Milnesium matheusi

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Kaczmarek Ł, Grobys D, Kulpa A, Bartylak T, Kmita H, Kepel M, Kepel A, Roszkowska M (2019) Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar. ZooKeys 884 : 1–22, doi. Versioned wiki page: 2019-10-30, version 180386, https://species-id.net/w/index.php?title=Milnesium_matheusi&oldid=180386 , contributors (alphabetical order): Pensoft Publishers.

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BibTeX:

@article{Kaczmarek2019ZooKeys884,
author = {Kaczmarek, Łukasz AND Grobys, Daria AND Kulpa, Adam AND Bartylak, Tomasz AND Kmita, Hanna AND Kepel, Marta AND Kepel, Andrzej AND Roszkowska, Milena},
journal = {ZooKeys},
publisher = {Pensoft Publishers},
title = {Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar},
year = {2019},
volume = {884},
issue = {},
pages = {1--22},
doi = {10.3897/zookeys.884.29469},
url = {https://zookeys.pensoft.net/articles.php?id=29469},
note = {Versioned wiki page: 2019-10-30, version 180386, https://species-id.net/w/index.php?title=Milnesium_matheusi&oldid=180386 , contributors (alphabetical order): Pensoft Publishers.}

}

RIS/ Endnote:

TY - JOUR
T1 - Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar
A1 - Kaczmarek Ł
A1 - Grobys D
A1 - Kulpa A
A1 - Bartylak T
A1 - Kmita H
A1 - Kepel M
A1 - Kepel A
A1 - Roszkowska M
Y1 - 2019
JF - ZooKeys
JA -
VL - 884
IS -
UR - http://dx.doi.org/10.3897/zookeys.884.29469
SP - 1
EP - 22
PB - Pensoft Publishers
M1 - Versioned wiki page: 2019-10-30, version 180386, https://species-id.net/w/index.php?title=Milnesium_matheusi&oldid=180386 , contributors (alphabetical order): Pensoft Publishers.

M3 - doi:10.3897/zookeys.884.29469

Wikipedia/ Citizendium:

<ref name="Kaczmarek2019ZooKeys884">{{Citation
| author = Kaczmarek Ł, Grobys D, Kulpa A, Bartylak T, Kmita H, Kepel M, Kepel A, Roszkowska M
| title = Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar
| journal = ZooKeys
| year = 2019
| volume = 884
| issue =
| pages = 1--22
| pmid =
| publisher = Pensoft Publishers
| doi = 10.3897/zookeys.884.29469
| url = https://zookeys.pensoft.net/articles.php?id=29469
| pmc =
| accessdate = 2020-10-29

}} Versioned wiki page: 2019-10-30, version 180386, https://species-id.net/w/index.php?title=Milnesium_matheusi&oldid=180386 , contributors (alphabetical order): Pensoft Publishers.</ref>

See also the citation download page at the journal.


Taxonavigation

Ordo: Apochela
Familia: Milnesiidae
Genus: Milnesium

Name

Milnesium matheusi Kaczmarek & Grobys & Kulpa & Bartylak & Kmita & Kepel & Kepel & Roszkowska, 2019 sp. nov.Wikispecies linkZooBank linkPensoft Profile

Material examined

Holotype and 18 paratypes, all from sample No 139: Ivohibory forest, Madagascar, lichen sample from quartz rocks, coll. Marta Kepel and Andrzej Kepel.

Description

Adult females (Fig. 1, Table 4) with no modified claws I. Body light yellow before fixation and transparent afterwards, eyes present (in 89% of measured specimens). Dorsal cuticle sculptured with pseudopores, not arranged in bands, sparsely distributed and not forming a reticular design (Fig. 2). Six peribuccal papillae and six peribuccal lamellae present around the mouth opening. Two cephalic papillae positioned laterally. Peribuccal papillae slightly longer than lateral papillae.
The buccal apparatus of the Milnesium type (Figs 1, 3). The buccal tube wide and short (standard width, on average 46% of its length), and slightly funnel-shaped, wider anteriorly (posterior diameter on average 89% of the anterior diameter) (Table 4). The pharyngeal bulb elongated, pear-shaped and without placoids or septulum.

