taxonID	type	description	language	source
586D3B6C0237615CEFF9CE9AFCC0F8E5.taxon	description	(FIGS 1 – 3, 7 I, 8 I, 9 I, 10 I, 11 K, L; TABLES 3, 4) Zoobank registration: u r n: l s i d: z o o b a n k. org: act: A 11 E 6 BDF- 7 E 2 A- 4 F 0 A-BD 6 A- 9 BC 87 FCE 147 A Morphometric data: w w w. t a r d i g r a d a. n e t / register / 0109. htm	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C0237615CEFF9CE9AFCC0F8E5.taxon	materials_examined	Type material: Holotype (adult female, slide AR. 303.02) and 12 paratypes (eight adult females on slides AR. 302.02, AR. 303.01, 3, 5 – 6 and four juveniles on slides AR. 303.03 – 4). Found together with B. ollantaytamboensis. Type locality: 24 ° 47 ′ 14 ″ S, 65 ° 43 ′ 30 ″ W, 2150 m asl: Argentina, Salta Province, Rosario de Lerma Department, vicinity of Río Rosario; lichen on rock in a shrubland (see also Table 1).	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C0237615CEFF9CE9AFCC0F8E5.taxon	etymology	Etymology: From Latin paucus, few, and granulatus, grained, alluding to the scarcity of epicuticular granulation on the dorsal plates. An adjective in nominative singular.	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C0237615CEFF9CE9AFCC0F8E5.taxon	description	Description: Adult females (i. e. from the third instar onwards, measurements in Table 3). Dark orange body with large red eyes; the pigment and eyes dissolve quickly after mounting in Hoyer’s medium. Body massive with stumpy limbs (Figs 1 A, B, 2). Cylindrical, Echiniscus - type cephalic papillae (secondary clavae) and (primary) clavae; cirri embedded in bulbous cirrophores (Fig. 3 D). Cirrus A is short (<20 % of the body length) and with evident, but small cirrophore (Fig. 1 A, B). Dorsal plate sculpturing of the bigranulata - type, comprising minute, poorly developed pillars (pseudogranulation) and pores (Figs 1 C, 2, 3 A – C, 7 I, 8 I, 9 I, 10 I). Pillars are densely packed and additionally interconnected by thin striae in the scapular (Fig. 7 I) and caudal (terminal) plates (Fig. 10 I). Pores are slightly larger in the scapular plate, median plate 1, posterior portion of median plate 2 and centroposterior portions of paired segmental plates compared with posterolateral portions of paired segmental plates and the caudal plate (Figs 1, 2); pores of similar diameter are rarely present in the entire dorsum. Pores are always absent in the anterior portion of median plate 2 (Fig. 8 I), paired segmental plates (Fig. 9 I) and the entirety of median plate 3; and all these areas are covered with epicuticular, multangular granules. Lateralmost portions of the scapular and paired segmental plates can be poreless or with single minute pores (Figs 1 A, B, 2). The cephalic plate is large, with a pronounced chalice-shaped anterior incision and with only pillars in the posterior portion of the plate. A broad and strongly sclerotized cervical plate is divided into sculptured anterior portion and smooth posterior portion bordering with the scapular plate (Figs 1 A, 2 A). Lateral sutures in the scapular plate demarcate lateralmost, trapezoidal portions (Figs 1 A, B, 2 B). Median plates 1 and 3 are unipartite (the latter with strongly developed granules), median plate 2 is bipartite (Fig. 8 I). Paired segmental plates are without transverse unsculptured bands, as epicuticular granules of anterior portion transition gradually into pillars of the posterior portions (Fig. 9 I). The caudal plate with short, poorly sclerotized incisions and no signs of faceting (Figs 1 A, 2, 10 I). Ventral cuticle with minute endocuticular pillars (Fig. 11 K) distributed evenly throughout the entire venter, lacking plates, beside of rarely developed subcephalic plates (Fig. 3 D). Sexpartite gonopore placed anteriorly to a trilobed anus between legs IV. Pedal