Next Article in Journal
Reptile Biodiversity and Vulnerability in Bolivia’s Beni Department: Informing Conservation Priorities in a Neglected Frontier
Previous Article in Journal
What Factors Determine the Natural Fruit Set of Cephalanthera longifolia and Cephalanthera rubra?
Previous Article in Special Issue
Investigation of Potential Effects of Ibuprofen on the Storage Cells and Anhydrobiosis Capacity of the Tardigrade Paramacrobiotus experimentalis
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Diversity
Volume 16
Issue 6
10.3390/d16060334
diversity-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles thumb_up
...
Endorse textsms
...
Comment
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Tardigrades of North America: Additions to Montana’s Biodiversity Including a New Species, Platicrista loloensis nov. sp. (Parachela, Hypsibioidea, Itaquasconinae) †

by
Chelsea N. Scheirer
1,
William R. Miller
2,* and
Jeffrey D. Miller
3
1
Department of Biological Sciences, University of Alabama, Tuscaloosa, AL 35401, USA
2
Department of Biology, Baker University, Baldwin City, KS 66006, USA
3
Biological Research and Education Consultants, Missoula, MT 59801, USA
*
Author to whom correspondence should be addressed.
†
urn:lsid:zoobank.org:pub:B8BECA01-877A-4ADD-B836-589CBA15CE1E.
Diversity 2024, 16(6), 334; https://doi.org/10.3390/d16060334
Submission received: 18 January 2024 / Revised: 5 April 2024 / Accepted: 9 April 2024 / Published: 6 June 2024
(This article belongs to the Special Issue Investigating the Biodiversity of the Tardigrada)
Browse Figures
Review Reports Versions Notes

Abstract

:
A total of 205 tardigrades representing two orders, five families, nine genera and ten species were extracted from a moss sample (104 tardigrades) and a lichen sample (101 tardigrades) collected near Missoula, Montana, in 2016. Three of the species are new to Montana and one is new to science, Platicrista loloensis nov. sp., which is distinguished by its smooth cuticle, the presence of internal cuticular bars at the base of the claws of legs II and III and a median cuticular bar between the claws of leg IV.

1. Introduction

There are only five reports of the phylum Tardigrada in the state of Montana. Leetham et al. [1,2] twice reported the same three genera (Macrobiotus, Diphascon and Hexapodibius) but did not identify the species. Beasley [3] recorded the first species, when he described Hypsibius macrocalcaratus [3] from New Mexico and added Montana as another place he had found the animal. Miller [4] listed 20 species from Montana west of the continental divide. Miller and J.D. Miller [5] extended the known range of eight species into the Bob Marshall Wilderness Complex. Two were additions to Montana’s biodiversity list and one was new to science, Placticrista brunsoni [5]. This report adds three species to the states’ biodiversity list, including one that is new to science.

2. Study Area and Methods

A moss habitat sample was collected from a site on the north side of Lolo Creek (a tributary of the Bitterroot River) near Fort Fizzle south of Missoula, Montana (Figure 1). The moss sample was taken from the center of a plot that measured approximately 30 × 30 cm. The moss was pulled from the gravelly soil substrate and clinging debris was removed using scissors. The moss was put into a plastic bag that contained 510 mL of distilled water and taken to the lab, where it was stored in a refrigerator at approximately 1–2 °C until being processed to extract tardigrades.
A lichen habitat sample was scraped from a lichen-covered fence rail on the north side of an irrigation ditch located on the east side of Rattlesnake Creek (a tributary of the Clark Fork River) north of Missoula (Figure 1). The lichen sample was dry when collected and stored in a paper bag at room temperature (21–22 °C) until processing.
In the laboratory, approximately one-third of each sample was soaked in distilled water. After 24 h, three 2.5 mL aliquots of debris and fluid were extracted from each sample, examined in a black-bottomed, rectangular petri dish using a 25× dissecting microscope illuminated by LED lights to search for tardigrades [6,7]. Tardigrades were removed with an Irwin loop [8] and placed into a drop of polyvinyl alcohol (PVA) on a glass slide, which was covered with a round glass coverslip. The coverslip was sealed with fingernail polish after five days of drying [6,7].
Each specimen was examined using an Olympus BX-60 microscope equipped with Nomarski Differential Interference Contrast objectives (DIC). Images were 5.0 MP taken with a Miotic M-500 digital camera and saved in PNG format. The images were cropped to fit the page and only adjusted for brightness and contrast for printability [9].
Measurements were taken µm using AmScope version 64x (2003–2022) software Data were stored and formatted in Excel spreadsheets. A data table was used to characterize species in which N = number of specimens, range = (min–max), mean = average of N specimens, pt = character length/buccal tube length, b = slope of regression of data, and a* = Y intercept per Bartels et al. [10]. We accept the use of the mean of the pt value [11] of the set of measurements used to describe a species and published as the statement of diagnosis for that variable. This, while slightly imperfect, pt value allows variable characteristics to be compared within a taxon and among taxa.
Nomenclature was based on the “Actual Checklist of Tardigrades” by Guidetti and Bertolani [12], Degma and Guidetti [13], Degma, et al. [14], and Degma and Guidetti [15]. Identification was based on the work of Ramazzotti and Maucci [16], Pilato and Binda [17], and current literature, including Gąsiorek et al. [18]. Buccal tube measurements followed the work of Pilato [11]. Abbreviations followed the work of Perry et al. [19,20]. North American distribution and biodiversity were based on the work of McInnes [21], Meyer [22], Kaczmarek et al. [23], and Miller and Perry [24].

