RMNH.5070808</td></tr><tr><td>France, Corsica, Ruisseau de Tuara, crossing road D81</td><td>42.3496° N, 8.6650° E</td><td>NLACA1051-17</td><td>RMNH.5070809</td></tr><tr><td>France, Corsica, Ruisseau de Tuara, crossing road D81</td><td>42.3496° N, 8.6650° E</td><td>NOVMB001-21/ OM952168</td><td>CCDB 38559 A01</td></tr><tr><td>France, Corsica, Ruisseau de Tuara, crossing road D81</td><td>42.3496° N, 8.6650° E</td><td>NOVMB002-21/ OM952167</td><td>CCDB 38559 A02</td></tr><tr><td>France, Corsica, Ruisseau de Sattu</td><td>41.6907° N, 9.1525° E</td><td>NOVMB033-21/ OM952166</td><td>CCDB 38559 C09</td></tr><tr><td>France, Corsica, Riviere La Solenzara</td><td>41.8383° N, 9.3306° E</td><td>NOVMB048-21/ OM952165</td><td>CCDB 38559 D12</td></tr><tr><td>France, Corsica, Riviere La Solenzara</td><td>41.8383° N, 9.3306° E</td><td>NOVMB049-21/ OM952164</td><td>CCDB 38559 E01</td></tr><tr><td>France, Corsica, Ruisseau de Battesta at crossing with road D50</td><td>41.5449° N, 9.0229° E</td><td>NOVMB068-21/ OM952163</td><td>CCDB 38559 F08</td></tr><tr><td>France, Corsica, Ruisseau de Battesta at crossing with road D50</td><td>41.5449° N, 9.0229° E</td><td>NOVMB069-21/ OM952162</td><td>CCDB 38559 F09</td></tr><tr><td>France, Corsica, Ruisseau d' Enova, at crossing with road D124</td><td>42.2659° N, 8.7141° E</td><td>NOVMB076-21/ OM952161</td><td>CCDB 38559 G04</td></tr><tr><td colspan="4"><strong>Limnolegeria longiseta Motaş, 1928</strong></td></tr><tr><td>Turkey, Burdur, Bucak at Kizilli Village</td><td>37.3425° N, 30.9256° E</td><td>SEPTA083-21/ OM321062</td><td>CCDB 38363 G11</td></tr></tbody></table>
<figure><img src="image_8.png" /></figure>

**Figure 1** Neighbour-Joining tree of the genus *Teutonia*, obtained from 17 nucleotide COI sequences and the results of species delimitation analyses. Values near branches show bootstrap support (BS). The results of species delimitation by ASAP procedure are indicated by vertical bars. Country codes (alpha-2 code): DE – Germany, FR – France, MN – Montenegro, NO – Norway, TR – Turkey.

In order to assess the genetic differentiation of species we used the online ASAP (assem-
ble species by automatic partitioning) version (https://bioinfo.mnhn.fr/abi/public/asap/asapweb.html) with default settings and the K2P distance model. The latter procedure was
designated to a list of partitions of species hypotheses using genetic distances, calculated
between DNA sequences and ranked by their ASAP-scores: the lower the score, the better the
partition (Puillandre et al. 2021).

**Results and discussion**

**Species delimitation using DNA-barcodes**

The final alignment for species delimitation using COI sequence data included 17 *Teutonia*
specimens listed in Table 1 and one outgroup, *Limnolegeria longiseta* from Turkey. The final
alignment consisted of 837 nucleotide positions. The NJ tree is presented in Figure 1.

The sequences retrieved from the *Teutonia* specimens from Corsica appeared as a sister clade of the clade that encompasses specimens of *T. cometes* from Norway, Germany and Montenegro (Figure 1). The average K2P genetic distance between *T. corsicana* sp. nov. from Corsica and *T. cometes* was estimated 18.7±1.9%. Both species possesses a low intraspecific distance, 0.15±0.07% K2P for *T. corsicana* sp. nov. and 0.36±0.15% K2P for *T. cometes*.
<figure><img src="image_9.png" /><figcaption>**Figure 2** Results of ASAP analysis for COI sequences. (A) Distribution of pairwise differences, (B) Ranked pairwise differences.</figcaption></figure>

The ASAP method observed a barcode gap of about 1-18% K2P distances (Figure 2A-B). The applied ASAP procedure identified two MOTUs (hypothetical species) corresponding to *T. cometes* and *T. corsicana* sp. nov. at the threshold distance of 9.51% (K2P) which has the best ASAP-score (1.00) within the available molecular data.

