taxonID	type	description	language	source
D9081578FF9EFF99FC8AF7F1FAA45A0D.taxon	type_taxon	Type species – Litoribates caelestis Pfingstl & Schatz, 2017	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF99FF9CFFD0FAEDFB945168.taxon	materials_examined	Type material / locality Holotype: adult female, Florida U. S. A, Florida Keys, Islamorada (FL _ 08), leaf litter under red mangrove (Rhizophora mangle); 13 February 2017, coll. T. Pfingstl and A. Lienhard, preserved in ethanol, deposited at the Naturhistorisches Museum Wien, Vienna. Four paratypes from the same sample, deposited in the collections of the US National Museum and the Senckenberg Museum für Naturkunde Görlitz, respectively. Additional non-type specimens are stored in the collections of the Institute of Biology, University of Graz.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF99FF9CFFD0FAEDFB945168.taxon	etymology	Etymology The specific epithet is a noun in the genitive case, referring to Florida, the type locality of this species.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF99FF9CFFD0FAEDFB945168.taxon	diagnosis	Diagnosis Adult instar brown sclerotized. Length 336 – 375 µm, width 231 – 259 µm. Notogaster rounded, almost circular in dorsal view. Slender lamellar ridges anteriorly converging. Sensillus slightly clavate and spinose at tip. Cerotegument finely granular, obviously larger granules in humeral areas. Irregular longitudinal cuticular ridges flanking lenticulus, conspicuous cuticular pattern absent in other notogastral areas. Fourteen pairs of setiform, notogastral setae. Epimeral setation 3 - 1 - 2 - 2, setae 1 b and 3 b the longest.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF99FF9CFFD0FAEDFB945168.taxon	discussion	Remarks Presently, there are only three species of this genus known. Litoribates floridae can be easily distinguished from the Pacific L. caelestis by the absence of the obvious reticulate cuticular surface pattern present in the latter and its larger size (mean length 360 µm vs. 322 µm). It is morphologically very similar with L. bonairensis but has several irregular longitudinal cuticular ridges in the humeral region that are either absent or only faintly developed in the latter species.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF99FF9CFFD0FAEDFB945168.taxon	description	Description Measurements. Females (N = 9), length: 336 – 375 μm (mean 368 μm), width: 240 – 259 μm (mean 249 μm); males (N = 7), length: 350 – 363 μm (mean 357 μm), width: 231 – 246 μm (mean 238 μm). Integument. Colour dark brown. Prodorsum. (Figures 3 (a) and 4 (a )) Cerotegument finely granular, larger granules next to anterior notogastral border and bothridia. Rostrum nearly triangular in dorsal view, projecting anteroventrally in lateral view; demarcated from remainder of prodorsum by faint transverse ridge. Pair of converging, slender lamellar ridges in slightly lateral position, reaching from bothridium to lamellar seta. Rostral seta (ro) robust, setiform, smooth (approx. 30 µm). Lamellar seta (le) setiform, short (approx. 15 µm), and smooth. Interlamellar seta (in) setiform (approx. 20 µm), exobothridial seta (ex) minute. Bothridium hardly protruding, borders not clearly defined; orifice narrow and circular. Sensillus (ss) long (approx. 75 µm), slightly bent caudally, slightly clavate, and spinose at the tip. Gnathosoma. Typical for the genus. Distal part of rutellum developed as thin triangular membrane, slightly curved inward with longitudinal incision (Figure 5 (a )). Setae a and m long (approx. 30 µm), robust, and smooth. Mentum regular, seta h setiform, thin (approx. 20 µm). Palp setation 0 - 2 - 1 - 3 - 8 (solenidion not included), trochanter very short, femur by far longest segment, genu, tibia, and tarsus of almost equal length (Figure 5 (b )). Chelicera with mobile digit darker sclerotized; distinct teeth interlocking. Träghård ’ s organ (tg) slender blunt finger-like oncophysis. Seta cha and chb dorsally slightly pectinate both of the same length (approx. 