Table 4. Measurements and pt values of selected morphological structures of adult females of Milnesium matheusisp. nov. mounted in Hoyer’s medium (N – number of specimens/structures measured, RANGE refers to the smallest and the largest structure among all measured specimens; SD – standard deviation, pt – ratio of the length of a given structure to the length of the buccal tube expressed as a percentage).
Character N Range Mean SD Holotype
µm pt µm pt µm pt µm pt
Body length 6 630 766 691 45 766
Peribuccal papillae length 5 10.0 12.0 18.6 22.1 11.0 19.9 0.8 1.5 11.8 18.9
Lateral papillae length 7 9.4 10.7 16.5 19.7 10.0 18.1 0.4 1.2 10.3 16.5
Buccal tube
Length 9 51.3 62.5 56.6 3.8 62.5
Stylet support insertion point 9 34.5 42.3 66.1 69.4 38.4 67.8 2.4 1.3 41.5 66.4
Anterior width 9 25.2 35.9 47.6 57.9 28.9 51.0 3.2 3.1 31.4 50.2
Standard width 9 23.1 31.1 42.4 50.8 26.3 46.5 2.7 3.0 29.4 47.0
Posterior width 9 23.0 30.2 41.1 50.3 25.7 45.3 2.6 3.1 28.9 46.2
Standard width/length ratio 9 42% 51% 46% 3% 47%
Posterior/anterior width ratio 9 84% 94% 89% 4% 92%
Claw 1 lengths
External primary branch 9 17.2 21.8 30.2 35.2 18.9 33.3 1.5 1.6 21.8 34.9
External base + secondary branch 9 13.3 16.7 23.5 27.9 15.0 26.5 1.2 1.5 16.6 26.6
External spur 7 3.5 5.3 6.4 9.6 4.4 7.8 0.7 1.3  ?  ?
External branches length ratio 9 76% 82% 80% 2% 76%
Internal primary branch 9 16.0 21.1 30.2 34.5 18.3 32.3 1.6 1.6 21.1 33.8
Internal base + secondary branch 9 13.3 16.6 24.5 27.3 14.8 26.2 1.1 1.0 16.3 26.1
Internal spur 9 3.3 5.5 6.1 10.5 4.4 7.7 0.8 1.4 5.5 8.8
Internal branches length ratio 9 77% 88% 81% 4% 77%
Claw 2 lengths
External primary branch 8 17.4 21.2 32.9 36.5 19.5 34.9 1.4 1.4 21.2 33.9
External base + secondary branch 7 13.7 17.0 24.5 27.5 15.0 26.7 1.1 1.2 17.0 27.2
External spur 3 3.9 4.9 7.2 7.8 4.4 7.6 0.5 0.4 4.9 7.8
External branches length ratio 7 72% 81% 77% 3% 80%
Internal primary branch 8 16.8 20.5 31.1 35.7 18.7 33.3 1.3 1.5 20.2 32.3
Internal base + secondary branch 9 13.0 16.3 25.0 27.9 14.7 26.0 1.1 0.9 16.3 26.1
Internal spur 9 3.4 5.8 6.1 10.3 4.4 7.8 0.8 1.5 4.7 7.5
Internal branches length ratio 8 74% 81% 78% 3% 81%
Claw 3 lengths
External primary branch 5 19.7 21.0 32.3 38.3 20.5 35.7 0.6 2.5  ?  ?
External base + secondary branch 6 14.2 16.3 24.5 28.4 15.4 27.1 0.7 1.5  ?  ?
External spur 5 3.5 5.2 6.4 9.3 4.4 7.7 0.7 1.1  ?  ?
External branches length ratio 5 72% 82% 75% 4%  ?
Internal primary branch 5 18.9 20.4 31.3 36.5 19.7 34.3 0.6 2.0  ?  ?
Internal base + secondary branch 6 13.7 16.0 23.7 28.2 14.9 26.2 0.8 1.9  ?  ?
Internal spur 5 3.8 5.6 7.0 9.7 4.8 8.3 0.7 1.1  ?  ?
Internal branches length ratio 5 70% 79% 75% 4%  ?
Claw 4 lengths
Anterior primary branch 7 19.6 23.0 35.1 39.8 20.9 37.2 1.3 1.5 23.0 36.8
Anterior base + secondary branch 7 14.6 17.2 26.3 29.4 15.8 28.2 0.9 1.1 17.2 27.5
Anterior spur 6 4.1 6.3 7.5 11.5 5.4 9.7 0.9 1.6 6.0 9.6
Anterior branches length ratio 7 71% 80% 76% 4% 75%
Posterior primary branch 7 20.5 24.0 38.1 41.3 21.8 38.9 1.1 1.1 24.0 38.4
Posterior base + secondary branch 7 15.2 17.7 26.9 29.6 16.1 28.6 0.8 1.0 17.7 28.3
Posterior spur 7 4.4 5.8 7.6 10.3 5.2 9.3 0.6 1.1 5.5 8.8
Posterior branches length ratio 7 70% 76% 74% 2% 74%
Claws of the Milnesium type, slender (Figs 4, 5). Primary branches on all legs with small, but distinct accessory points detaching from the branch at its greatest curvature (Fig. 5, arrowhead). Secondary branches with rounded basal thickenings (Figs 4, 5). All secondary branches on all legs with three points (claw configuration: [3-3]–[3-3]). Single, long transverse, cuticular bars present under claws I–III (Fig. 4, arrow). Adult males (Table 5) with modified claws I. Similar to females but clearly smaller, with secondary branches of claws I modified into strong hooks and with a different proportion of peribuccal and lateral papillae length (peribuccal papillae clearly shorter than lateral), eyes present only in 33% of measured specimens.