3. Results

Both samples were positive for tardigrades. The three aliquots from the moss sample yielded 104 individuals and the three aliquots from the lichen sample yielded 101. The moss sample yielded eight species from three families and five specimens in simplex and five specimens that could not be identified because they were distorted when positioned on the slide. Of those eight species, three were new to Montana and one was new to science (Table 1). The lichen sample yielded four species from four families and three specimens that could not be identified because they were distorted when positioned on the slide (Table 1).
The species of tardigrades recovered were not uniformly distributed among the two samples representing two different types of habitats (Table 1). All three members of the family Hypsibiidae were found only in the moss. Three of five Macrobiotidae species were also found only in the moss. Mesobiotus harmsworthi sensu lato, known for its intercontinental distribution, was recovered from both the moss and lichen samples. Milnesium and Ramazzottius were found only in the lichen sample. No pattern of association among the species and their habitats can be determined with only two samples.

4. Taxonomic Treatment of New Species

When a specimen is identified, it is assigned to a genus by confirming the specimen has the diagnostic characteristics for that classification. Then, it is assigned to a species based on the variation in generic characteristics or the addition of a unique specific character(s). Thus, when one or more of a specimen’s characters do not correspond with the characters of previously described species and cannot be reasonably explained by variation, then a new species should be created [25].
PhylumTardigrada Doyère, 1840
ClassEutardigrada Richters, 1926
OrderParachela Schuster, Nelson, Grigarick & Christenberry, 1980
SuperfamilyHypsibioidea Pilato, 1969
FamilyHypsibiiidea Pilato, 1969
SubfamilyItaquasconinae Rudescu, 1964
GenusPlaticrista Pilato, 1987
SpeciesPlaticrista loloensis nov. sp.
 Figure 2 and Figure 3, Table 2 and Table 3
The number in parenthesis in the description is the measurement of the holotype.
Diagnosis: Large, gray Platicrista with a smooth cuticle, without eyes before mounting. Smooth, ridged buccal tube, flexible pharyngeal tube, two long, thin macroplacoids in a row, with the second twice as long as the first and no microplacoid or septulum. Claws of Hypsibius type, internal cuticular bars at the base of claws on legs II and III, median cuticular bar between claws on legs IV and ragged pseudolunules present on the base of claws IV (Figure 2).
Description: Large (615 µm), gray Platicrista, without eyes before mounting. Sub-terminal mouth. Cuticle smooth but may show wrinkles. Pointed, spoon-like furcae away from buccal tube. Buccopharyngeal apparatus with smooth (73.05 µm), ridged buccal tube (36.46 µm) to stylet support attachment point (36.46 µm) and ringed, flexible pharyngeal tube (36.59 µm) posterior to the stylet support attachment point into the oval pharynx and terminating with small distal thickenings (Figure 2A). Buccal tube with an external narrow width (9.24 µm) (Figure 2A) measured at the stylet attachment point (Table 3). Elongated, oval pharynx without apophysis (72.71 µm). Pharyngeal cuticular bars pointing forward in the pharynx anterior to first placoid (Figure 2A, a). Two thin macroplacoids in placoid stack (42.46 µm) with the first measuring 13.29 µm and the second measuring 26.25 µm. Microplacoid and septulum absent. Claws of legs I–III are the Hypsibius type (1212), with the largest being the external claws with a stout primary branch, light transmission areas, smooth bases with pseudolunula, and topped with small accessory points (29.39 µm) (Figure 2B–D). Internal claws are smaller but robust with light transmission areas (Figure 2B–F), smooth bases with pseudolunula, and strong accessory points (17.63 µm). No cuticular bars at the base of claws on leg I, with an internal cuticular bar next to the base of internal claws of legs II and III (Figure 2C,D). Claws of leg IV are the Hypsibius type (1212), with the posterior claws being the largest (39.59 µm) with a stout primary branch, light transmission areas, topped with small accessory points, and smooth indented bases with thin, flat, serrated pseudolunula. Anterior claws are smaller but more robust (26.95 µm) with stout primary claws, light transmission areas, topped with large accessory points and smooth indented bases with thin, flat, serrated pseudolunula. Median cuticular bar between the anterior and posterior claws of legs IV (Figure 2E,F).
Holotype: Collected from a sample of moss on 28 December 2016 by Chelsea Scheirer on the south side of Lolo Creek near Fort Fizzle, Montana, of the Bitterroot River (46.74410, −114.17442) (Figure 1). Deposited at the University of California Davis, Bohart Museum of Entomology, Davis California (Number S-0025810).
Paratypes: Five additional specimens were extracted from the same moss sample. Deposited at the University of California Davis, Bohart Museum of Entomology, Davis California (Numbers S-0025811, S-0025812, S-0025813, S-0025814, S-0025815).
Etymology: The new species is named for the creek that runs beside the collection site. The area has cultural significance for American Indian tribes, specifically the Séliš, QÍispé, Ksanka and Nimiipuu peoples. The creek and canyon are located within the aboriginal territories of these tribes from time immemorial as a pathway between present day Montana and Idaho. In addition, the Lewis and Clark Expedition used the canyon while travelling west in 1805 and east in 1806.
Differentiation: The new species is similar to the nine other species of the genus Platicrista but is different from Platicrista cheleusis [26], Platicrista brunsoni [5] and Platicrista carpathica [18] in Gąsiorek et al. [18] by its smooth vs. tubercated cuticle. Platicrista loloensis nov. sp. has internal cuticular bars under the claws that are not found on Platicrista nivea [18] in Gąsiorek et al. [18], Platicrista angustata [27] or Platicrista horribilis [28]. Platicrista ramsayi [29] presents internal cuticular bars on legs I, II and III while Platicrista loloensis nov. sp. presents internal cuticular bars only on legs II and III. Platicrista borneensis [18] in Gąsiorek et al. [18] presents median cuticular bars on legs I-IV, while Platicrista loloensis nov. sp. has a median cuticular bar only between the claws on leg IV (Table 3). The new species has serrated lunules that are similar to Pla. angustata and Pla. horribilis species with a smooth cuticle. It differs from both by the presence of median cuticular bars in legs IV.