25 µm) (Figure 5 (c )). Gastronotic region. (Figures 3 (a) and 4 (a )) Conspicuously rounded in dorsal view, convex in lateral view; anterior notogastral margin distinct, forming a slightly overhanging bulge in area of lenticulus. Lenticulus slightly trapezoid with slightly irregular borders. Cerotegument finely granular, obviously larger granules in humeral areas next to lenticulus. Irregular short longitudinal cuticular ridges flanking lenticulus. Several small circular light spots posterior to seta c 1, representing sigilla of dorsoventral muscles. Fourteen pairs of setiform notogastral setae (8 – 16 µm), c 1 - 2, da, dm, dp, la, lm, lp, h 1 - 3, p 1 - 3; seta c 3 absent. Five pairs of notogastral lyrifissures present; ia laterad of seta c 2, hardly discernible due to strong cerotegumental granulation and cuticular ornamentation; im laterad of seta lm; ih laterad and anterior to h 3; lyrifissures ip and ips laterally of seta p 3 and p 2, respectively. Orifice of opisthonotal gland (gla) laterad, between seta lm and lp. Lateral aspect. (Figures 3 (c) and 4 (c )) Cerotegument mostly finely granular, larger granules on pedotectum I and in areas surrounding acetabula. Pedotectum I small rounded, slightly projecting. Pedotectum II absent. Discidium present, not conspicuously expressed. Van der Hammen ’ s organ present; typical for the genus (for details, see Pfingstl and Schatz 2017). Podosoma and venter. (Figures 3 (b) and 4 (b )) Cerotegument finely granular. Epimeral setation 3 - 1 - 2 - 2, all setae setiform and smooth. Setae 1 b and 3 b longest (approx. 18 µm), others shorter (10 µm). Internal borders of all epimera well visible. Median longitudinal small ridge on epimeron I, other sternal surface lines slightly protruding with small indentations at each transition from one epimeral segment to the other. Genital and anal opening closely adjacent, both surrounded by slightly darker cuticle. Rounded genital plates with four pairs of fine setae (approx. 10 µm). One pair of setiform aggenital setae ag. Anal valves strongly triangular. Outer part of preanal organ rectangular with rounded edges, inner part shaped like transverse bar. Two pairs of short anal setae, an 1 - 2 (approx. 10 µm). Three pairs of short adanal setae, ad 1 - 3 (approx. 8 µm). Lyrifissure iad flanking posterior third of anal plates. Legs. (Figure 6) Monodactylous and slender. Long, strong hook-like claws, with inconspicuous dorsal dentition. Cerotegument generally finely granular, larger granules only on distal third of all femora. Trochanter III and IV with obvious dorsal spur. All genua with ventral transversal ridge. Large elongate porose areas on ventral paraxial side of femora I and II and on paraxial dorsal aspect of femora III and IV. Kidney-shaped porose areas on paraxial dorsal aspect of trochanters III and IV. Dorsal seta d on all femora slightly thickened and barbed on outer curvature. Lateral setae of femora and genua I and II blunt, short, broadened, and slightly serrate. Ventral setae of all tibiae and tarsi long and slightly serrate ventrally. Tibia IV with two setae. Solenidia φ 1 - 2 on tibia I borne on vaguely delimited mound. Chaetome and solenidia given in Table 2.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF9CFC8AFC86FAE951BC.taxon	type_taxon	Type species – Thalassozetes riparius Schuster, 1963	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	materials_examined	Type material / locality Holotype: adult female preserved in ethanol, Panama, Bocas del Toro, Isla Colon, Paunch (PA _ 37), intertidal algae (Bostrychia) from rock, 7 February 2017, deposited at the Naturhistorisches Museum Wien. Four paratypes preserved in ethanol: adult females; Panama, Bocas del Toro, Isla Colon, Boca del Drago (PA _ 43), intertidal algae (Bostrychia) from rock, deposited at Museo de Invertebrados Fairchild, Universidad de Panamá. Additional non-type specimens from all locations are stored in the collections of the Institute of Biology, University of Graz.