Barbaria weglarskae, GASIOREK, WILAMOWSKI,, 2022

Gąsiorek, Piotr, Wilamowski, Andrzej, Vončina, Katarzyna & Michalczyk, Łukasz, 2022, Neotropical jewels in the moss: biodiversity, distribution and evolution of the genus Barbaria (Heterotardigrada: Echiniscidae), Zoological Journal of the Linnean Society 195, pp. 1037-1066 : 1048-1054

publication ID	https://doi.org/ 10.1093/zoolinnean/zlab087
publication LSID	lsid:zoobank.org:pub:87E03803-7D69-4393-93CD-24A39D607972
DOI	https://doi.org/10.5281/zenodo.6993209
persistent identifier	https://treatment.plazi.org/id/586D3B6C-0239-6145-EC64-CBFDFDA3FA70
treatment provided by	Plazi
scientific name	Barbaria weglarskae
status	SP. NOV.

Treatment

BARBARIA WEGLARSKAE GĄSIOREK, WILAMOWSKI, View in CoL

VONČINA & MICHALCZYK, SP. NOV.

( FIGS 4 View Figure 4 , 7L View Figure 7 , 8L View Figure 8 , 9L View Figure 9 , 10L View Figure 10 , 11M, N; TABLE View Figure 11 5)

Zoobank registration: u r n: l s i d: z o o b a n k. org:act: F083907B-741F-48C0-A127-2FC28482C9AA

Morphometric data: w w w. t a r d i g r a d a. n e t / register/0110.htm

Ty p e m a t e r i a l: Holotype (adult female, slide AR.059.04) and GoogleMaps two paratypes (adult females, slides AR.059.01, 05). GoogleMaps

Type locality: 48°25′42″S, 71°44′48″W, 803 m asl: Argentina, Patagonia , Santa Cruz Province, Río Chico Department, vicinity of La Florida; lichen from on in the Andean Patagonian forest (see also Table 1 View Table 1 ) GoogleMaps .

Etymology: A patronym honouring Professor Barbara Węglarska, 20.02.1922 – 02.10.2020, whose death left a void in the community of tardigradologists. A noun in the genitive case.

Description: Adult females (i.e. from the third instar onwards, measurements in Table 5). Orange body with large, red eyes; the entire pigment and eyes dissolve quickly after mounting in Hoyer’s medium. Body massive ( Fig. 4A View Figure 4 ). Cylindrical, Echiniscus - type cephalic papillae (secondary clavae) and (primary) clavae; cirri embedded in bulbous cirrophores. Cirrus A is long (> 50% of the body length) and with evident, conical cirrophore ( Fig. 4A, B View Figure 4 ).

Dorsal plate sculpturing of the bigranulata - type, composed of pillars present in all plate portions (pseudogranulation) and pores and pseudopores present in different elements of armour ( Figs 4A, B View Figure 4 , 7L View Figure 7 , 8L View Figure 8 , 9L View Figure 9 , 10L View Figure 10 ). Pseudopores can be present or absent exclusively in the anterior portion of the median plate 2 and paired segmental plates and the entirety of median plate 3 ( Fig. 4A, B View Figure 4 ). Minute pores of equal size are regularly distributed in the remaining plate portions ( Figs 4A, B View Figure 4 , 7L View Figure 7 , 8L View Figure 8 , 9L View Figure 9 , 10L View Figure 10 ). The cephalic plate large, with a pronounced chalice-shaped anterior incision and lateral sutures demarcating roughly triangular lateralmost portions of the plate ( Fig. 4A, B View Figure 4 ). Thin cervical plate with developed pillars and pseudopores. Lateral sutures in the scapular plate demarcate lateralmost, rectangular portions with identical sculpturing as on the rest of the plate ( Fig. 4A, B View Figure 4 ). Median plates 1 and 3 unipartite (the latter strongly reduced and partially covered by the caudal plate), median plate 2 bipartite ( Figs 4A, B View Figure 4 , 8L View Figure 8 ). Paired segmental plates with broad, transverse, unsculptured bands ( Figs 4A, B View Figure 4 , 9L View Figure 9 ). The caudal plate with short, poorly sclerotized incisions clearly joined by a transversal suture ( Figs 4A, B View Figure 4 , 10L View Figure 10 ). Ventral cuticle with minute endocuticular pillars distributed evenly throughout the entire venter; a pair of small, subcephalic plates present ( Fig. 4C–D View Figure 4 ). Sexpartite gonopore ( Fig. 4E View Figure 4 ) placed anteriorly to a trilobed anus between legs IV.

Pedal plates I–IV with evident pillars and pseudopores ( Figs 4A View Figure 4 , 11M, N View Figure 11 ). Evident pulvini on outer sides of all legs. Dentate collar IV with numerous irregular short teeth ( Fig. 11N View Figure 11 ). A tiny spine I and a tubby papilla IV ( Fig. 11N View Figure 11 ). Claws slightly heteronych, because primary spurs on internal claws IV are positioned higher than those on claws I–III ( Fig. 11M, N View Figure 11 ). The shape and angle at which spurs diverge from branches are almost identical on all limbs. Cuticular bars below claw bases on the inner side of legs present. Buccal apparatus with a rigid tube and round pharynx containing placoids. Lacking stylet supports.

Adult males, juveniles, larvae or eggs not found.

