Fig 1.
Light micrographs of the holotypic adult male of Nanaloricus valdemari sp. nov.
Anterior faces up in all aspects. (A) Ventral view of the extended specimen. Note that the mouth tube is not fully extended. (B) Dorsal view of the specimen. Note the pair of longitudinal stripes (ls) spanning the anterior two thirds of the dorsal plate of the lorica. (C) Anterior region of the specimen, dorsal view. (D) Close-up of the posterior region of the introvert and neck, ventral view. White arrowhead points to the small, midventral anterior spike. Arrows point to trichoscalid sensory organs. Abbreviations: af, anal field; bpa/b, basal plate of type a or b; cs, clavoscalid; dp, dorsal plate; fu, oral furca; in, introvert; lo, lorica; mc, mouth cone; mt, mouth tube; sp (and black arrowheads), anterior spike; sr8/9, spinoscalid of 8th or 9th row; ss, spinoscalid; tp, trichoscalid plate; tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 2.
Line art drawing of the adult male habitus of Nanaloricus valdemari sp. nov.
Ventral view of the body, anterior faces up. Note that only a selected number of scalids from rows 1 to 8 are represented for clarity. Abbreviations: af, anal field; bpa/b, basal plates of type a and b; bu, buccal tube; cs, clavoscalid; dlp, dorsolateral plate; fu, oral furca; gl?, putative gland outlet; in, introvert; lo, lorica; mc, mouth cone; mo, mouth aperture; mt, mouth tube; mvs, midventral anterior spike; ne, neck; or1/2, oral ridges of type 1 and 2; sr2–9, spinoscalids of 2nd to 9th row (including sr4a/b, i.e. type a and b spinoscalids of the 4th row); sp (and black arrowheads), anterior spike; to, trichoscalid sensory organ; tp, trichoscalid plate; tr1, single trichoscalid; tr2, double trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 3.
Line art drawing of the adult male partial habitus of Nanaloricus valdemari sp. nov.
Dorsal view of the lorica, anterior faces up. Abbreviations: ac, anal cone; dlp, dorsolateral plate; dp, dorsal plate; fl, flosculum; go?, putative gonopore; ls, longitudinal stripe; sp (and arrowhead), anterior spike.
Fig 4.
Light micrographs of the allotypic adult female of Nanaloricus valdemari sp. nov.
Anterior faces up in all aspects. (A) Overview of the specimen, dorsal view. Note that the mouth tube is not fully extended. Note also the pair of longitudinal stripes (ls) spanning the anterior two thirds of the dorsal plate of the lorica. (B) Close-up of the anterior region of the specimen, dorsal view. Arrow points to the two most distal segments of a clavoscalid. (C) Close-up of the introvert, neck and anterior region of the abdomen, dorsal view. (D) Posterior region, ventral view. Double-headed arrows point to large pores (gland outlets?) situated on the posterior region of the ventral plate. (E) Posterior region, dorsal view. Note the putative gonopores (go?) located on the posteriormost region of the dorsolateral plates. Abbreviations: af, anal field; bu, buccal tube; cc, cuticularized crest; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; lo, lorica; mc, mouth cone; or, oral ridge; sp (and black arrowheads), anterior spike; ss, spinoscalid; tr, trichoscalid; vp, ventral plate.
Fig 5.
Light micrographs of a paratypic adult female of Nanaloricus valdemari sp. nov.
Anterior faces up in both aspects, dorsal view. (A) Overview of the specimen with extended introvert and fully extended mouth tube. (B) Close-up of the anterior region of the body. Abbreviations: bu, buccal tube; cs, clavoscalids; in, introvert; lo, lorica; ls, longitudinal stripe; mc, mouth cone; mt, mouth tube; or, oral ridge; sp, anterior spike; ss spinoscalid; tr, trichoscalid.
Fig 6.
Light micrographs of three live specimens of Nanaloricus valdemari sp. nov.
Anterior faces up in all aspects. (A) Overview of a female adult with head almost fully retracted. Internally, note the presence of the large ovaries (ov) in the abdomen. (B) Overview of a female adult (different from that shown in A). Note the pair of longitudinal stripes (ls) spanning the anterior two thirds of the dorsal plate of the lorica. (C and D) Overview of a male adult with head almost fully retracted. Note the presence, within the abdomen, of a testis (te) containing spermatozoa (sz). Note also the relative position of the pharyngeal bulb (pb) in this retracted animal. Abbreviations: af, anal field; bu, buccal tube; cs, clavoscalid; dp, dorsal plate; lo, lorica; sp, anterior spike; ss spinoscalid; tr, trichoscalid.