Discussion: The genus Platicrista [30] was erected within the subfamily Itaquasconinae Bartoš in Rudescu, [31] to separate the species Pla. angustata, originally described by Murray [27] with a flexible bucco-pharyngeal tube with flat ridges for the insertion of stylet muscles and pointed, spoon-like furcae. Pilato [30] included Diphascon itaquasconoide [32] and Diphascon affine [33] in the new genus. Over the next 16 years, two additional species were described, including Pla. cheleusis by Kathman [26] from Vancouver Island and Pla. horribilis by Kaczmarek and Michalczyk [28] from Mongolia. Three years later, Marley [29] added Pla. ramsayi from Ecuador and followed Dastych’s [34] recommendation to declare Pla. affine to be a nomen dubium. Miller and J.D. Miller [5] found Pla. brunsoni in the southern part of the Bob Marshall Wilderness Complex of Montana. Recently, Gąsiorek et al. [18] added three additional species: Pla. carpathica from Poland, Pla. nivea from Taiwan, and Pla. borneensis from Borneo.
The genus Meplitumen [35] was separated from Platicrista by Lisi et al. [35] based on the presence of annulations on the base of the buccal tube. Meplitumen aluna [35] was described as the type species for the genus. Massa et al. [36] transferred Pla. itaquasconoide [32] from Platicrista to Meplitumen. Tumanov and Tsvetkova [37] identified faint annulations on the buccal tube of Pla. angustata specimens and by using 18S and 28S rRNA fragments, they moved all the Meplitumen species back to the genus Platicrista because there were no other significant morphological differences, declaring Meplitumen to be a junior synonym to Platicrista. Gąsiorek et al. [18] found no differences and formally synonymized Pla. itaquasconoide to Pla. angustata.
Gąsiorek et al. [18] advanced a re-description of the genus Platicrista, (page 27), which contains several diagnostic statements that are not inclusive of some of the species within the genus. The sentence “Cuticle smooth, without pores or undulations…” does not include the three species (Pla. brunsoni, Pla. carpathica, and Pla. cheleusis) with notable polyform tubercles. The statement “…smooth, ridged buccal tube…” does not include the buccal tube annulations of Pla. aluna from Meplitumen. The sentence “The macroplacoid margins are serrated” should be a species character because some are smooth. The sentence “Pseudolunulae present and well developed” does not apply to all species. Finally, the sentence “Cuticular bars absent.” does not apply to four species in the genus with internal and median cuticular bars (Pla. brunsoni, Pla. ramsayi, Pla. cheleusis, Pla. loloensis nov. sp.).
Thus, we advance the following amended re-description of the genus Platicrista to include all ten species currently assigned to the genus (Table 3).
Amended description of Platicrista: Adult body medium-sized to very large, up to 1000 µm, elongated and slightly narrowed anteriorly. Body white to brownish. Head blunt and mouth sub-terminal. Eyes absent. Cuticle without pores, but may be smooth or display irregular wrinkles, fine granulation or shallow polygonal tubercles. Legs very short and plump; legs IV barely distinguishable from the posterior body. Buccopharyngeal apparatus with a wide ridged buccal tube that may be smooth or faintly annulated; flexible, annulated pharyngeal tube; apodeme at the border of the buccal and pharyngeal tube is absent. Stylet supports present and inserted at the caudal position of the buccal tube. Furcae cup shaped with pointed ends. Oral cavity armature absent. Caudal apophyses are wide and show flat ridges. Pharyngeal apophyses absent but the terminal end pharyngeal tube may be slightly enlarged. Pharynx oval and broad, with two long, thin macroplacoids, with the second being about twice as long as the first (1 < 2). Claws of the Hypsibius type (1212), large and robust, primary branches of exterior and posterior claws with barely divergent accessory points, slender at their tips. Primary branches of internal and anterior claws are particularly massive with large accessory spines. Pseudolunulae may be present or absent and cuticular bars may be present at the base of the claws in internal or median positions.
The genus seems to have a largely Holarctic distribution with no records of occurrence from Central America [38], or Australasia [39], and only one record from northern Africa [40]. To date, there are three records from South America [41] including Pla. ramsayi from Ecuador [29] near the equator; thus, only the two records from Argentina are significantly below the equator. All three of the new species reported by Gąsiorek et al. [18] are from the equator or above (Borneo, Taiwan, and Poland).
The genus remains infrequently recorded across North America (Canada, Greenland, and USA); there are only 57 records of occurrence for five of the ten species over a period of 120 years. Three of the five species are endemic to North America. Forty-one of those records are Pla. angustata from places in Alaska, Kansas, New Brunswick and Greenland, suggesting a continental distribution range (Figure 3), but many records are older and may represent newly described or additional undescribed species. They need verification. Platicrista cheleusis has been recorded 11 times from Alaska to Alberta to Vancouver to Colorado, suggesting a northwestern North American range (Figure 3). Platicrista brunsoni has two records, including one from western Montana and another from northern Wyoming, suggesting a limited regional range (Figure 3). The remaining record is a single occurrence in Tennessee of Pla. horribili, a species known originally from Mongolia. Platicrista loloensis nov. sp. has only one record so far (Figure 3).
Platicrista specimens seldom occur in great numbers. Most species have been described based on very few specimens. This new species is no exception, with just six specimens recovered so far and because it has been found at only one location, the presence of a regional population is not yet suggested. The description of five new species in the last four years and the clarification of specific characteristics suggest that the previously reported specimens of Platicrista species should be reexamined to have their characteristics confirmed.