plates without pores, their sculpturing consists of poorly developed endocuticular pillars formed as belts in the central portions of the legs (Figs 1 A, B, 11 K, L). Thick pulvini on outer side of all legs (Figs 1 A, B, 2 B). Dentate collar IV has numerous irregular short teeth (Fig. 11 L). A small, elongated spine I and a tubby papilla IV (Figs 1 B, 2 B, 11 K, L). Claws slightly heteronych with claws IV (Fig. 3 F) higher and more robust than claws I – III (Fig. 3 E). Internal claws IV have needle-like spurs more divergent from branches compared to spurs I – III (Fig. 11 K, L). Cuticular bars below claw bases on the inner side of legs present. Buccal apparatus with a rigid tube and round pharynx containing placoids. Flexible stylet supports present. Juveniles (i. e. the second instar, measurements in Table 4). Clear morphometric gap between juveniles and adult females. Qualitatively alike adult females, excluding the lack of gonopore. Adult males, larvae or eggs not found.	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C0237615CEFF9CE9AFCC0F8E5.taxon	diagnosis	Differential diagnosis: The new species B. paucigranulata is distinguished from its congeners (alphabetically): • Barbaria bigranulata, by the distribution of pores in plates [absent in the anterior portion of median plate 2 (Fig. 8 I), paired segmental plates (Fig. 9 I) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m 2 (Fig. 8 A), paired segmental plates (Fig. 9 A) and the entirety of m 3 in B. bigranulata], the shape of papilla IV [tubby in B. paucigranulata (Fig. 11 L) vs. elongated in B. bigranulata (Fig. 11 B)] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. robust, hook-shaped and divergent from the claw branch in B. bigranulata (Fig. 11 A, B)]. • Barbaria charrua, by the shape of papilla IV [tubby in B. paucigranulata (Fig. 11 L) vs. elongated in B. charrua (Fig. 11 D)] and by the primary spurs [present in B. paucigranulata (Fig. 11 K, L) vs. typically absent in B. charrua (Fig. 11 C, D)]. • Barbaria danieli, by the shape of papilla IV [tubby in B. paucigranulata (Fig. 11 L) vs. elongated in B. danieli (Fig. 12)] and by the primary spurs [present in B. paucigranulata (Fig. 11 K, L) vs. absent in B. danieli (Fig. 11 O)]. • Barbaria ganczareki, by the dorsal sculpturing [both pillars and pores easily identifiable in B. paucigranulata (Figs 7 I, 8 I, 9 I, 10 I) vs. pillars so poorly developed that pores become the dominant element of the sculpture in B. ganczareki (Figs 7 C, 8 D, 9 C, 10 C)], the pores in the subcephalic region (absent in B. paucigranulata vs. present in B. ganczareki) and in pedal plates (absent in B. paucigranulata vs. present in B. ganczareki) and by the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. robust, hook-shaped and divergent from the claw branch in B. ganczareki, fig. 21 – 22 in Michalczyk & Kaczmarek (2007)]. • Barbaria hannae, by the dorsal sculpturing [no smooth plate portions in B. paucigranulata (Figs 8 I, 9 I, 10 I) vs. thickened plate portions devoid of sculpturing present in B. hannae (Figs 8 E, 9 D, E, 10 D, E)] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. robust, hook-shaped and divergent from the claw branch in B. hannae (Fig. 11 P)]. • Barbaria jenningsi, by the cirrus A length (<20 % of the body length in B. paucigranulata vs.