Fig 7.
Scanning electron micrographs of three paratypic adult males of Nanaloricus valdemari sp. nov.
(A) Overview of extended specimen, ventral view, anterior faces up. Same specimen as in C and D. Note the multiform clavoscalids: the ventralmost pair is unbranched (and similar to the eight female clavoscalids), while the others are divided into three branches. (B) Close up of the mouth tube (mt). Same specimen as in E. Note the honeycomb sculpture (hs) of the cuticle. (C) Anterior region of the body, ventral view. Note the difference between single (tr1) and double trichoscalids (tr2). White arrowhead points to the small midventral anterior spike. White arrows point to lorical pores. (D) Close up of the midventral pair of single trichoscalids, each of which protrudes from a plate characterized by a large pore (white double-headed arrow). The inset shows a magnification of the double trichoscalid sensory organ (to), which possesses a small pore (black arrows). (E) Close up of spinoscalids of 9th row (sr9) and basal plates, one of each type (bpa and bpb). Note the long hairs (ha) scattered along the length of the distal segments of a spinoscalid of 7th (?) row. (F) Detail of a cluster of flosculi arranged in a rectangular pattern on the dorsolateral plate. Same specimen as in G. (G) Overview of the posterodorsal region of the lorica. Note the pair of longitudinal stripes (ls) that span the anterior two thirds of the dorsal plate (dp). Black double arrowheads indicate the clusters of flosculi situated on the dorsolateral plates (dlp), while the white double arrowheads point to the small flosculum situated on the dorsal plate. Abbreviations: ac, anal cone; cs, clavoscalid; dlp, dorsolateral plate; fu, oral furca; lo, lorica; mc, mouth cone; or, oral ridge; ss, spinoscalid; sp, anterior spike; tp, trichoscalid plate; tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 8.
Schematic diagrams of the distribution of introvert and neck appendages in adult forms of Nanaloricus valdemari sp. nov. and Nanaloricus mathildeae sp. nov.
Note that for sake of clarity clavoscalid sexual dimorphism is not depicted. Note also that the shaded area indicates the neck region. (A) Polar diagram. (B) Planar projection. Abbreviations: bpa/b, basal plates of type a or b; cs, clavoscalid; MD, middorsal line; MV, midventral line; sr1–9, scalids of 1st to 9th row; ss, spinoscalid; tp, trichoscalid plate; tr1, single trichoscalid; tr2, double trichoscalid.
Fig 9.
Light micrographs of the holotypic adult male of Nanaloricus mathildeae sp. nov.
Anterior faces up in all aspects. (A) Ventral view of the extended specimen. Black arrowheads indicate anterior spikes. The inset shows a detail of the mouth tube (mt), which is not fully extended. (B) Dorsal view of the specimen. (C) Close-up of multiform clavoscalids (cs). (D) Close-up of the introvert and anterior spikes of the dorsal plate. Note the presence of basal plates of type a and b (bpa/b), and spinoscalids of 9th row (sr9); (E) Close-up of the anterior spikes of the ventral and ventrolateral plates. White arrowhead points to the small midventral anterior spike. Abbreviations: af, anal field; cc, cuticularized crest; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; in, introvert; lo, lorica; mc, mouth cone; mt, mouth tube; or, oral ridge; ss spinoscalid; to, trichoscalid sensory organ; tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 10.
Line art drawing of the adult male habitus of Nanaloricus mathildeae sp. nov.
Ventral view, anterior faces up. Note that only a selected number of scalids from rows 1 to 8 are represented for clarity. Abbreviations: af, anal field; ba, mouth cone bar; bpa/b, basal plates of type a and b; cs, clavoscalid; dlp, dorsolateral plate; fu, oral furca; gl?, putative gland outlets; in, introvert; lo, lorica; mc, mouth cone; mo, mouth aperture; mt, mouth tube; mvs, midventral anterior spike; ne, neck; or1/2, oral ridges of type 1 and 2; sr2–9, spinoscalids of 2nd to 9th row (including sr4a/b, i.e. spinoscalids of 4th row of type a and b); sp (and black arrowheads), anterior spike; to, trichoscalid sensory organ; tp, trichoscalid plate; tr1, single trichoscalid; tr2, double trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 11.
Light micrographs of the allotypic adult female of Nanaloricus mathildeae sp. nov.