5. Comments

There are about 1500 described species of tardigrades worldwide [15]. However, only 320 of these species have been found in North America [22,23,24]. The records of occurrence of these species are clumped into a few areas because systematic, wide-scale sampling of tardigrade habitats has not been conducted. Generally, collecting occurs through the efforts of a primary researcher and their associates as they address specific research questions or during collecting expeditions (Alaska [42], California [43], Kansas [44], Illinois [45], Tennessee [46,47], New Jersey [48], New Brunswick [49,50,51], and Greenland [52]).
Recent zoogeographic reviews of the records of occurrence in the literature have provided some understanding of the distribution and diversity of these species [23,24,38,39,40,41]. Although there are a few thousand records of occurrence of a tardigrade species being collected at a specific location in North America, there remain great expanses of the continent with no records. There are even fewer data concerning the environments, habitats, and/or relationships to temperature, moisture, or vegetation type. While geopolitical records (states) are important to humans, they mean little to the animals. It would be more useful to consider EPA ecoregions [53] that relate to the conditions of the earth. For example, Missoula is in the 17s Bitterroot-Frenchtown Valley Ecoregion of the level IV EPA.
The diversity of tardigrades in our samples of moss and lichen are typical of habitat samples from around the world (WRM, personal experience). The numbers of one or two species dominate the diversity within the sample, while most species are represented by far fewer specimens. This new species is not an exception. Because it has been found at only one location so far, there is just a single record of occurrence until additional specimens from additional locations are reported.
The addition of three species to the biodiversity list for the state of Montana represents a 10% increase and suggests there are many more species to be found in the geographic boundaries of the state [54].
The Montana biodiversity list for tardigrades has been updated and adjusted for the taxonomic structure and names to reflect the latest guidelines from the “Actual Checklist of Tardigrade Names Version 42” [15] (Table 4).

Author Contributions

C.N.S. conceived the project, C.N.S. and J.D.M. collected, extracted, and mounted the specimens. W.R.M. identified the specimens and prepared the descriptions. C.N.S., W.R.M. and J.D.M. prepared the manuscript. All authors have read and agreed to the published version of the manuscript.

Funding

Microscope used at the Department of Biology of Baker University were acquired by NSF Grant DEB 0640847. The APC was funded by the journal Diversity. Support for C.N.S., including laboratory facilities, was provided by Big Sky-High School, Missoula, Montana, while C.N.S. was a student.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Data is contained within the article.