> 50 % of the body length in B. jenningsi), the type of perforation in the dorsal plates [pores in B. paucigranulata (Fig. 7 I) vs. pseudopores in B. jenningsi (Fig. 7 E, F)] and by the primary spur morphology [slightly heteronych, needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. fully heteronych, robust, hook-shaped and divergent from the claw branch in B. jenningsi (Fig. 11 E, F)]. • Barbaria madonnae, by striae between pillars in the scapular and the caudal plate [present in B. paucigranulata (Figs 7 I, 10 I) vs. absent in B. madonnae (Figs 7 G, 10 G)], the shape of papilla IV [tubby in B. paucigranulata (Fig. 11 L) vs. elongated in B. madonnae (Fig. 11 H)] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. robust, hook-shaped and divergent from the claw branch in B. madonnae (Fig. 11 G, H)]. • Barbaria ollantaytamboensis, by the distribution of pores in plates [absent in the anterior portion of median plate 2 (Fig. 8 I), paired segmental plates (Fig. 9 I) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m 2 (Fig. 8 H), paired segmental plates (Fig. 9 H) and the entirety of m 3 in B. ollantaytamboensis] and claw isomorphy [slightly heteronych (heteromorphic) in B. paucigranulata (Fig. 11 K, L) vs. isonych (homomorphic) in B. ollantaytamboensis (Fig. 11 I, J)]. • Barbaria quitensis, by the type of perforation in the dorsal plates [pores in B. paucigranulata (Figs 7 I, 8 I, 9 I, 10 I) vs. pseudopores in B. quitensis (Figs 7 J, 8 J, 9 J, 10 J)] and the distribution of pores / pseudopores [scarcer on lateralmost portions of the caudal plate (Fig. 10 I) in B. paucigranulata vs. roughly equally distributed in all portions of the caudal plate (Fig. 10 J) in B. quitensis]. • Barbaria ranzii, by the cirrus A length (<20 % of the body length in B. paucigranulata vs.> 50 % of the body length in B. ranzii), the distribution of pores in plates [absent in the anterior portion of median plate 2 (Fig. 8 I), paired segmental plates (Fig. 9 I) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m 2 (Fig. 8 K), paired segmental plates (Fig. 9 K) and the entirety of m 3 in B. ranzii] and the by secondary spurs directed upwards on external claws IV [absent in B. paucigranulata (Fig. 11 L) vs. present in B. ranzii, Fig. 11 R]. • Barbaria weglarskae, by the cirrus A length (<20 % of the body length in B. paucigranulata vs.> 50 % of the body length in B. weglarskae) and by the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata (Fig. 11 K, L) vs. robust, hook-shaped and divergent from the claw branch in B. weglarskae (Fig. 11 M, N)].	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	description	(FIGS 4, 7 L, 8 L, 9 L, 10 L, 11 M, N; TABLE 5) Zoobank registration: u r n: l s i d: z o o b a n k. org: act: F 083907 B- 741 F- 48 C 0 - A 127 - 2 FC 28482 C 9 AA Morphometric data: w w w. t a r d i g r a d a. n e t / register / 0110. htm	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	materials_examined	Ty p e m a t e r i a l: Holotype (adult female, slide AR. 059.04) and two paratypes (adult females, slides AR. 059.01, 05). 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).	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	etymology	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.	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	description	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. 4 A). 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. 4 A, B). 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 4 A, B, 7 L, 8 L, 9 L, 10 L). 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. 4 A, B). Minute pores of equal size are regularly distributed in the remaining plate portions (Figs 4 A, B, 7 L, 8 L, 9 L, 10 L). The cephalic plate large, with a pronounced chalice-shaped anterior incision and lateral sutures demarcating roughly triangular lateralmost portions of the plate (Fig. 4 A, B). 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. 