Anterior faces up in all aspects. (A) Ventral view of the extended specimen. Note that the mouth tube is not fully extended. Black arrowheads indicate anterior spikes. (B) Close-up of anterior body half, ventral view. (C) Introvert, neck and anterior region of the abdomen, dorsal view. (D) Close-up of anterior spikes of the ventral and ventrolateral plates. White arrowhead points to the very small midventral anterior spike. (E) Close up of the posterior region of the lorica, dorsal view. Note the putative gonopores (go?) located on the posteriormost region of the dorsolateral plate. Double arrowheads point to the small flosculum situated on the dorsal plate. (F) Detail of a cluster of flosculi arranged in a rectangular pattern on the right dorsolateral plate. Abbreviations: af, anal field; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; lo, lorica; mc, mouth cone; mt, mouth tube; ss spinoscalid; tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 12.
Light micrographs of a paratypic adult female of Nanaloricus mathildeae sp. nov.
Anterior faces up in all aspects. (A) Overview of the fully extended specimen. Note that the mouth tube is fully extended. (B) Close up of the anterior region of the body. (C) Detail of the anteriormost region of the mouth tube. Short, black arrowheads point to ring-like thickenings of the fully extended mouth tube. Abbreviations: bu, buccal tube; cs, clavoscalids; fu, oral furca; lo, lorica; mc, mouth cone; mo, mouth aperture; mt, mouth tube; or, oral ridge; sp, anterior spike; ss spinoscalid; tr, trichoscalid.
Fig 13.
Light micrographs of a live adult female of Nanaloricus mathildeae sp. nov.
Anterior faces up in both aspects. Note the video sequence of this animal in the S1 Video. (A and B) Overview of the body with partly extended introvert and mouth cone. Internally, note the presence of a large oocyte (oo) within each ovary (ov) in the abdomen. Note also the shape and relative position of the pharyngeal bulb (pb). Abbreviations: af, anal field; cs, clavoscalid; lo, lorica; mc, mouth cone; mt, mouth tube; sp, anterior spike; ss spinoscalid.
Fig 14.
Scanning electron micrographs of a paratypic adult male of Nanaloricus mathildeae sp. nov.
(A) Overview of the extended specimen, ventral view, anterior faces up. (B) Close up of the anterior body region. Note the broad, multiform clavoscalids (cs). Clavoscalid bases are marked with asterisks (*). (C) Close up of spinoscalids from different rows, each with distinct gross morphology. Note the numerous long hairs (ha) scattered along the length of the various spinoscalids. (D) Close up of a dorsolateral single trichoscalid (tr1) and a midlateral (right-hand side) double trichoscalid (tr2). Note the relatively small sensory organ (to) of the double trichoscalid. (E) Close up of the trichoscalid sensory organ with a small pore (black arrow). Note also a type b basal plate (bpb) of the anterior neck region. (F) Overview of the posterodorsal region of the lorica. Black double arrowheads indicate the cluster of flosculi situated on each dorsolateral plate, while the white double arrowheads point to the small flosculum of the dorsal plate. (G) Detail of a dorsolateral cluster of flosculi arranged in a rectangular pattern. Abbreviations: ac, anal cone; af, anal field; ba, mouth cone bar; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; fu, oral furca; mc, mouth cone; mt, mouth tube; or, oral ridge; ss, spinoscalid; sp, anterior spike; sr2-4a/b,7,8, spinoscalid of 2nd to 4th (type a and b), 7th and 8th rows; vlp, ventrolateral plate; vp, ventral plate.
Fig 15.
Light micrographs of the holotypic adult female of Scutiloricus hugoi gen. et sp. nov.
Anterior faces up in all aspects. (A) Overview of the extended specimen, with focus on the internal buccal tube (bu). Double-headed arrow points to large pores (gland outlets?) situated on the posterior region of the ventral plate. (B) Ventral view of the specimen. (C) Close-up of the introvert, neck and anterior region of the lorica, ventral view. Note the fenestrated aspect of the anterior spikes (sp). (D) Close-up of the anal field. (E) Dorsal view of the extended specimen. Abbreviations: ac, anal cone; af, anal field; bp, basal plate; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; in, introvert; lo, lorica; mt, mouth tube; oo, oocyte; sp, anterior spike; sr8/9, spinoscalids of 8th or 9th row; ss spinoscalid; su, anal field spurs tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 16.
Line art drawing of the adult female habitus of Scutiloricus hugoi gen. et sp. nov.