Eggs oval, smooth and deposited in the exuvium as in all other known Milnesium species.

Table 5. Measurements and pt values of selected morphological structures of adult males (with modified claws I) of Milnesium matheusi sp. nov. mounted in Hoyer’s medium (N – number of specimens/structures measured, RANGE refers to the smallest and the largest structure among all measured specimens; SD – standard deviation, pt – ratio of the length of a given structure to the length of the buccal tube expressed as a percentage).
Character N Range Mean SD
µm pt µm pt µm pt
Body length 2 409 428 419 13
Peribuccal papillae length 3 3.0 3.9 8.9 11.3 3.5 10.2 0.5 1.2
Lateral papillae length 3 5.6 6.0 16.2 17.8 5.9 17.1 0.2 0.8
Buccal tube
Length 3 33.8 34.5 34.2 0.4
Stylet support insertion point 2 21.2 22.3 62.7 64.6 21.8 63.7 0.8 1.4
Anterior width 3 9.4 11.2 27.8 32.6 10.5 30.8 1.0 2.6
Standard width 3 9.1 9.8 26.9 28.5 9.5 27.8 0.4 0.8
Posterior width 3 9.4 10.2 27.8 29.6 9.8 28.7 0.4 0.9
Standard width/length ratio 3 27% 28% 28% 1%
Posterior/anterior width ratio 3 88% 100% 94% 6%
Claw 1 lengths
External primary branch 2 15.8 16.3 45.9 48.2 16.1 47.1 0.4 1.6
External base + secondary branch 3 14.1 15.0 41.7 43.5 14.6 42.7 0.5 0.9
External spur 2 3.2 3.4 9.3 9.9 3.3 9.6 0.1 0.4
External branches length ratio 2 87% 94% 90% 5%
Internal primary branch 3 14.9 15.7 43.2 46.4 15.4 45.1 0.5 1.7
Internal base + secondary branch 3 14.0 14.5 40.6 42.9 14.2 41.4 0.3 1.3
Internal spur 3 3.0 3.7 8.9 10.7 3.4 9.9 0.4 0.9
Internal branches length ratio 3 89% 94% 92% 2%
Claw 2 lengths
External primary branch 2 16.9 17.9 49.0 53.0 17.4 51.0 0.7 2.8
External base + secondary branch 1 13.2 13.2 39.1 39.1 13.2 39.1  ?  ?
External spur 1 3.5 3.5 10.4 10.4 3.5 10.4  ?  ?
External branches length ratio 1 74% 74% 74%  ?
Internal primary branch 3 16.4 16.9 47.5 50.0 16.7 48.8 0.3 1.2
Internal base + secondary branch 2 12.7 12.8 37.2 37.6 12.8 37.4 0.1 0.3
Internal spur 2 3.5 5.0 10.4 14.5 4.3 12.4 1.1 3.0
Internal branches length ratio 2 75% 76% 76% 1%
Claw 3 lengths
External primary branch 3 16.2 17.4 47.1 51.5 16.8 49.0 0.6 2.3
External base + secondary branch 2 12.1 12.8 35.2 37.9 12.5 36.5 0.5 1.9
External spur 1 3.9 3.9 11.3 11.3 3.9 11.3  ?  ?
External branches length ratio 2 74% 75% 74% 1%
Internal primary branch 3 14.8 17.0 43.0 50.3 16.0 46.7 1.1 3.6
Internal base + secondary branch 2 12.7 13.0 37.6 37.8 12.9 37.7 0.2 0.2
Internal spur 2 2.9 4.0 8.4 11.8 3.5 10.1 0.8 2.4
Internal branches length ratio 2 75% 88% 81% 9%
Claw 4 lengths
Anterior primary branch 3 16.3 17.0 47.4 49.3 16.6 48.5 0.4 1.0
Anterior base + secondary branch 2 12.4 12.9 36.7 37.5 12.7 37.1 0.4 0.6
Anterior spur 1 3.8 3.8 11.0 11.0 3.8 11.0  ?  ?
Anterior branches length ratio 2 75% 79% 77% 3%
Posterior primary branch 3 17.7 18.8 51.3 54.7 18.3 53.4 0.6 1.8
Posterior base + secondary branch 3 12.7 13.7 37.1 39.8 13.1 38.2 0.6 1.5
Posterior spur 2 3.0 4.1 8.7 11.9 3.6 10.3 0.8 2.3
Posterior branches length ratio 3 69% 73% 72% 2%

