Neotropical jewels in the moss: biodiversity, distribution and evolution of the genus Barbaria (Heterotardigrada: Echiniscidae) Author Gąsiorek, Piotr Author Wilamowski, Andrzej Author Vončina, Katarzyna Author Michalczyk, Łukasz text Zoological Journal of the Linnean Society 2022 2022-01-25 195 1037 1066 journal article 121550 10.1093/zoolinnean/zlab087 bafaf468-919e-467d-a31a-f8af68920e24 0024-4082 6993173 87E03803-7D69-4393-93CD-24A39D607972 BARBARIA PAUCIGRANULATA WILAMOWSKI, VONČINA, GĄSIOREK & MICHALCZYK, SP. NOV. ( FIGS 1–3 , 7I , 8I , 9I , 10I , 11K, L; TABLES 3, 4) Zoobank registration: u r n: l s i d: z o o b a n k. org:act: A11E6BDF-7E2A-4F0A-BD6A-9BC87FCE147A Morphometric data: w w w. t a r d i g r a d a. n e t / register/0109.htm 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 ). Etymology: From Latin paucus , few, and granulatus , grained, alluding to the scarcity of epicuticular granulation on the dorsal plates. An adjective in nominative singular. 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 1A, B , 2 ). Cylindrical, Echiniscus - type cephalic papillae (secondary clavae) and (primary) clavae; cirri embedded in bulbous cirrophores ( Fig. 3D ). Cirrus A is short (<20% of the body length) and with evident, but small cirrophore ( Fig. 1A, B ). Dorsal plate sculpturing of the bigranulata - type , comprising minute, poorly developed pillars (pseudogranulation) and pores ( Figs 1C , 2 , 3A–C , 7I , 8I , 9I , 10I ). Pillars are densely packed and additionally interconnected by thin striae in the scapular ( Fig. 7I ) and caudal (terminal) plates ( Fig. 10I ). Pores are slightly larger in the scapular plate, median plate1, 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. 8I ), paired segmental plates ( Fig. 9I ) 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 1A, 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 1A , 2A ). Lateral sutures in the scapular plate demarcate lateralmost, trapezoidal portions ( Figs 1A, B , 2B ). Median plates 1 and 3 are unipartite (the latter with strongly developed granules), median plate 2 is bipartite ( Fig. 8I ). Paired segmental plates are without transverse unsculptured bands, as epicuticular granules of anterior portion transition gradually into pillars of the posterior portions ( Fig. 9I ). The caudal plate with short, poorly sclerotized incisions and no signs of faceting ( Figs 1A , 2 , 10I ). Ventral cuticle with minute endocuticular pillars ( Fig. 11K ) distributed evenly throughout the entire venter, lacking plates, beside of rarely developed subcephalic plates ( Fig. 3D ). 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 1A, B , 11K, L ). Thick pulvini on outer side of all legs ( Figs 1A, B , 2B ). Dentate collar IV has numerous irregular short teeth ( Fig. 11L ). A small, elongated spine I and a tubby papilla IV ( Figs 1B , 2B , 11K, L ). Claws slightly heteronych with claws IV ( Fig. 3F ) higher and more robust than claws I–III ( Fig. 3E ). Internal claws IV have needle-like spurs more divergent from branches compared to spurs I– III ( Fig. 11K, 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. Figure 1. Habitus of Barbaria paucigranulata (PCM) : A, holotype (female) in dorsolateral view; B, paratype (female) in lateral view; C, dorsal sculpturing in close-up. Abbreviations: Ia–IIb, paired segmental plates; c, caudal plate; cA, cirrus A ; ce, cirrus externus; ci, cirrus internus; cl, (primary) clava; co, dentate collar IV; cp, cephalic plate; cv, cervical plate; m1–3, median plates; p, cephalic papilla (secondary clava); pIV, papilla IV; pl, pulvinus; pp, pedal plate; sI, spine I; sc, scapular plate. Scale bars in µm. Figure 2. Habitus of Barbaria paucigranulata (SEM) : A, paratype (sex undetermined) in dorsal view; B, paratype (female) in lateral view. Scale bars in µm. 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. Figure 3. Details of Barbaria paucigranulata (SEM) : A, pores in the scapular plate; B, pores in the posterior portion of the paired segmental plate II; C, pores in the caudal plate; D, cephalic appendages and a pair of subcephalic plates; E, claws I; F, claws IV. Scale bars in µm. 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. 