**Systematics**

**Family Teutoniidae Koenike, 1910**

**Diagnosis** — Smit 2020: 268.

**Genus Teutonia Koenike, 1889**

**Diagnosis** — Smit 2020: 269.

**Teutonia corsicana sp. nov.**

Zoobank: 5877B3D9-E264-4102-A442-3356BCE46C10

Figures 3-4

**Material examined**

Holotype ♀, France, Corsica, Ruisseau de Tuara, crossing road D81, 42.3496° N, 8.66497° E, 238 m a.s.l., 14 Apr. 2015, leg. Smit (sequenced [voucher code: CCDB 38559 A01; BOLD/GenBank Nos.: NOVMB001-21/OM952168]), dissected and slide mounted. Paratype: 1 ♀, same place and data as holotype, sequenced (Table 1); 1♂, 1♀, Ruisseau de Sattu, Corsica, 41.6907° N, 9.1525° E, 370 m a.s.l., 10 Apr. 2015, leg. Smit, 1♂ sequenced (Table 1); 1♂, 1♀, Rivière La Solenzara, Corsica, 41.8383° N, 9.33055° E, 90 m a.s.l., 7 Apr. 2015, leg. Smit, 1♂, 1♀ sequenced (Table 1); 2♂, Ruisseau de Battesta at crossing with road D50, Corsica, 41.5449° N, 9.02292° E, 220 m a.s.l., 8 Apr. 2015, leg. Smit, 2♂ sequenced (Table 1), 1♂ (voucher code: CCDB 38559 F09) dissected and slide mounted; 2♀, Ruisseau de Enova, at crossing with road D124, Corsica, 42.2659° N, 8.71405° E, 105 m a.s.l., 13 Apr. 2015, leg. Smit, 1♀ sequenced (Table 1).
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Teutonia corsicana* sp. nov. (A-B, D-G - holotype ♀, Ruisseau de Tuara, France; C - ♂ [CCDB 38559 D12], preserved specimen, Riviere La Solenzara, France); A - coxal and genital field; B, C - genital field; D - palp, medial view (P-1 lacking); E - palp, lateral view; F - I-L-5 and -6; G - IV-L-5 and -6. Scale bars = 100 µm.</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Figure 4** *Teutonia corsicana* sp. nov., ♂ [CCDB 38559 F09], Ruisseau de Battesta, France: A – coxal and genital field, partial view; B – photograph of ejaculatory complex; C – palp, medial view; D – gnathosoma. Scale bars = 100 µm.</figcaption></figure>

**Other material**

France, Corsica: Ruisseau de Battesta, at crossing with road D50, 41.54487° N, 9.02292° E, 220 m a.s.l., 8 Apr. 2015, 3♀, 1 deutonymph; Ruisseau de Campaghiu, Porto, 42.24128° N, 8.73240° E, 188 m a.s.l., 11 Apr. 2015, 1♀, 1 deutonymph; Ruisseau d'Aitone, at crossing with road D84, 42.26365° N, 8.84597° E, 1115 m a.s.l., 12 Apr. 2015, 1♀; Ruisseau de Casa Infurcata, at crossing with road D124, 42.25600° N, 8.76513° E, 217 m a.s.l., 13 Apr. 2015, 1♀; Ruisseau d'Enova, at crossing with road D124, 42.26590° N, 8.71405° E, 105 m a.s.l., 13 Apr. 2015, 1♂, 5♀, 2 deutonymphs; Rivière Le Fango, at crossing with road D81, 42.41503° N, 8.68300° E, 6 m a.s.l., 14 Apr. 2015, 1♀, 4 deutonymphs; Ruisseau de Tuara, at crossing with road D81, 42.34958° N, 8.66497° E, 230 m a.s.l., 14 Apr. 2015, 3♀, 4 deutonymphs; tributary of Ruisseau de Canne, at crossing with road D81, 42.38307° N, 8.69903° E, 225 m a.s.l., 14 Apr. 2015, 1♀, 2 deutonymphs; Rivière de Tartagine, at Maison Forestière Tartagine, 42.49410° N, 8.99288° E, 706 m a.s.l., 16 Apr. 2015, 1♀; unnamed stream N of Suarella, at crossing with road D18, 42.39033° N, 9.13762° E, 387 m a.s.l., 17 Apr. 2015, 1♀.
**Diagnosis**

Female gonopore pregenital sclerite partly covered by genital flaps; in both sexes P-2 with broad-based ventrodistal projection, P-3 slender (L/H ratio 1.6-1.7).