Acknowledgments

The project was conducted under US Forest Service permit SP0106.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Leetham, J.W.; McNary, T.J.; Dodd, J.L.; Laurenroth, W.K. Response of soil nematodes, rotifers, and tardigrades to three levels of season long sulfur dioxide exposures. Water Air Soil Pollut. 1980, 17, 343–356. [Google Scholar] [CrossRef]
  2. Leetham, J.W.; McNary, T.J.; Dodd, J.L.; Laurenroth, W.K. Response of field populations of Tardigrada to various levels of chronic low-level sulphur dioxide exposure. In Soil Biology as Related to Land Use Practices; Dindal, D.L., Ed.; Office of Pesticides and Toxic Substances, Environmental Protection Agency: Washington, DC, USA, 1982; pp. 382–390. [Google Scholar]
  3. Beasley, C.W. Altitudinal distribution of Tardigrada of New Mexico with the description of a new species. Am. Midl. Nat. 1988, 120, 436–440. [Google Scholar] [CrossRef]
  4. Miller, W.R. Tardigrades of North America: Western Montana. Intermt. J. Sci. 2006, 12, 45–57. [Google Scholar]
  5. Miller, W.R.; Miller, J.D. Tardigrades of North America: Platicrista brunsoni nov. sp. (Parachela, Hypsibiidae, Itaquasconinae) from the Bob Marshall Wilderness Area of Montana. Northwest Sci. 2021, 95, 98–105. [Google Scholar] [CrossRef]
  6. Miller, W.R. Tardigrades: Bears of the Moss. Kans. Sch. Nat. 1997, 43, 1–16. [Google Scholar]
  7. Miller, W.R. Tardigrades: Stalking the Wild Moss Bear. Kans. Sch. Nat. 2023, 67, 1–16. [Google Scholar]
  8. Schram, M.D.; Davidson, P.G. A History and Method of constructing homemade loops. Trans. Kans. Acad. Sci. 2012, 115, 35–40. [Google Scholar] [CrossRef]
  9. McInnes, S.J. Is It Real? Zool. Anz. 2001, 240, 467–468. [Google Scholar] [CrossRef]
  10. Bartels, P.J.; Nelson, D.R.; Ryan, P.; Exline, R.P. Allometry and the removal of body size effects in the morphometric analysis of tardigrades. J. Zoolgical Syst. Evol. Res. 2011, 49, 17–25. [Google Scholar] [CrossRef]
  11. Pilato, G. Analisi di nuovi caratteri nello studio degli Eutardigradi. Animalia 1981, 8, 51–57. [Google Scholar]
  12. Guidetti, R.; Bertolani, R. Tardigrade taxonomy: An updated check list of the taxa and a list of characters for their identification. Zootaxa 2005, 845, 1–46. [Google Scholar] [CrossRef]
  13. Degma, P.; Guidetti, R. Notes to the current checklist of Tardigrada. Zootaxa 2007, 1579, 41–53. [Google Scholar] [CrossRef]
  14. Degma, P.; Bertolani, R.; Guidetti, R. Actual Checklist of Tardigrada Species. Ver. 29: 15 December 2015. p. 37. Available online: http://www.tardigrada.modena.unimo.it/miscellanea/Actual%20checklist%20of%20Tardigrada.pdf (accessed on 1 October 2016).
  15. Degma, P.; Bertolani, R.; Guidetti, R. Actual Checklist of Tardigrada Species; University of Modena and Reggio Emilia: Modena, Italy, 2023. [Google Scholar] [CrossRef]
  16. Ramazzotti, G.; Maucci, W. Il Phylum Tardigrada. III Ed. Riveduta Eggiornata-Mem. Dell’Istituto Ital. Idrobiol. 1983, 41, 1–1011. [Google Scholar]
  17. Pilato, G.; Binda, M. Definition of families, subfamilies, genera and subgenera of the Eutardigrada and keys to their identification. Zootaxa 2010, 2404, 1–54. [Google Scholar] [CrossRef]
  18. Gąsiorek, P.; Blagden, B.; Morek, W. Michalczyk, Ł. What is a ‘strong’ synapomorphy? Redescriptions of Murray’s type species and descriptions of new taxa challenge the systematics of Hypsibiidae (Eutardigrada: Parachela). Zool. J. Linn. Soc. 2023, zlad151. [Google Scholar] [CrossRef]
  19. Perry, E.S.; Miller, W.R.; Kaczmarek, L. Recommended abbreviations for the names of genera of the phylum Tardigrada. Zootaxa 2019, 4608, 145–154. [Google Scholar] [CrossRef] [PubMed]
  20. Perry, E.S.; Miller, W.R.; Kaczmarek, Ł. Additional recommended abbreviations for the names of genera of the phylum Tardigrada. Zootaxa 2021, 4981, 398–400. [Google Scholar] [CrossRef] [PubMed]
  21. McInnes, S.J. Zoogeographic Distribution of Terrestrial/freshwater Tardigrades from current literature. J. Nat. Hist. 1994, 116, 185–205. [Google Scholar] [CrossRef]
  22. Meyer, H.A. Terrestrial and freshwater Tardigrada of the Americas. Zootaxa 2013, 3747, 1–71. [Google Scholar] [CrossRef] [PubMed]
  23. Kaczmarek, Ł.; Michalczyk, Ł.; McInnes, S.J. Annotated zoogeography of non-marine Tardigrada. Part III: North America and Greenland. Zootaxa 2016, 4203, 1–249. [Google Scholar]
  24. Miller, W.R.; Perry, E. Adjustments to the Annotated zoogeography of non-marine Tardigrada. Part III: North America and Greenland by Kaczmarek, Michalczyk & McInnes (Zootaxa 4203). Zootaxa 2019, 4345, 99–114. [Google Scholar]
  25. Winston, J.E. Describing Species: Practical Taxonomic Procedure for Biologists; Columbia University Press: New York, NY, USA, 1999; p. 518. [Google Scholar]
  26. Kathman, D.R. Eutardigrada from Vancouver Island, British Columbia, Canada, including a description of Platicrista cheleusis n.sp. Can. J. Zsoology 1990, 68, 1880–1895. [Google Scholar] [CrossRef]
  27. Murray, J. Tardigrades of the Scottish lochs. Trans. R. Soc. Edinb. 1905, 61, 100–108. [Google Scholar]