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	etymology	Etymology As this species was found in Panama and Florida, the name was chosen to somehow relate to both locations. The specific epithet “ balboa ” (given as noun in apposition) refers to the Spanish explorer Vasco Núñez de Balboa; he led the first European expedition that crossed the Isthmus of Panama in 1513 and the Panamanian currency is named after him. It also refers to Rocky Balboa, a well-known movie character, a descendant of immigrants who made his fortune in the USA. The new species similarly may be an immigrant that has now successfully settled in North America. Moreover, T. balboa sp. nov. was predominantly found on “ rocky ” substrate.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	diagnosis	Diagnosis Dark brown sclerotized mites. Length 277 – 312 µm, width 154 – 194 µm. Notogaster oval in dorsal view with characteristic reticulate-foveate pattern. Lamellar ridges present. Sensillus clavate, distal half spinose. Pair of weak anterior notogastral ridges present, mediad of elliptic depressions. Fourteen pairs of setiform, notogastral setae. Median sternal depression on epimeron I, shaped like inverted pear. Three pairs of genital setae, two pairs of adanal and anal setae. Lyrifissure iad oblique, next to anterior corner of anal orifice. Legs monodactylous, claws with two ventral teeth.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	discussion	Remarks T. balboa sp. nov. can be distinguished from the other three Thalassozetes species by its specific reticulate-foveate cuticular pattern, with the centre of notogaster covered with regularly distributed conglomerates of irregular circular or elliptic depressions, fading laterally and caudally in smaller foveate pattern; by two pairs of adanal setae instead of three; and by having two ventral teeth on each claw instead of one.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	description	Description Measurements. Females (N = 26), length: 280 – 312 μm (mean 303 μm), width: 172 – 194 μm (mean 182 μm); males (N = 64), length: 277 – 305 μm (mean 289 μm), width: 154 – 182 μm (mean 170 μm). Integument. Colour dark brown. Prodorsum. (Figures 7 (a) and 8 (a )) Cerotegument finely granular. Rostrum nearly triangular in dorsal view, projecting anteroventrally in lateral view. Rostrum demarcated from remainder of prodorsum by faint transverse ridge. Pair of inward-curving, slender lamellar ridges, reaching from bothridium to lamellar seta. Rostral seta (ro) and lamellar seta (le) short, setiform, smooth (approx. 5 – 6 µm). Interlamellar seta (in) short and setiform (approx. 5 µm), exobothridial seta (ex) minute. Bothridium large and strongly protruding, orifice circular. Sensillus of normal length (approx. 55 µm), clavate, distal half spinose. Gnathosoma. Typical for genus. Distal part of rutellum developed as thin triangular membrane, slightly curved inward with obvious longitudinal incision (Figure 9 (a )). Setae a and m robust and slightly barbed unilaterally (approx. 15 µm). Mentum regular, seta h setiform, robust (approx. 20 µm). Palp setation 0 - 2 - 1 - 3 - 8 (solenidion not included), trochanter very short, femur longest segment, genu, tibia and tarsus of almost equal length (Figure 9 (b )). Cheliceral digits with two teeth. Seta cha dorsally and chb laterally slightly pectinate, both setae of the same length (approx. 25 µm) (Figure 9 (c )). Träghård ’ s organ (tg) slender blunt finger-like oncophysis. Gastronotic region. (Figures 7 (a) and 8 (a )) Oval in dorsal view, convex in lateral view; dorsosejugal suture incomplete. Nearly triangular clear spot (vestigial lenticulus) with irregular borders on anterior notogastral median area. Cuticle with specific reticulate-foveate pattern, centre of notogaster covered with regularly distributed conglomerates of irregular circular or elliptic depressions, fading laterally and caudally in smaller foveate pattern. Pair of weak and highly variable (in shape) concave notogastral ridges framing light spot. Laterad of ridges, elliptic depressions lined with fine and densely packed granules. Fourteen pairs of setiform notogastral setae (approx. 10 µm), c 1 - 2, da, dm, dp, la, lm, lp, h 1 - 3, p 1 - 3; seta c 3 absent. Five pairs of notogastral lyrifissures present; ia in humeral area adjacent to anterior border of elliptic cavity; im laterad of seta la; ih laterad and anterior to h 3; lyrifissures ip and ips laterally of seta p 3 and p 2 respectively. Orifice of opisthonotal gland (gla) laterad between seta la and lm. Lateral aspect. (Figures 7 (c) and 8 (c )) Cerotegument basically finely granular, larger granules on pedotectum I and in areas surrounding acetabula. Lateral sejugal furrow broad and deep, with dense granulation. Next to lateral border of bothridium triangular protrusion orientated caudally, slightly projecting above lateral sejugal furrow. Pedotectum I small rounded, slightly projecting. Pedotectum II absent. Lateral enantiophysis present, anterior projection triangular, well developed, posterior protrusion triangular and small. Discidium present, developed as strongly projecting triangular bulge between acetabulum III and IV. Podosoma and venter. (Figures 7 (b) and 8 (b )) Cerotegument with larger granules next to acetabula and fine granules surrounding anal opening. Epimeral setation 1 - 0 - 1 - 1, all setae setiform and smooth, seta 1 b longest (approx. 45 µm), others significantly shorter (ca. 7 µm). Internal borders of all epimera well visible. A densely granulated median sternal cavity on epimeron I with slightly concave lateral borders resulting in inverted pear-like shape. Rounded genital plates with three pairs of fine setae (approx. 8 µm) arranged in longitudinal rows. Aggenital setae absent. Lyrifissure iad oblique adjacent to anterior corner of anal orifice. Outer part of preanal organ rectangular with rounded edges, inner part shaped like a transverse bar. Two pairs of short anal setae, an 1 - 2 and two pairs of short adanal setae, ad 1 - 3 (all approx. 8 µm). Legs. (Figure 10) Monodactylous. Long, strong hook-like claws, dorsally with slight dentation, hardly visible. Two ventral teeth on each claw, proximal one conspicuous and distal one weakly developed. Cerotegument generally finely granular, larger granules on all trochanters and femora. Femora with ventral carina. No porose areas discernible. Femoral setae serrate. Lateral setae of all genua thickened, blunt and slightly serrate. Ventral setae of tarsus serrate. Famulus ε developed as short broad knob. Solenidion ω 2 in lateral position. Chaetome and solenidia see Table 2.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
D9081578FF9BFF90FC8AFC11FBFD56A7.taxon	discussion	Morphometrics Univariate Statistics: Litoribates. The two Litoribates species L. bonairensis and L. floridae differ significantly in only 5 of the 15 measured variables (Table 3). The variability as indicated by cv is moderate (cv ≤ 0.10) in all characters. Multivariate analyses: Litoribates. The two Litoribates species can be separated only partly by PCA on raw data and totally overlap in the PCA on size-corrected data (Figure 11). In the PCA on raw data, PC 1 (explaining 38.5 % of the total variation) is responsible for the slight separation. The variable with by far the highest loading on PC 1, and thus contributing most to the separation, is the lenticulus length ll (Table 4). PC 1 also correlates slightly with “ size ” as defined by the geometric mean (r = 0.81), indicating that L. floridae individuals are slightly larger than those of L. bonairensis. The combination of PC 1 and PC 2 (explaining 25.0 % of the total variation) reveals a sexual dimorphism. In L. floridae, males and females are clearly separated, while the two sexes overlap in L. bonairensis. The variable contributing the most to separation along PC 2 is the length of the genital opening gl (Table 4). Size correction resulted in a decrease of 87.1 % of the total variation. After size correction, PCA no longer separates the two Litoribates species (Figure 11). The sexual dimorphism of L. floridae is not visible in the size-corrected data, but a slight separation of the sexes becomes apparent along PC 2 in L. bonairensis. The variable with the highest loading on PC 2 is dcg (Table 4). The LDA on both raw and size-corrected data revealed that the two species are mainly separated by the variable ll, and the Hotelling ’ s T-test showed highly significant differences (p <0.001) between them. In the raw data, 100 % of the individuals can be correctly classified by all-samples LDA and 80 % after leave-one-out cross-validation. After size correction, all-samples LDA still correctly classifies 97.14 % and leave-one-out cross-validation LDA correctly classifies 74.29 %. Univariate Statistics: Thalassozetes. Thalassozetes barbara from Barbados and T. balboa from Panama differ significantly in 11 of the 20 measured variables (Table 5). The variability of almost all characters is moderate with a cv ≤ 0.10 in all populations, only efw 2 shows slightly higher values (cv between 0.11 and 013). Multivariate analyses: Thalassozetes. In PCA on both raw and size-corrected data, PC 1 (explaining 33.6 % of the total variation in raw data and 29.1 % in size-corrected data) separated T. barbara from T. balboa with a very small overlapping area (Figure 12). The variation within T. balboa seems to be larger than that of T. barbara, and the larger variation is not caused by different sample sites as the three populations from Panama always overlap. In PCAs on both raw and size-corrected data, the variables with the highest loadings on PC 1 were ll and efw 2 (Table 6). A pronounced sexual dimorphism is revealed by PC 2 (explaining 25.6 % and 17.0 % of the total variation, respectively) in both raw and size-corrected data (Figure 12). The differences between the two sexes are more pronounced in T. barbara than in T. balboa PC 2 is strongly correlated with size (geometric mean) in the raw data (r = 0.96), indicating that females of both species are considerably larger than males. Variables contributing most to PC 2 were gl and gw, length and width of the genital opening. In the size-corrected data, there is only a weak correlation between PC 2 and size (r = 0.68). Here, the differences between the two sexes are again mainly caused by the variables gl and gw, which have the highest loadings for PC 2 (Table 6). The total variation decreased strongly, by 88.7 %, after size correction. LDA showed again that separation of T. barbara from T. balboa was mainly caused by the variables ll and efw 2 in both raw and size-corrected data. All-samples LDA on raw data correctly classified 100 % of all specimens and 96.3 % after leave-one-out cross-validation. After size correction, the percentages of correctly classified individuals slightly dropped to 99.1 % in all-samples LDA and increased to 97.3 % after leave-one-out cross-validation, Hotelling ’ s T-test revealed highly significant differences between the two species. Molecular genetic analyses To ensure the genetic distinctness of the investigated Litoribates species, classical barcoding by means of COI data was applied (Figure 13). The COI topology clearly separates L. floridae and L. bonairensis. The distinctness of these two species is underlined by a clear “ barcoding gap ” with high genetic differences. The highest intraspecific distance (uncorrected p - distance) amounted to 1.4 % and the lowest interspecific distance to 9.6 %. The 18 S topology clearly demonstrates the monophyly of all investigated genera (Figure 14). Although the 18 S rRNA gene is typically not suitable for species identification, T. balboa and T. barbara could be clearly separated with high statistical support. Thalassozetes species form the sister group of the Schusteria and Thasecazetes clade and together with the recently described Indopacifica species and Rhizophobates (= “ Thalassozetes shimojanai, ” for explanation of diverging taxon name, please refer to discussion) they build a monophyletic clade of Selenoribatidae. Litoribates spp. are placed as sister group of the Pacific Alismobates species; however, members of Fortuyniidae are shown to be paraphyletic with the genus Fortuynia being placed closer to Selenoribatidae.	en	Pfingstl, Tobias, Lienhard, Andrea, Baumann, Julia (2019): New and cryptic species of intertidal mites (Acari, Oribatida) from the Western Caribbean - an integrative approach. International Journal of Acarology 45 (1 - 2): 10-25, DOI: 10.1080/01647954.2018.1532458, URL: https://doi.org/10.1080/01647954.2018.1532458