**Description**

**General features** — Integument finely striated. Coxae medially touching, but not fused, with a few setae. Ac-1-2 rectangular, Ac-3 roundish. P-2 with a generally broad-based ventrodistal projection, P-3 with nearly straight, or slightly convex ventral margin, P-3 lateral seta long, shifted to medial surface, P-4 slender, bearing several fine hair-like setae, mainly in anteroventral part. Legs: claws with claw blade and ventral clawlets; swimming setae restricted to III-/IV-L-3-4, numbers: III-L-3, 3; III-L-4, 8; III-L-5, 8-9; IV-L-3, 3; IV-L-4, 8-9; IV-L-5, 11-12. *Male* — Glandularium on Cx-IV completely surrounded by coax sclerotization. Genital field: genital flaps medially with a dense cover of fine setae, pregenital sclerite small, postgenital sclerite as wide as genital field. *Female* — Glandularium on Cx-IV partly surrounded by coax sclerite, medially in contact with the perigenital membrane. Genital field: genital flaps with a row of regularly arranged medial setae, pregenital sclerite large, crescent shaped, partly covered by genital flaps (but only slightly covered in Koenike-fixed specimens), gonopore shorter than the genital flaps, postgenital sclerite shorter than pregenital sclerite.

**Description** — *Female* (holotype) — Idiosoma L 1190, W 905. Coxal field: L 581, W 878; Cx-III W 563; L Cx-I+II 228, Cx-III+IV 434. Genital flaps L 177-181; pregenital sclerite W 200, postgenital sclerite W 106; L Ac-1-3: 59, 56-60, 39-43. Eggs maximum diameter (n=2) 173-178.

Palp total L 684, dL/H, dL/H ratio: P-1, 33/44, 0.75; P-2, 142/117, 1.21; P-3, 164/102, 1.61; P-4, 289/41, 7.1; P-5, 56/15, 3.8; L ratio P-2/P-4 0.49. Gnathosoma vL 194; chelicera L 256, basal segment L 219, claw L 44, L ratio chelicera basal segment/claw 5.0. Leg segments dL: I-L-2-6: 122, 166, 228, 244, 253; IV-L-1-6: 156, 156, 200, 253, 264, 331.

*Male* (paratype, CCDB 38559 F09) — Idiosoma L 750, W 588. Coxal field: L 495, W 703; Cx-III W 459; L Cx-I+II 197, Cx-III+IV 380. Genital field L/W 166/147, genital flaps L 156-163, pregenital sclerite W 119; L Ac-1-3: 50, 52-53, 34-38. Ejaculatory complex (Figure 3B) L 192.

Palp total L 581, dL/H, dL/H ratio: P-1, 28/36, 0.77; P-2, 120/96, 1.25; P-3, 142/84, 1.69; P-4, 244/34, 7.2; P-5, 47/14, 3.3; L ratio P-2/P-4 0.49. Gnathosoma vL 165; chelicera L 230, basal segment L 186, claw L 44, L ratio chelicera basal segment/claw 4.3. Leg segments dL: I-L-1-6: 71, 111, 135, 191, 200, 209; IV-L-1-6: 131, 144, 175, 215, 236, 284.

**Etymology**

Named after the island where the new species was discovered.

**Remarks**

The female of the new species from Corsica, is easily distinguished from the two other known species of the genus, *T. cometes* and *T. subalpina* in the shape of genital field having a pregenital sclerite partly covered by the genital flaps. Moreover, in both sexes *T. cometes* differs in P-2 generally with a narrow-based ventrodistal projection (but in larger populations, the latter projection appearing broad-based in some specimens, especially in juveniles) and stouter P-3 (L/H ratio 1.35-1.5 vs. 1.6-1.7 in *T. corsicana* sp. nov.). The high genetic distance between the new species and *T. cometes* (COI 18.7±1.9 K2P%) suggests a long independent history of the new species. The former records of *T. cometes* from Corsica (Angelier 1959; Santucci 1971) likely should be assigned to the new species.