  28. Kaczmarek, Ł.; Michalczyk, Ł. Platicrista horribilis, a new species of Tardigrada (Eutardigrada: Hypsibiidae) from Mongolia. Zootaxa 2003, 132, 1–8. [Google Scholar] [CrossRef]
  29. Marley, N.J. A new species of Tardigrada, Platicrista ramsayi sp. nov. from the páramo of Volcán Chiles, Ecuador. Zootaxa 2006, 1166, 35–48. [Google Scholar] [CrossRef]
  30. Pilato, G. Revision of the genus Diphascon Plate, 1889, with remarks on the subfamily Itaquasconinae (Eutardigrada, Hypsibiidae). In Biology of Tardigrades; Bertolani, R., Ed.; Selected Symposia an Monographs, U.Z.I.; Mucchi: Modena, Italy, 1987; pp. 337–357. [Google Scholar]
  31. Rudescu, L. Tardigrada. Fauna Rep. Pop. Rom. Bucur. 1964, 4, 1–398. [Google Scholar]
  32. Durante Pasa, M.V.; Maucci, W. Descrizione di tre nuove specie di Tardigradi della Scandinavia. Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano 1975, 116, 244–250. [Google Scholar]
  33. Mihelcic, F. Beitrag zur Systematik de Tardigraden. Arch. Zool. Ital. 1951, 36, 57–103. [Google Scholar]
  34. Dastych, H. Redescription of the cryoconital tardigrade Hypsibius klebelsbergi Miheli, 1959, with notes on the microslide collection of the late Dr. F. Mihelcic (Tardigrada). Sonderdr. Veröffentlichungen Mus. Ferdinandeunm 1993, 73, 5–12. [Google Scholar]
  35. Lisi, O.; Daza, A.; Londoño, R.; Quiroga, S.; Pilato, G. Meplitumen aluna gen. nov., sp. nov. an interesting eutardigrade (Hypsibiidae, Itaquasconinae) from the Sierra Nevada de Santa Marta, Colombia. ZooKeys 2019, 865, 1–20. [Google Scholar] [CrossRef] [PubMed]
  36. Massa, E.; Guidetti, R.; Cesari, M.; Rebecchi, L.; Jönsson, I. Tardigrades of Kristianstads Vattenrike Biosphere Reserve with description of four new species from Sweden. NaturePortfolio Sci. Rep. 2021, 11, 4861. [Google Scholar] [CrossRef] [PubMed]
  37. Tumanov, D.V.; Tsvetkova, A.Y. Some have drops and some do not, but can we rely on that? Re-ivestigation of Diphascon tenue (Tardigrada: Eutardigrada) with discussion of phylogeny and taxonomy of the superfamiy Hypsibioidea. Zoosytematica Ross. 2023, 32, 50–74. [Google Scholar] [CrossRef]
  38. Kaczmarek, Ł.; Michalczyk, Ł.; McInnes, S.J. Annotated zoogeography of non-marine Tardigrada. Part I: Central America. Zootaxa 2014, 3763, 1–62. [Google Scholar] [CrossRef]
  39. Michalczyk, Ł.; Kaczmarek, Ł.; McInnes, S.J. Annotated zoogeography of non-marine Tardigrada. Part V: Australasia. Zootaxa 2022, 5107, 1–119. [Google Scholar] [CrossRef] [PubMed]
  40. McInnes, S.J.; Michalczyk, Ł.; Kaczmarek, Ł. Annotated zoogeography of non-marine Tardigrada. Part IV: Africa. Zootaxa 2021, 4284, 1–74. [Google Scholar] [CrossRef]
  41. Kaczmarek, Ł.; Michalczyk, Ł.; McInnes, S.J. Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa 2015, 3923, 1–107. [Google Scholar]
  42. Johansson, C.; Miller, W.R.; Linder, E.T.; Adams, B.Y.; Boreliz-Alvarado, E. Tardigrades of Alaska: Distribution patterns, diversity and species richness. Polar Res. 2013, 32, 18793. [Google Scholar] [CrossRef]
  43. Schuster, R.O.; Grigarick, A.A. Tardigrada from Western North America: With emphasis on the Fauna of California. Univ. Calif. Publ. Zool. 1965, 76, 1–67. [Google Scholar]
  44. Lehmann, R.E.; Shively, S.D.; Miller, W.R. Tardigrades of North America: An historical collection from Kansas and Missouri. Trans. Kans. Acad. Sci. 2007, 110, 169–178. [Google Scholar] [CrossRef]
  45. Pugila, C.R. Some tardigrades from Illinois. Trans. Am. Microsc. Soc. 1964, 83, 300–311. [Google Scholar] [CrossRef]
  46. Bartels, P.J.; Nelson, D.R. A large-scale, multihabitat inventory of the Phylum Tardigrada in the Great Smokey Mountains National Park, USA: A preliminary report. Hydrobiological 2006, 558, 111–118. [Google Scholar] [CrossRef]
  47. Bartels, P.J.; Nelson, D.R. An evaluation of species richness estimators for tardigrades of the Great Smoky Mountains National Park, Tennessee and North Carolina, USA. Proceedings of the Tenth International Symposium on Tardigrada. J. Limnol. 2007, 66 (Suppl. S1), 104–110. [Google Scholar] [CrossRef]
  48. Shaw, M.W.; Miller, W.R. Tardigrades of North America: A preliminary survey of New Jersey, U.S.A. Newsl. N. Y. Microsc. Soc. 2013, 7, 36–56. [Google Scholar]
  49. Argue, C.W. Some terrestrial tardigrades from New Brunswick, Canada. Can. J. Zool. 1971, 49, 401–415. [Google Scholar] [CrossRef]
  50. Argue, C.W. Tardigrades of New Brunswick, Canada 2. Can. J. Zool. 1972, 50, 87–94. [Google Scholar] [CrossRef]
  51. Argue, C.W. Tardigrades of from New Brunswick, Canada 3. Can. J. Zool. 1974, 52, 919–922. [Google Scholar] [CrossRef]
  52. Petersen, B. The Tardigrade Fauna of Greenland. Meddelelser Grønland 1951, 150, 1–1194. [Google Scholar]
  53. Omernik, J.M. and Griffith. G.M. Ecoregions of the conterminous United States: Evolution of a hierarchical spatial framework. Environ. Manag. 2014, 54, 1249–1266. [Google Scholar] [CrossRef]
  54. Bartels, P.J.; Apodacal, J.J.; Mora, C.; Nelson, D.R. A global biodiversity estimates of a poorly known taxon: Phylum Tardigrada. Zool. J. Linn. Soc. 2016, 178, 730–736. [Google Scholar] [CrossRef]
Diversity 16 00334 g001