4 A, B). Median plates 1 and 3 unipartite (the latter strongly reduced and partially covered by the caudal plate), median plate 2 bipartite (Figs 4 A, B, 8 L). Paired segmental plates with broad, transverse, unsculptured bands (Figs 4 A, B, 9 L). The caudal plate with short, poorly sclerotized incisions clearly joined by a transversal suture (Figs 4 A, B, 10 L). Ventral cuticle with minute endocuticular pillars distributed evenly throughout the entire venter; a pair of small, subcephalic plates present (Fig. 4 C – D). Sexpartite gonopore (Fig. 4 E) placed anteriorly to a trilobed anus between legs IV. Pedal plates I – IV with evident pillars and pseudopores (Figs 4 A, 11 M, N). Evident pulvini on outer sides of all legs. Dentate collar IV with numerous irregular short teeth (Fig. 11 N). A tiny spine I and a tubby papilla IV (Fig. 11 N). Claws slightly heteronych, because primary spurs on internal claws IV are positioned higher than those on claws I – III (Fig. 11 M, N). 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.	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	diagnosis	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. 7 L) vs. pseudopores in B. jenningsi (Figs 7 E, F, 8 F, 9 F, 10 F)] and claw isomorphy [anisonych / slightly heteronych in B. weglarskae (Fig. 11 M, N) vs. strongly heteronych in B. jenningsi (Fig. 11 E, F)]. • Barbaria ranzii, by the pedal plate sculpturing [with evident pillars in B. weglarskae (Fig. 11 M, N) vs. without pillars in B. ranzii (Fig. 11 R)], the shape of papilla IV [tubby in B. weglarskae (Fig. 11 N) vs. elongated in B. ranzii] and by the presence of secondary spurs directed upwards on external claws IV [absent in B. weglarskae (Fig. 11 N) vs. present in B. ranzii (Fig. 11 R)].	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
586D3B6C02396145EC64CBFDFDA3FA70.taxon	discussion	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). 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 (HM 193406; Jørgensen et al., 2011) ranged between 2.6 and 2.9 % (alignment length = 585 bp). Analogous index for the pair B. weglarskae – B. jenningsi (KP 013596; 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: 18 S rRNA: 0.1 % (in B. bigranulata and B. charrua); 28 S rRNA: 0.1 – 0.4 % (B. bigranulata, B. ollantaytamboensis); ITS 1: 0.5 % (B. bigranulata), 0.1 – 1.0 % (B. charrua); ITS 2: 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: • 18 S rRNA: 0.0 – 2.5 % (1.2 % on average), with the most similar being B. charrua (MZ 820796) and B. danieli (MZ 820800); and the least similar being B. madonnae (MZ 820803) and B. ollantaytamboensis (MZ 820804). • 28 S rRNA: 0.0 – 4.3 % (2.4 % on average), with the most similar being B. charrua (MZ 820814) and B. danieli (MZ 820818); and the least similar being B. madonnae (MZ 820821) and B. ollantaytamboensis (MZ 820823). • ITS 1: 0.3 – 10.3 % (5.5 % on average), with the most similar being B. charrua (MZ 820833) and B. danieli (MZ 820836); and the least similar being B. madonnae (MZ 820839) and B. bigranulata (MZ 820828). • ITS 2: 1.6 – 10.8 % (8.1 % on average), with the most similar being B. charrua (MZ 822380) and B. danieli (MZ 822384); and the least similar being B. danieli + B. madonnae (MZ 822384, MZ 822387) and B. ollantaytamboensis (MZ 822388 – 91). • COI: 9.1 – 20.4 % (16.0 % on average), with the most similar being B. charrua (MZ 820850) and B. danieli (MZ 820853); and the least similar being B. ollantaytamboensis (MZ 820855) and B. weglarskae (MZ 820860). 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. 6 A) and welldelimited pedal plates on legs I – III with densely packed pillars and pseudopores (Fig. 6 B). Its papillae on legs IV were elongated (Fig. 6 C), meaning that they were much longer than wide. Internal claws were exhibiting heteronychy or lacking primary spurs (Fig. 6 D). In other words, among the extant species, B. bigranulata is morphologically overall the most similar to the last common ancestor of the analysed species.	en	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: 1037-1066, DOI: 10.1093/zoolinnean/zlab087