DNA sequences

We obtained good quality sequences for the applied molecular markers: 28S rRNA sequence (GenBank: MN191503), 756 bp long; COI sequence (GenBank: MN187056), 628 bp long; ITS-2 sequence (GenBank: MN239906), 218 bp long.

Type locality

Madagascar, 22°37'07.7"S, 46°43'14.5"E, ca. 1187 m asl, Fianarantsoa Province, Ivohibory forest.

Etymology

The second author with great pleasure dedicates this species to her fiance – Mateusz Wojciechowski.

Type depositories

The holotype and 13 paratypes (slides: MAD139/14, MAD139/16, MAD139/18, MAD139/19, MAD139/34, MAD139/35, MAD139/42, MAD139/56, MAD139/72) are deposited at the Department of Animal Taxonomy and Ecology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, Poznań, Poland; five paratypes (slides: MAD139/12, MAD139/13, MAD139/15) are deposited at the Natural History Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark.

Morphological differential diagnosis

The new species with three points on the secondary branches of all claws (claw configuration [3-3]–[3-3]) and a rather wide buccal tube, in relation to its length, is most similar to: Mil. beatae Roszkowska, Ostrowska & Kaczmarek, 2015, Mil. bohleberi Bartels, Nelson, Kaczmarek & Michalczyk, 2014, Mil. eurystomum Maucci, 1991, Mil. shilohae Meyer, 2015 and Mil. tumanovi Pilato, Sabella & Lisi, 2016, but it differs from:
1. Milnesium beatae, only reported from Argentina and USA (Roszkowska et al. 2015[1]; Tibbs et al. 2016[2]) by: narrower buccal tube (25.2–35.9 [47.6–57.9] and 23.1–31.1 [42.4–50.8] anterior and standard width, respectively, in the new species vs. 37.0–53.5 [70.3–78.9] and 32.0–42.5 [58.1–65.6] anterior and standard width respectively in Mil. beatae), smaller standard width/length ratio of the buccal tube (42%–51% in new species vs. 58%–66% in Mil. beatae) and larger posterior/anterior width ratio (84%–94% in new species vs. 69%–76% in Mil. beatae).
2. Milnesium bohleberi, only recorded from North Carolina and Tennessee (USA) (Bartels et al. 2014[3]) by: presence of pseudopores on dorsal cuticle, shorter peribuccal papillae (10.0–12.0 [18.6–22.1] in new species vs. 15.5–20.3 [27.2–32.3] in Mil. bohleberi), smaller pt values of anterior, standard and posterior widths of the buccal tube (47.6–57.9, 42.4–50.8, 41.1–50.3, respectively, in new species vs. 63.4–74.7, 54.5–64.0, 52.4–62.0, respectively, in Mil. bohleberi), smaller standard width/length ratio of the buccal tube (42%–51% in new species vs. 54%–64% in Mil. bohleberi) and slightly shorter claws (see Table 4 below and Bartels et al. (2014[3]: Table 1) for the exact differences in claw dimensions).
3. Milnesium eurystomum reported from a few localities in Argentina, Chile, Greenland, Mongolia and USA (see review by Kaczmarek et al. 2016[4]) by: shorter buccal tube (51.3–62.5 in new species vs. 70.8–77.5 in Mil. eurystomum), stylet supports inserted in a more posterior position (pt = 66.1–69.4 in new species vs. ca. pt = 60.0–60.3 in Mil. eurystomum), narrower buccal tube (25.2–35.9 [47.6–57.9], 23.1–31.1 [42.4–50.8] and 23.0–30.2 [41.1–50.3] anterior, standard and posterior width, respectively, in new species vs. 53.7–55.9 [72.1–75.8], 45.9–47.9 [61.8–64.8] and 33.9–41.0 [43.7–57.9] anterior, standard and posterior width, respectively, in Mil. eurystomum), smaller standard width/length ratio of the buccal tube (42%–51% in new species vs. 62%–65% in Mil. eurystomum) and larger posterior/anterior width ratio (84%–94% in new species vs. 61%–76% in Mil. eurystomum).
4. Milnesium shilohae, only reported from the type locality in Hawaii (USA) (Meyer 2015[5]) by: presence of pseudopores on dorsal cuticle, presence of similar in length spurs on internal and external claws (internal and posterior spurs larger than external and anterior spurs in Mil. shilohae), slightly longer lateral papillae (9.4–10.7 in new species vs. 5.0–9.0 in Mil. shilohae), slightly longer buccal tube (51.3–62.5 in new species vs. 38.4–50.3 in Mil. shilohae), stylet supports inserted in a more anterior position (pt = 66.1–69.4 in new species vs. pt = 75.5–77.5 in Mil. shilohae) and larger spurs on some external and anterior claws (see Table 4 below and Table 3 in Meyer (2015)[5] for the exact differences in claw dimensions).
5. Milnesium tumanovi, only recorded from the type locality in the Crimea (Ukraine) (Pilato et al. 2016[6]) by: presence of pseudopores on dorsal cuticle, funnel-shaped buccal tube (cylindrical in Mil. tumanovi) and stylet supports inserted in a more posterior position (pt = 66.1–69.4 in new species vs. ca. pt = 52–54 in Mil. tumanovi).