Differential diagnosis: There are only two other species of Barbaria with a cirrus A /body length ratio> 50% [the titles and values in the last five rows of table 1 in Michalczyk & Kaczmarek (2007) are mismatched in the case of sexes treated separately, but the ratio statistics for all measured specimens of B. ganczareki stands valid: min = 15%, max = 23%, mean = 19%]: B. jenningsi and B. ranzii , but B. weglarskae can be distinguished from:

• Barbaria jenningsi , by the type of perforation in the dorsal plates [dominant pores in B. weglarskae ( Fig. 7L View Figure 7 ) vs. pseudopores in B. jenningsi ( Figs 7E, F View Figure 7 , 8F View Figure 8 , 9F View Figure 9 , 10F View Figure 10 )] and claw isomorphy [anisonych/ slightly heteronych in B. weglarskae ( Fig. 11M, N View Figure 11 ) vs. strongly heteronych in B. jenningsi ( Fig. 11E, F View Figure 11 )].

• Barbaria ranzii , by the pedal plate sculpturing [with evident pillars in B. weglarskae ( Fig. 11M, N View Figure 11 ) vs. without pillars in B. ranzii ( Fig. 11R View Figure 11 )], the shape of papilla IV [tubby in B. weglarskae ( Fig. 11N View Figure 11 ) vs. elongated in B. ranzii ] and by the presence of secondary spurs directed upwards on external claws IV [absent in B. weglarskae ( Fig. 11N View Figure 11 ) vs. present in B. ranzii ( Fig. 11R View Figure 11 )].

MOLECULAR PHYLOGENY

Phylogeny based on the five concatenated markers brought fully resolved relationships between all eight analysed species of Barbaria , which form clades characterized by low intraspecific and large interspecific genetic variability ( Fig. 5 View Figure 5 ). The topology is as follows: B. madonnae is a sisterspecies to all other sequenced Barbaria spp. , which are clustered in two clades: the first comprising ( B. paucigranulata ( B. danieli + B. charrua )) and the second grouping (( B. weglarskae + B. jenningsi ) + ( B. ollantaytamboensis + B. bigranulata )).

INTRA- AND INTERSPECIFIC GENETIC VARIABILITY

Regarding COI sequences deposited in GenBank, the data are available only for B. bigranulata and B. jenningsi . COI p -distances between populations of B. bigranulata and the previously published data for a population from Chile ( HM193406 View Materials ; Jørgensen et al., 2011) ranged between 2.6 and 2.9% (alignment length = 585 bp). Analogous index for the pair B. weglarskae – B. jenningsi ( KP013596 View Materials ; Velasco-Castrillón et al., 2015) was 18.9% (alignment length = 472 bp).

More than one haplotype per marker has been found for all markers, but only in a few species. The intraspecific p -distances are as follows: 18S rRNA: 0.1% (in B. bigranulata and B. charrua ); 28S rRNA: 0.1–0.4% ( B. bigranulata , B. ollantaytamboensis ); ITS 1: 0.5 % ( B. bigranulata ), 0.1 – 1.0 % ( B. charrua ); ITS2: 0.2% ( B. bigranulata ), 0.2–0.4% ( B.ollantaytamboensis ); COI: 0.1–2.3% ( B. bigranulata ) and 0.7% ( B. charrua ).

Interspecific p -distances in the analysed dataset are as follows:

• 18S rRNA: 0.0–2.5% (1.2% on average), with the most similar being B. charrua ( MZ820796 View Materials ) and B. danieli ( MZ820800 View Materials ); and the least similar being B. madonnae ( MZ820803 View Materials ) and B. ollantaytamboensis ( MZ820804 View Materials ).

• 28S rRNA: 0.0–4.3% (2.4% on average), with the most similar being B. charrua ( MZ820814 View Materials ) and B. danieli ( MZ820818 View Materials ); and the least similar being B. madonnae ( MZ820821 View Materials ) and B. ollantaytamboensis ( MZ820823 View Materials ).

• ITS1: 0.3–10.3% (5.5% on average), with the most similar being B. charrua ( MZ820833 View Materials ) and B. danieli ( MZ820836 View Materials ); and the least similar being B. madonnae ( MZ820839 View Materials ) and B. bigranulata ( MZ820828 View Materials ).

• ITS2: 1.6–10.8% (8.1% on average), with the most similar being B. charrua ( MZ822380 View Materials ) and B. danieli ( MZ822384 View Materials ); and the least similar being B. danieli + B. madonnae ( MZ822384 View Materials , MZ822387 View Materials ) and B. ollantaytamboensis ( MZ822388 View Materials –91).

• COI: 9.1–20.4% (16.0% on average), with the most similar being B. charrua ( MZ820850 View Materials ) and B. danieli ( MZ820853 View Materials ); and the least similar being B. ollantaytamboensis ( MZ820855 View Materials ) and B. weglarskae ( MZ820860 View Materials ).

MORPHOLOGICAL EVOLUTION

Mapping morphological traits on to the phylogeny suggests that the ancestor of Barbaria was most probably covered with uniform dorsal sculpturing comprising both pillars and pores ( Fig. 6A View Figure 6 ) and welldelimited pedal plates on legs I–III with densely packed pillars and pseudopores ( Fig. 6B View Figure 6 ). Its papillae on legs IV were elongated ( Fig. 6C View Figure 6 ), meaning that they were much longer than wide. Internal claws were exhibiting heteronychy or lacking primary spurs ( Fig. 6D View Figure 6 ). In other words, among the extant species, B. bigranulata is morphologically overall the most similar to the last common ancestor of the analysed species.