8I ), paired segmental plates ( Fig. 9I ) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m2 ( Fig. 8A ), paired segmental plates ( Fig. 9A ) and the entirety of m 3 in B. bigranulata ], the shape of papilla IV [tubby in B. paucigranulata ( Fig. 11L ) vs. elongated in B. bigranulata ( Fig. 11B )] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata ( Fig. 11K, L ) vs. robust, hook-shaped and divergent from the claw branch in B. bigranulata ( Fig. 11A, B )]. • Barbaria charrua , by the shape of papilla IV [tubby in B. paucigranulata ( Fig. 11L ) vs. elongated in B. charrua ( Fig. 11D )] and by the primary spurs [present in B. paucigranulata ( Fig. 11K, L ) vs. typically absent in B. charrua ( Fig. 11C, D )]. • Barbaria danieli , by the shape of papilla IV [tubby in B. paucigranulata ( Fig. 11L ) vs. elongated in B. danieli ( Fig. 12 )] and by the primary spurs [present in B. paucigranulata ( Fig. 11K, L ) vs. absent in B. danieli ( Fig. 11O )]. • Barbaria ganczareki , by the dorsal sculpturing [both pillars and pores easily identifiable in B. paucigranulata ( Figs 7I , 8I , 9I , 10I ) vs. pillars so poorly developed that pores become the dominant element of the sculpture in B. ganczareki ( Figs 7C , 8D , 9C , 10C )], 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. 11K, 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 8I , 9I , 10I ) vs. thickened plate portions devoid of sculpturing present in B. hannae ( Figs 8E , 9D, E , 10D, E )] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata ( Fig. 11K, L ) vs. robust, hook-shaped and divergent from the claw branch in B. hannae ( Fig. 11P )]. • 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. 7I ) vs. pseudopores in B. jenningsi ( Fig. 7E, F )] and by the primary spur morphology [slightly heteronych, needle-like and adjacent to the claw branch in B. paucigranulata ( Fig. 11K, L ) vs. fully heteronych, robust, hook-shaped and divergent from the claw branch in B. jenningsi ( Fig. 11E, F )]. • Barbaria madonnae , by striae between pillars in the scapular and the caudal plate [present in B. paucigranulata ( Figs 7I , 10I ) vs. absent in B. madonnae ( Figs 7G , 10G )], the shape of papilla IV [tubby in B. paucigranulata ( Fig. 11L ) vs. elongated in B. madonnae ( Fig. 11H )] and the primary spur morphology [needle-like and adjacent to the claw branch in B. paucigranulata ( Fig. 11K, L ) vs. robust, hook-shaped and divergent from the claw branch in B. madonnae ( Fig. 11G, H )]. Figure 4. Habitus of Barbaria weglarskae (PCM) : A, holotype (female) in dorsal view; B, dorsal sculpturing in close-up (paratype); C, subcephalic plates (paratype); D, subcephalic plates (holotype); E, gonoporal area (holotype). Scale bars in µm. Figure 5. Phylogeny of the genus Barbaria based on five concatenated markers conducted in MrBayes and W-IQ-TREE (values above the nodes signify posterior probabilities, whereas bootstraps are provided below the nodes; maximal supports are denoted with asterisks). Diploechiniscus oihonnae and Testechiniscus spitsbergensis tropicalis were used as an outgroup. The scale bar represents 0.2 substitutions per nucleotide position and refers to the Bayesian inference. • Barbaria ollantaytamboensis , by the distribution of pores in plates [absent in the anterior portion of median plate 2 ( Fig. 8I ), paired segmental plates ( Fig. 9I ) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m2 ( Fig. 8H ), paired segmental plates ( Fig. 9H ) and the entirety of m 3 in B. ollantaytamboensis ] and claw isomorphy [slightly heteronych (heteromorphic) in B. paucigranulata ( Fig. 11K, L ) vs. isonych (homomorphic) in B. ollantaytamboensis ( Fig. 11I, J )]. • Barbaria quitensis , by the type of perforation in the dorsal plates [pores in B. paucigranulata ( Figs 7I , 8I , 9I , 10I ) vs. pseudopores in B. quitensis ( Figs 7J , 8J , 9J , 10J )] and the distribution of pores/pseudopores [scarcer on lateralmost portions of the caudal plate ( Fig. 10I ) in B. paucigranulata vs. roughly equally distributed in all portions of the caudal plate ( Fig. 10J ) 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. 8I ), paired segmental plates ( Fig. 9I ) and the entirety of median plate 3 in B. paucigranulata vs. present in the anterior portion of m2 ( Fig. 8K ), paired segmental plates ( Fig. 9K ) 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. 11L ) vs. present in B. ranzii , Fig. 11R ]. • 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. 11K, L ) vs. robust, hook-shaped and divergent from the claw branch in B. weglarskae ( Fig. 11M, N )].