Genotypic differential diagnosis

The ranges of uncorrected genetic p-distances between Mil. matheusi sp. nov. and species of the genus Milnesium, for which molecular marker sequences are available from GenBank (see Table 6 for details), are as follows:

Table 6. Sequences of 28S rRNA, COI and ITS-2 of Milnesium taxa available in GenBank and used in differential diagnosis.
DNA marker Taxon Accession number Source
28S rRNA Milnesium sp. JX888585.1 Adams et. al. unpublished
JX888586.1 Adams et. al. unpublished
JX888587.1 Adams et. al. unpublished
Milnesium tardigradum JX888541.1 Adams et. al. unpublished
JX888540.1 Adams et. al. unpublished
KC138808.1 Zawierucha unpublished
KC138809.1 Zawierucha unpublished
Milnesium sp. AY210826.1 Mallatt et. al. 2004[7]
Milnesium tardigradum FJ435780.1 Guil and Giribet 2012[8]
FJ435779.1 Guil and Giribet 2012[8]
Milnesium berladnicorum KT951661.1 Morek et. al. 2016[9]
Milnesium variefidum KT951665.1 Morek et. al. 2016[9]
COI Milnesium sp. KX306950.1 Fox et al. unpublished
Milnesium tardigradum EU244603.1 Schill unpublished
EU244604 Schill unpublished
FJ435810.1 Guil and Giribet 2012[8]
Milnesium t. tardigradum JN664950.1 Michalczyk et al. 2012[10]
Milnesium cf. tardigradum JX683824.1 Vicente et al. 2013[11]
JX683823.1 Vicente et al. 2013[11]
JX683822.1 Vicente et al. 2013[11]
Milnesium sp. KJ857002.1 Velasco-Castrillón et al. 2015[12]
KJ857001.1 Velasco-Castrillón et al. 2015[12]
Milnesium cf. alpigenum KU513422.1 Kosztyła et al. 2016[13]
Milnesium variefidum KT951663.1 Morek et al. 2016[9]
Milnesium berladnicorum KT951659.1 Morek et al. 2016[9]
Milnesium sp. EF632553.1 Sands et. al unpublished
Milnesium cf. granulatum MH751517.1 Jackson and Meyer 2019[14]
Milnesium lagniappe MH751518.1 Jackson and Meyer 2019[14]
Milnesium tardigradum MG923558.1 Morek et al. 2019[15]
MG923559.1 Morek et al. 2019[15]
MG923560.1 Morek et al. 2019[15]
MG923561.1 Morek et al. 2019[15]
MG923562.1 Morek et al. 2019[15]
MG923563.1 Morek et al. 2019[15]
MG923564.1 Morek et al. 2019[15]
MG923565.1 Morek et al. 2019[15]
Milnesium dornensis MG923566.1 Morek et al. 2019[15]
ITS-2 Milnesium alpigenum MH000382.1 Morek et al. unpublished
Milnesium sp. MH000386.1 Morek et al. unpublished
MH000387.1 Morek et al. unpublished
Milnesium tardigradum HM150648.2 Wełnicz et. al. 2010[16]
GQ403682.1 Schill et al. 2010[17]
GQ403681.1 Schill et al. 2010[17]
Milnesium t. tardigradum JF951049 Michalczyk et al. 2012[10]
Milnesium variefidum KT951667.1 Morek et. al. 2016[9]
KT951666.1 Morek et. al. 2016[9]
Milnesium berladnicorum KT951662.1 Morek et. al. 2016[9]
Milnesium cf. granulatum MK681875.1 Jackson and Meyer 2019[14]
MK681876.1 Jackson and Meyer 2019[14]
MK681877.1 Jackson and Meyer 2019[14]
MK681878.1 Jackson and Meyer 2019[14]
MK681879.1 Jackson and Meyer 2019[14]
MK681880.1 Jackson and Meyer 2019[14]
MK681881.1 Jackson and Meyer 2019[14]
MK681882.1 Jackson and Meyer 2019[14]
MK681883.1 Jackson and Meyer 2019[14]
MK681884.1 Jackson and Meyer 2019[14]
MK681885.1 Jackson and Meyer 2019[14]
MK681886.1 Jackson and Meyer 2019[14]
1. 28S rRNA: 4.5–6.7% (5.4% on average), with the most similar being Milnesium sp. from North America (JX888585.1, JX888586.1, JX888587.1) (unpublished) and the least similar being Mil. wrightae sp. nov. (MN191504.1) (present studies);