Figure 1. Location of moss and lichen collection sites in western Montana. Based on Natural Earth and Google Earth Pro maps and the images.
Figure 1. Location of moss and lichen collection sites in western Montana. Based on Natural Earth and Google Earth Pro maps and the images.
Diversity 16 00334 g001
Diversity 16 00334 g002
Figure 2. DIC images Platicrista loloensis nov. sp. (A) Buccal apparatus, a = pharyngeal cuticular bar, b = buccal tube, p = pharyngeal tube, i = first placoid, ii = second placoid (400×), (B) claws of leg I; (C) claws of leg II, c = internal cuticular bar; (D) claws of leg III, c = internal cuticular bar; (E) claws of leg IV, l = pseudolunule; (F) claws of leg IV, l = pseudolunule, m = median cuticular bar. (B–F) = 1000×, Scale bar = 10 µm.
Figure 2. DIC images Platicrista loloensis nov. sp. (A) Buccal apparatus, a = pharyngeal cuticular bar, b = buccal tube, p = pharyngeal tube, i = first placoid, ii = second placoid (400×), (B) claws of leg I; (C) claws of leg II, c = internal cuticular bar; (D) claws of leg III, c = internal cuticular bar; (E) claws of leg IV, l = pseudolunule; (F) claws of leg IV, l = pseudolunule, m = median cuticular bar. (B–F) = 1000×, Scale bar = 10 µm.
Diversity 16 00334 g002
Diversity 16 00334 g003
Figure 3. North American records of the genus Platicista. Red dots = Pla. angustata, light blue dots = Pla. cheleusis, green dots = Plal. brunsoni, black dot = Pla. horribilis, yellow dot = Pla. loloensis nov. sp. Locations taken from the work of Kaczmarek et al., 2016 [23].
Figure 3. North American records of the genus Platicista. Red dots = Pla. angustata, light blue dots = Pla. cheleusis, green dots = Plal. brunsoni, black dot = Pla. horribilis, yellow dot = Pla. loloensis nov. sp. Locations taken from the work of Kaczmarek et al., 2016 [23].
Diversity 16 00334 g003
Table 1. Tardigrade species reported from samples collected in 2016 at Lolo Creek and Rattlesnake Creek sites near. Missoula Montana. * = New to Montana. Taxonomy follows Degma and Guidetti [15].
Table 1. Tardigrade species reported from samples collected in 2016 at Lolo Creek and Rattlesnake Creek sites near. Missoula Montana. * = New to Montana. Taxonomy follows Degma and Guidetti [15].
ClassMossLichen
 Order  
  Superfamily  
   Family  
    Subfamily  
     Genus species  
Eutardigrada Richters, 1926  
 Apochela Schuster, Nelson, Grigarick and Christenberry, 1980   
   Milnesiidae Ramazzotti, 1962  
     Milnesium sp. 024
 Parachela Schuster, Nelson, Grigarick & Christenberry, 1980   
  Hypsibioidea Plato. 1969  
   Hypsibiidae Pilato, 1969  
    Diphasconinae Dastych, 1992  
     Diphascon pingue (Marcus, 1936) sensu lato *10
    Itaquasconinae Bartoš in Rudescu, 1964  
     Adropion scoticum (Murray, 1905)20
     Platicrista loloensis nov. sp. *60
   Ramazzottiidae Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
     Ramazzottius oberhaeuseri (Doyère, 1840)064
  Isohypsibioidea Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
   Isohypsibioidae Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
     Isohypsibius sp.23
  Macrobiotoidea Thulin, 1928  
   Macrobiotidae Thulin, 1928  
     Macrobiotus hufelandi C.A.S. Schultze, 1834 sensu lato490
     Mesobiotus harmsworthi (Murray, 1907) sensu lato154
     Paramacrobiotus richtersi (Murray, 1911) sensu lato40
     Paramacrobiotus tonollii (Ramazzotti, 1956) *150
Identification undetermined53
Total 104101
Table 2. Morphometric data for Platicrista loloensis nov. sp.
Table 2. Morphometric data for Platicrista loloensis nov. sp.
MinMaxMeanSlope (b)Y Intercept (a*)
Body Length (BL)6530.00750.00621.33  
   pt = BL/BTL (%)61341%1869%1666%0.621768%
Bucco-Pharyngeal Apparatus (BPA)670.7284.8477.860.4175.51
   pt = BPA/BTL (%)6194%237%210%0.85206%
Buccal Tube Length (BTL)629.8941.2637.380.3837.38
Buccal Tube Width (BTW)68.5910.249.240.259.34
   pt = BTW/BTL (%)622%29%25%−0.1326%
Stylet Support Attachment (SSA)629.8941.2637.380.3837.98
   pt = SSA/BTL (%)6100%100%100%0.00100%
Pharyngeal Tube Length (PTL)636.5946.3240.480.1438.63
   pt = PTL/BTL (%)694%137%110%0.04100%
Pharynx Length (PhL) 667.3280.6274.040.3774.08
   pt = PhL/BTL (%)6170%246%200%0.28193%
Placoid Row Length (PRL)641.8752.3546.420.5647.60
   pt = PRL/BTL (%)6106%146%125%−0.20125%
Placoid 1 Length (P1L)610.9015.6013.450.7513.51
   pt = P1L/BTL (%)631%39%36%0.3737%
Placoid 2 Length (P2L)526.2536.0731.180.5130.77
   pt = P2L/BTL (%)572%87%80%0.4280%
Exterior Claw I Length (ECI)529.3933.1831.360.2331.85
   pt = ECI/BTL (%)576%83%81%0.1483%
Interior Claw I Length (ICI)616.3724.1919.980.9020.71
   pt = ICI/BTL (%)641%59%51%0.8154%
Anterior Claw IV Length (ACL-IV) 621.6526.9525.100.4127.16
   pt = ACL-IV/BTL (%)655%79%68%0.0079%
Posterior Claw IV Length (PCL-IV) 632.2843.9739.590.1740.01
   pt = PCL-IV/BTL (%)694.43%114.15%106.18%−0.23109.26%
Table 3. Differential comparison of species of Platicristia.
Table 3. Differential comparison of species of Platicristia.
Platicrista brunsoniPlaticrista cheleusisPlaticrista carpathicaPlaticrista barneensisPlaticrista ramsayiPlaticrista loloensis nov. sp.Platicrista niveaPlaticrista angustataPlaticrista horribilisPlaticrista aluna