Ventral view, anterior faces up. Note that only a selected number of scalids from rows 2 to 8 are represented for clarity. Abbreviations: ac, anal cone; af, anal field; ag?, putative adhesive gland; bp, basal plate; bu, buccal tube; cr, cuticularized triangular ridge; cs, clavoscalid; dlp, dorsolateral plate; fu, oral furca; gl?, putative gland outlet; in, introvert; lo, lorica; mc, mouth cone; mo, mouth aperture; mt, mouth tube; ne, neck; or1/2, oral ridges of type 1 and 2; ov, ovary; ser, seminal receptacle; sr2–9, spinoscalids of 2nd to 9th row; sp (and arrowheads), anterior spike; su, anal field spurs; tp, trichoscalid plate; tr1, single trichoscalid; tr2, double trichoscalid; vlp, ventrolateral plate; vp, ventral plate; ws, wave-shaped lateral reinforcement.
Fig 17.
Light micrographs of a paratypic adult female of Scutiloricus hugoi gen. et sp. nov.
(A) Overview of the fully extended specimen, anterior faces up. Note the presence of a large oocyte (oo) and two seminal receptacles (ser). (B) Close up of the oocyte and the two seminal receptacles (arrowheads). (C) Close up of the posterior region of the lorica. Note the linear arrangement of the four flosculi (black double arrowheads) on each dorsolateral plate (dlp) and the two flosculi (white double arrowheads) located medially on the dorsal plate (dp). Abbreviations: cs, clavoscalids; in, introvert; lo, lorica; mc, mouth cone; mt, mouth tube; sp, anterior spike; ss, spinoscalid; vlp, ventrolateral plate; vp, ventral plate; ws, wave-shaped lateral reinforcement.
Fig 18.
Light micrographs of the allotypic adult male of Scutiloricus hugoi gen. et sp. nov.
Anterior faces up in all aspects. (A) Overview of the extended specimen, dorsal view. (B) Anterior region of the body, ventral view. (C) Close-up of the anterior spikes of the dorsal and dorsolateral plates. Note the fenestrated aspect of the anterior spikes (arrowheads). (D) Introvert, neck and anterior region of the abdomen; ventral view. (E) Posterior region of the lorica, dorsal view. Note the putative gonopores (go?) located in the posteriormost region of the dorsolateral plates. (F) Posterior region of the lorica, ventral view. Double-headed arrow points to large pore (gland outlet?) situated in the posterior region of the ventral plate. Double arrowheads indicate an anal field spur. Abbreviations: ac, anal cone; af, anal field; bu, buccal tube; cs, clavoscalid; dlp, dorsolateral plate; dp, dorsal plate; in, introvert; lo, lorica; mc, mouth cone; mt, mouth tube; sp and black arrowheads, anterior spike; sr9, spinoscalids of 9th row; ss spinoscalid; tr, trichoscalid; vlp, ventrolateral plate; vp, ventral plate.
Fig 19.
Schematic diagrams of the distribution of introvert and neck appendages in adult forms of Scutiloricus hugoi gen. et sp. nov.
Note that for the sake of clarity sexual dimorphism of the clavoscalids is not depicted. The shaded area indicates the neck region. (A) Polar diagram. (B) Planar projection. Abbreviations: bp, basal plate; cs, clavoscalid; MD, middorsal line; MV, midventral line; mv bp, midventral pair of basal plates; sr1–9, scalids of 1st to 9th row; ss, spinoscalids; tp, trichoscalid plate; tr1, single trichoscalid; tr2, double trichoscalid.
Fig 20.
Light micrographs of four paratypic postlarvae belonging to genus Nanaloricus.
Anterior faces up in all aspects. All specimens characterized by a dorsal plate divided into three subplates. (A and B) Postlarvae tentatively assigned to Nanaloricus valdemari sp. nov. (C and D) Postlarvae tentatively assigned to Nanaloricus mathildeae sp. nov. Abbreviations: bu, buccal tube; cs, clavoscalids; dp, dorsal plate; fl, flosculi; in, introvert; lo, lorica; mc, mouth cone; mt, mouth tube; sp, anterior spike; ss spinoscalid; tr, trichoscalid.
Table 1.
Comparison of adult characters of the species assigned to the genus Nanaloricus (family Nanaloricidae).
Nanaloricus khaitatus is not included in this table because its original description is probably based on observations of at least three different species.
Table 2.
Comparison of adult (or in certain cases postlarval) morphological characters between the genera assigned to family Nanaloricidae.