2. COI: 20.1–38.8% (23.3% on average), with the most similar being Mil. variefidum Morek, Gąsiorek, Stec, Blagden & Michalczyk, 2016 from UK (KT951663.1) (Morek et al. 2016[9]) and the least similar being Mil. t. tardigradum from Spain (FJ435810.1) (Guil and Giribet 2012[8]);
3. ITS-2: 17.8–31.1% (23.7% on average), with the most similar being Mil. t. tardigradum from Germany (JF951049.1) (Michalczyk et al. 2012[10]) and the least similar being Mil. cf. granulatum from USA (MK681879.1) (Jackson and Meyer 2019[14]).

Original Description

  • Kaczmarek, Ł; Grobys, D; Kulpa, A; Bartylak, T; Kmita, H; Kepel, M; Kepel, A; Roszkowska, M; 2019: Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar ZooKeys, 884: 1-22. doi

Images

Other References

  1. Roszkowska M, Ostrowska M, Kaczmarek Ł (2015) The genus Milnesium Doyère, 1840 (Tardigrada) in South America with descriptions of two new species from Argentina and discussion of the feeding behaviour in the family Milnesiidae.Zoological Studies54(1): 1–12. https://doi.org/10.1186/s40555-014-0082-7
  2. Tibbs L, Emanuels A, Miller W (2016) Tardigrades of the canopy: Argentine species Milnesium beatae Roszkowska, Ostrowska and Kaczmarek, 2015 (Eutardigrada, Milnesidae) discovered in the trees of Kansas, USA.Transactions of the Kansas Academy of Science119(2): 173–178. https://doi.org/10.1660/062.119.0207
  3. 3.0 3.1 Bartels P, Nelson D, Kaczmarek Ł, Michalczyk Ł (2014) The genus Milnesium (Tardigrada: Eutardigrada: Milnesiidae) in the Great Smoky Mountains National Park (North Carolina and Tennessee, USA), with the description of Milnesium bohleberi sp. nov.Zootaxa3826(2): 356–368. https://doi.org/10.11646/zootaxa.3826.2.5
  4. Kaczmarek Ł, Michalczyk Ł, Sandra J (2016) Annotated zoogeography of non-marine Tardigrada. Part III: North America and Greenland.Zootaxa4203(1): 1–249. https://doi.org/10.11646/zootaxa.4203.1.1
  5. 5.0 5.1 Meyer H (2015) Water bears (Phylum Tardigrada) of Oceania, with the description of a new species of Milnesium.New Zealand Journal of Zoology42(3): 173–186. https://doi.org/10.1080/03014223.2015.1062402
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