The little known *T. subalpina* can be distinguished in the female by the very long gonopore, anteriorly distinctly extending over the margin of the genital flaps (Di Sabatino *et al.* 2010). The taxonomic status of *T. subalpina* is still unclear, and according to Di Sabatino *et al.* (2010),
## Acknowledgements

We are indebted to Elisabeth Stur (Trondheim, Norway) and Reinhard Gerecke (Tübingen, Germany) for permission to use some unpublished sequences from Norway (Project "Water M&M (University of Trondheim)"). Special thanks to Milica Jovanović and Ana Manović (University of Podgorica) for their excellent routine laboratory processing work. HS would like to thank Truus van der Pal (Alkmaar) for her assistance with the field work on Corsica. This study is part of the "DNA-Eco" scientific project, supported by a grant of the Montenegrin Ministry of Science. We thank Joanna Mąkol (Wrocław), Hiroshi Abé (Fujisawa) and one anonymous reviewer, whose constructive comments greatly improved this work.

## References

* Angelier E. 1959. *Acariens (Hydrachnellae et Porohalacidae) des eaux superficielles*. In: Angelier E. et collaborateurs, *Hydrobiologie de la Corse. Vie et Milieu, Suppl.*, 8: 64-148.

* Cook D.R. 1974. Water mite genera and subgenera. Mem. Am. Entomol. Inst., 21: 1-860.

* Di Sabatino A., Gerecke R., Gledhill T., Smit H. 2010. Chelicerata: Acari II. In: Gerecke R. (ed.) Chelicerata: Araneae, Acari I. Süßwasserfauna von Mitteleuropa, 7, 2-2: 1-134. München: Elsevier Spektrum Akademischer Verlag. https://doi.org/10.1007/978-3-8274-2266-8_1

* Edgar R.C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high 679 throughput. Nucleic acids Res 32(5): 1792-1797. https://doi.org/10.1093/nar/gkh340

* Felsenstein J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39: 783-791. https://doi.org/10.1111/j.1558-5646.1985.tb00420.x

* Gerecke R., Gledhill T., Pešić V., Smit H. 2016. Chelicerata: Acari III. In: Gerecke R. (ed.) Süßwasserfauna von Mitteleuropa, Bd. 7/2-3. Springer-Verlag Berlin, Heidelberg, pp. 1-429. https://doi.org/10.1007/978-3-8274-2689-5

* Ivanova N.V., de Waard J.R., Hebert P.D.N. 2007. CCDB protocols, glass fiber plate DNA extraction. ccdb.ca/site/wp-content/uploads/2016/09/CCDB_DNA_Extraction.pdf

* Ivanova N.V., Grainger C.M. 2007a. CCDB protocols, COI amplification. Available at: ccdb.ca/site/wp-content/uploads/2016/09/CCDB_Amplification.pdf

* Ivanova N.V., Grainger C.M. 2007b. CCDB protocols, sequencing. Available at: ccdb.ca/site/wp-content/uploads/2016/09/CCDB_Sequencing.pdf

* Kimura M. 1980. A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. J. Mol. Evol., 16: 111-120. https://doi.org/10.1007/BF01731581

* Kumar S., Stecher G., Li M., Knyaz C., Tamura K. 2018. MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Mol. Biol. Evol., 35: 1547-1549. https://doi.org/10.1093/molbev/mxy096

* Puillandre N., Brouillet S., Achaz G. 2021. ASAP: assemble species by automatic partitioning. Mol. Ecol. Resour., 21:609-620. https://doi.org/10.1111/1755-0998.13281

* Santucci J. 1971. Contribution à l'étude de la répartition des Hydracariens (Hydrachnellae) des eaux superficielles d'un torrent de Corse - Le Porto. Ann. Fac. Sci. Marseille, 45: 81-99.

* Smit H. 2020. Water mites of the world with keys to the families, subfamilies, genera and subgenera (Acari: Hydrachnidia). Monogr. Nederl. Entom. Veren., 12: 1-774.