Specimens used in description3141073632835
Apophyses: Flat YesYesYesYesYesYesYesYesYesYes
Furcae: Spoon-like YesYesYesYesYesYesYesYesYesYes
Buccal tube: Ridged YesYesYesYesYesYesYesYesYesYes
Pharyngeal tube: Flexible and striated YesYesYesYesYesYesYesYesYesYes
Phaaryngeal apophysis: Absent YesYesYesYesYesYesYesYesYesYes
Claws: Hypsibius type (2121)YesYesYesYesYesYesYesYesYesYes
Cuticle: Smooth NoNoNoYesYesYesNoYesYesYes
Cuticle: Low tuberclesYesYesYesNoNoNoNoNoNoNo
Cuticle granulatedNoNoYesNoNoNoYesNoNoNo
Pharyngeal tube shorter than buccal tubeYesNoNoYesYesYes/NoYesYesYes/NoYes
pt SSA 100%100%100%82%97%100%100%99%100%100%
PT/BT ratio92%95%107%144%86%104%112%118%108%89%
Cuticular bars present YesYesNoYesYesYesNoNoNoNo
 Internal bars on legs I NoYesNoNoYesNoNoNoNoNo
 Median bars on legs INoYesNoNoNoNoNoNoNoNo
 Internal bars on legs II–III YesYesNoNoYesYesNoNoNoNo
 Median bars on legs I–IIINoYesNoYesNoNoNoNoNoNo
 Posterior bars on legs IV NoNoNoYesNoNoNoNoNoNo
 Median bars on legs IVNoYesNoYesNoYesNoNoNoNo
Base of claws on leg IV: Smooth NoYesNoYesYesYesYesNoNoYes
Base of claws on leg IV: Indented YesNoYesNoNoYesNoNoYesNo
Pseudolunulae on legs IV YesNoYesNoNoYesYesYesYesYes
Pseudolunulae on legs IV Serrated = Ser, smooth = SmoSerNoSmoNoNoSerSmoSmoSerSmo
Table 4. Tardigrade Biodiversity in Montana, U.S.A. in 2024. Taxonomy follows Degma and Guidetti [15].
Table 4. Tardigrade Biodiversity in Montana, U.S.A. in 2024. Taxonomy follows Degma and Guidetti [15].
Class     
 OrderLeetchamBeasleyMillerMillerThis
  Superfamilyet al.  andReport
   Family   Miller 
    Subfamily1980, 19821988200620212024
     Genus species     
Heterotardigrada Marcus, 1927     
 Echiniscoidea Richters, 1926     
   Ehiniscidae Thulin, 1928     
     Acanthechiniscus victor (Ehrenberg, 1853)  X  
     Echiniscus arctomys Ehrenberg, 1853   X  
     Echiniscus quadrispinosus Richters, 1902   X  
     Echiniscus tristosus Cuénot, 1932   X  
     Multipseudechiniscus raneyi (Grigarick, Mihelčič and Schuster, 1964) XX 
Eutardigrada Richters, 1926     
 Apochela Schuster, Nelson, Grigarick and Christenberry, 1980    
   Milnesiidae Ramazzotti, 1962     
     Milnesium sp.     X
     Milnesium tardigradum Doyère, 1840 sensu lato X  
 Parachela Schuster, Nelson, Grigarick and Christenberry, 1980  
  Hypsibioidea Plato. 1969     
   Hypsibiidae Pilato, 1969     
    Diphasconinae Dastych, 1992     
     Diphascon alpinum Murray, 1906  X  
     Diphascon pingue (Marcus, 1936) sensu lato    X
     Diphascon sp.X    
    Hypsibiinae Plato, 1969     
     Hypsibius convergens (Urbanowicz, 1925)  X  
     Hypsibius macrocalcaratus Beasley, 1988 X   
    Itaquasconinae Bartoš, 1962      
     Adropion scoticum (Murray, 1905)  X X
     Guidettion arduifrons (Thulin, 1928)  X  
     Mesocrista spitzbergensis (Richters, 1903  X  
     Platicrista angustata (Murray, 1905)  X  
     Placticrista brunsoni Miller and J.D. Miller, 2022  X 
     Platicrista loloensis nov. sp.    X
    Pilatobiinae Bertolani, Guidetti, Marchioro, Altiero, Rebecchi and Gąsiorek, 2014
     Pilatobius nodulosum (Ramazzotti, 1957)   X 
     Pilatobius oculatum (Murray, 1906)  X  
   Ramazzottiidae Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
     Ramazzottius oberhaeuseri (Doyère, 1840)  X X
  Isohypsibioidea Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
   Hexapodibiidae Cesari, Vechhi, Plamer, Bertolani, Rebecchi and Geodetti, 2016
     Hexapodibus sp.X    
   Isohypsibioidae Sands, McInnes, Marley, Goodall-Copestake, Convey and Linse, 2008
     Isohypsibius prosostomus Thulin, 1928  X  
     Isohypsibius sp.   XX
  Macrobiotoidea Thulin, 1928     
   Macrobiotidae Thulin, 1928     
     Macrobiotus hufelandi C.A.S. Schultze, 1834 sensu latoXXX
     Macrobiotus islandicus Richtersi, 1904  X  
     Macrobiotus sp. X    
     Mesobiotus harmsworthi (Murray, 1907) sensu lato XXX
     Paramacrobiotus areolatus (Murray, 1907) sinsu lato X  
     Paramacrobiotus richtersi (Murray, 1911) sensu lato X X
     Paramacrobiotus tonollii (Ramazzotti, 1956)    X
     Total Species = 273120610
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Scheirer, C.N.; Miller, W.R.; Miller, J.D. Tardigrades of North America: Additions to Montana’s Biodiversity Including a New Species, Platicrista loloensis nov. sp. (Parachela, Hypsibioidea, Itaquasconinae). Diversity 2024, 16, 334. https://doi.org/10.3390/d16060334

AMA Style

Scheirer CN, Miller WR, Miller JD. Tardigrades of North America: Additions to Montana’s Biodiversity Including a New Species, Platicrista loloensis nov. sp. (Parachela, Hypsibioidea, Itaquasconinae). Diversity. 2024; 16(6):334. https://doi.org/10.3390/d16060334

Chicago/Turabian Style

Scheirer, Chelsea N., William R. Miller, and Jeffrey D. Miller. 2024. "Tardigrades of North America: Additions to Montana’s Biodiversity Including a New Species, Platicrista loloensis nov. sp. (Parachela, Hypsibioidea, Itaquasconinae)" Diversity 16, no. 6: 334. https://doi.org/10.3390/d16060334

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop