Paronellidae with finely denticulate scales covering dorsum of head and body, and ventral face of furcula; Ant. 4 sometimes annulated, never subdivided in two; labial seta L2 normal, not reduced; eyes 0-8; EOS present; Abd. 2-4 with 2, 3, 3 bothriothricha; manubrium without spines, dens with 1-2 rows of spines; mucro short, with 3-5 more or less evenly spaced teeth.

As currently circumscribed (Thibaud and Najt 1988[1989]), the absence of a subdivision on Ant. 4 in Trogolpahysa is the only character that distinguishes this genus from Troglopedetes.

It is not known if the type species of the genus, Trogolaphysa maya Mills, 1938, has EOS, but the presence of this structure in all species discussed below, including the two troglomorphic forms, suggests it is likely also present in that species.

BELIZE: Toledo District: 29 km WNW of Punta Gorda, Blue Creek Cave, Hokeb Ha entrance, 11.IV.2011, SJT, MES, JJ, CMS, GBH & RS, coll.

Type material:Holotype, female on microscope slide preparation, INHS Collection Number 579, 406; Paratypes: BELIZE: Toledo District: 29 km WNW of Punta Gorda, Blue Creek Cave, Hokeb Ha entrance, 11.IV.2011, (3 in alcohol), SJT, MES, JJ, CMS, GBH & RS, coll.; 37 km WNW of Punta Gorda, cave near Pueblo Creek Cave, 16.IV.2011, (4 in alcohol-one headless), MES, JKK, CMS, GBH & GeC, coll.; 28 km NNW of Punta Gorda, Tiger Cave, 9.IV.2012, (1 on slide, 33 in alcohol), SJT, MES, JJ, CMS, GBH, BKK & GaC, coll.; 28 km NNW of Punta Gorda, Bat Cave, 10.IV.2011, (2 on slides, 29 in alcohol—some in poor condition, one headless), SJT, MES, JJ, CMS & GBH, coll.; 31 km WNW of Punta Gorda, Okebal Ha, 14.IV.2011, (3 on slides, 16 in alcohol), SJT, MES, JJ, CMS, GBH, BKK & RS, coll.

Trogolaphysa giordanoae sp. n. is unique among species with 6–8 eyes in having 5 dorsal head macrochaetae, 3 metathoracic macrochaetae and 4 inner macrochaetae on Abd. 4. Among species with known dorsal chaetotaxy, the new species is most similar to Trogolaphysa riopedrensis, but the two species are easily distinguished by the combination of characters given above and by the presence of a relatively shorter mucro in the new species (Table 2). Additional diagnostic characters distinguishing the new species from all other New World Trogolaphysa with 6–8 eyes and capitate/spatulate tenent hair are presented in Table 2. Among the species described before the introduction of chaetotaxy, the new species is most similar to Trogolaphysa distinguenda (Denis, 1931), but the two species can be separated by the presence of a relatively long mucro with 5 teeth in distinguenda, and a 4-toothed short mucro in Trogolaphysa giordanoae sp. n. Trogolaphysa belizeana is the only other New World Trogolaphysa with 3 metathoracic macrochaetae. However, Trogolaphysa belizeana is a troglobiont (sensu Sket 2008, Culver and Pipan 2009)—blind, with long antennae and modified ungues.

Size. Body length up to 2.1 mm.

Color. Pattern, if any, obscured by green dye present in the alcohol in which specimens were preserved (Fig. 1).

Scale distribution. Scales dark brown, present on Ant. 1-2 and base of Ant. 3, more abundant on dorsal face than on ventral face of segment. Scales absent from ventral tube, legs and dorsal face of manubrium.

Head. Antenna/cephalic diagonal ratio 2.0–2.5 (Fig. 1). Apical bulb of Ant. 4 absent; subapical sensillum capitate (Fig. 2), fully contained in circular depression; guard sensillum absent. Sense organ of Ant. 3 (Fig. 3) with sensilla 1 and 4 acuminate, thin-walled and translucent; sensillum 5 acuminate, dark (light dense), shorter than 1 and 4; sensilla 2–3 wide, leaf-like, resting in shallow grooves. Eyes 6+6 (Fig. 4), chaetotaxy of eyepatch well with 4, sometimes 6 ciliate setae, and 1 seta posterior to eye F. Head dorsally with 5 macrochaetae (A0, A2, M3, Pa5 and Pm3 — Figs 4–5). Series M with 2 setae (M3–4); series S with 5 setae (S1–5); seta M0 seen only in one individual; S0 absent. Prelabral setae serrate (Fig. 6). Labral setae smooth: setae in rows A and C subequal; seta B2 distinctly shorter than setae B0 and B1 (Fig. 7). Distal margin of labrum with 1+1 medial hooks, papillae absent (Fig. 8). Apical and subapical setae of maxillary palp smooth; sublobular plate with 2 seta-like appendages. Lateral process of labial papilla E weakly bent dorsally, barely reaching apex of papilla (Fig. 9). Labial triangle setae as M1M2rEL1–2A1–5 (Fig. 10); r short, stout and sparsely ciliate; L1 inserted close to E and distant from L2 when compared to other entomobryoids (Fig. 11). Postlabium covered by setae and scales, all postlabial setae ciliate, modified setae absent. Columns ICELO with 42221 setae (Fig. 11): column I with posterior seta detached from main group and much longer than anterior setae. Ventral cervical setae usually 8+8.

Body. Mesothoracic hood not developed. Complete dorsal macrochaetae as 32/73/0245+0+9. Mesothorax with 1 anterior (a5) and 6 posterior (p3 complex) macrochaetae arranged as is typical for genus (Fig. 12); microchaetae m2, m4, p1, p2, p4, p5 and p6 present. Inner chaetotaxy of metathorax with 3 macro- and 1 microchaetae (Fig. 13). First abdominal segment with 1 anterior (a6) and 4 posterior setae arranged in a single row (Fig. 14). Second abdominal segment (Fig. 15) inner bothriotrix with 3 fan-shaped setae, one microsensillum and macrochaeta m3; outer bothriotrix with 3 fan-shaped setae and macrochaeta m5; setae a6, m6 and p5 present. Third abdominal segment (Fig. 16) inner bothriotrix complex with 2 fan-shaped setae, 1 sensillum, and macrochaeta m3; external bothriotricha with 7 fan-shaped setae, and macrochaetae am6, pm6 and p6; sensillum d2 present, inserted near pm6. Fourth abdominal segment with 5 inner and 9 outer macrochaetae (Fig. 17): large inner macrochaetae A4, A5, B4, and B5 present; B6 a small macrochaeta; large outer macrochaetae D3, E2, E3, F1, F2, and F3 present; macrochaetae E4, F4 and one other seta probably homologous to Fe4, small. Anterior and medial bothriotricha with 7 and 3 fan-shaped supplementary setae, respectively. Posterior bothriotrix, corresponding to D4, without associated supplementary setae. Posterior setae 19–21+19–21. Intersegmental membrane between Abd. 4–5 with 4–10 lenticular organs (as in Trogolaphysa riopedrensis, Fig. 60).

Legs. Trochanteral organ with up to 36 setae. Metathoracic claw complex as in Fig. 18. Tenent hair weakly spatulate. Smooth posterior setae on metathoracic legs 0.76× as long as unguiculus. Unguis with 4 inner teeth: 1 basal tooth sometimes appearing slightly larger than other, both paired teeth ending near middle of inner edge; proximal unpaired tooth as large as basal paired teeth, ending on distal half of inner edge; distal unpaired tooth smallest of all inner teeth and ending on distal fourth of inner edge. Outer tooth ending on basal quarter of outer ungual edge. Unguiculus lanceolate, with outer margin serrate.

Ventral tube. Anterior face with 3+3 or 4+4 distal macrochaetae; lateral and posterior setae not seen clearly.

Furcula. Dens with 2 rows of ciliate spines: inner row with 35–42 spines; outer row with 25–28 spines. Basal outer spines longest (Figs 19–20). Mucro with 4 short, stout teeth (Fig. 21), ratio mucro length/width of dens tip 1.2–1.8×; basal outer tooth reaches to at least half length of basal inner tooth.

This species is dedicated to Rosanna Giordano, the senior author’s wife, for her years of support and contributions to science.

The species is known only fromBelize

Trogolaphysa giordanoae sp. n. is a guanophile, recorded from entrance, twilight (Fig. 22) and dark zones of caves (6.7, 53.3 & 40.0 % of 15 collections, respectively), often in association with fruit bat or other guano (Fig. 23) (noted for 40% of 15 collections). It was commonly found on the floor of caves (76.9% of 13 collections where position was noted), but also on cave walls (23.1% of 13 collections where position was noted).

BELIZE: Toledo District: 32 km WNW of Punta Gorda, Yok Balum Cave, 13.IV.2012, SJT, MES, JJ, CMS, GBH & AC, coll.

Holotype, female on microscope slide preparation, INHS collection number 579, 407; BELIZE: Toledo District: 32 km WNW of Punta Gorda, Yok Balum Cave, 13.IV.2012, SJT, MES, JJ, CMS, GBH & AC, coll.; Paratypes: BELZE: Toledo District: 32 km WNW of Punta Gorda, Yok Balum Cave, 13.IV.2012, (2 adults & 1 juvenile on slides, 3 adults or subadults & 3 juveniles in alcohol), SJT, MES, JJ, CMS, GBH & AC, coll.; 37 km WNW of Punta Gorda, cave near Pueblo Creek Cave, 16.IV.2011, (1 adult on slide—without legs), MES, JKK, CMS, GBH & GeC, coll.

Trogolaphysa jacobyi sp. n. is the only member of the genus that is blind, has 3-toothed mucro and unguis, and has a single macrochaeta on the metathorax. Trogolaphysa belizeana is the only other New World Trogolaphysa lacking eyes and having a 3-toothed mucro, but it differs from Trogolaphysa jacobyi sp. n. in having 3 metathoracic macrochaetae (1 in Trogolaphysa jacobyi sp. n.), in the arrangement and identity of inner macrochaetae on Abd. 4 (cf. Figs 38, 49 see discussion below), in having few postlabial scales (absent in Trogolaphysa jacobyi sp. n.) and setae (many in Trogolaphysa jacobyi sp. n., cf. Figs 30, 46), in the presence of sensillum d2 on Abd. 3 (absent in Trogolaphysa jacobyi sp. n.), in the absence of unpaired ungual teeth (1 tooth in Trogolaphysa jacobyi sp. n.) and in having a typical lanceolate unguiculus (basally swollen in Trogolaphysa jacobyi sp. n.). Table 3 provides a list of characters that distinguish Trogolaphysa jacobyi sp. n. from all other New World Trogolaphysa lacking eyes and having paired basal ungual teeth inserted near the basal fourth of the inner edge.

Size. Body length up to 2.0 mm.

Color. Living specimens yellowish, with pigment only on a small eyepatch and mesothorax (Fig. 24). Specimens in alcohol white, without trace of pigment.

Scale distribution. Scales transparent, present on Ant. 1–2. Scales absent from postlabial region of head, ventral tube, legs and dorsal face of manubrium.

Head. Antenna/cephalic diagonal ratio up to 5.8 (Fig. 24). Fourth antennomere with incomplete but clear constriction near middle, with many shallow whorls of setae (Fig. 25); apical bulb absent; subapical sensillum not seen. Sense organ of Ant. 3 with sensilla 1 and 4 short, acuminate, thin-walled and translucent; sensillum 5 acuminate, dark and shorter than 1 and 4; sensilla 2–3 broad, leaf-like, resting in shallow grooves. Eyes not seen on slide-mounted specimens, but 1–2 pigment patches visible in life (Fig. 24). Head dorsally with 8 macrochaetae (A0, A2, A3, M3, S3, S5, Pa5 and Pm3 — Fig. 26). Seta M4 displaced laterally towards cephalic sulcus. Series S with setae S1–5; S0 absent, macrochaeta S3 displaced anteriorly, away from cephalic sulcus (cf. Figs 5, 26). Prelabral and all labral setae smooth: setae within row A and C subequal; seta B2 shorter than B0 and B1 (Fig. 27). Distal margin of labrum smooth, papillae absent. Apical and subapical setae of maxillary palp smooth; sublobular plate without seta-like appendages. Lateral process of labial papilla E weakly bent dorsally and not nearly reaching apex of papilla (Fig. 28). Labial triangle setae as M1M2rEL1–2A1–5 (Fig. 29), seta M1 ciliate, all others smooth; r short; A2 close to r, L1 close to E and distant from L2. Postlabium without scales, polychaetotic, uniformly covered with many large and small, weakly ciliate or smooth setae (Fig. 30); modified setae absent. Columns ICELO ill defined due to polychaetosis. Ventral cervical setae usually 6+6.

Body. Mesothoracic hood not developed. Complete dorsal macrochaetae as 62/41/0244+0+9-11. Mesothorax with 1 anterior (a5) and usually 3 posterior (p1–3) macrochaetae forming an arch (Fig. 31); some individuals with only mesothoraxic macrochaeta p2 (Fig. 32); microchaetae m1, m2, m4, p4 and p5 present. Metathorax with 1 macro- and 5 microchaetae (Fig. 33). First abdominal segment seta a6 absent; 4 posterior setae arranged in a single row (Fig. 34–36). Inner bothriotrix complex of Abd. 2 with 3 fan-shaped setae, one microsensillum and macrochaeta m3; outer bothriotrix with 3 fan-shaped setae and macrochaeta m5; setae a6, m6 and p5 present. Inner bothriotrix complex of Abd. 3 with 3 fan-shaped setae, one sensillum and macrochaetae m3; external bothriotrichal complex (Fig. 37) with 6–7 fan-shaped setae, macrochaetae am6, pm6 and p6; sensillum d2 absent. Fourth abdominal segment with 4 inner (Fig. 38) and 9–11 outer (Fig. 39) macrochaetae: inner macrochaetae A5, B4, and B5 large, B6 small; B5 displaced towards A6 instead of B6; (Fig. 38). Outer macrochaetae D3, E2, E3, F1, and F2 large; small outer macrochaetae E4, F3, F4 and 3 others probably belonging to series Fe present. Abd. 4 anterior and medial bothriotricha with 4 and 2 fan-shaped supplementary setae, respectively (Fig. 39). Posterior bothriotrix corresponds to D4, without associated supplementary setae. Posterior setae 6–7+6–7. Intersegmental membrane between Abd. 4–5 with 4–7 lenticular organs.

Legs. Trochanteral organ with up to 25 setae. Claw complex as in Figs 41–42. Tenent hair acuminate, longer on L1 than L3. Smooth posterior setae on metathoraxic legs as long as unguiculus. Unguis with 3 inner teeth: basal teeth small, subequal and ending on basal fourth of inner edge; unpaired tooth distinctly larger than basal teeth, ending near middle of inner ungual edge. Outer tooth absent on all claws; lateral teeth present only on pro- and mesothoracic legs, and ending on basal quarter of outer edge of unguis (Fig. 41). Unguiculus basally swollen, with basal fifth of outer margin weakly serrate.

Anterior face with 2+2 distal macrochaetae; lateral and posterior setae not seen.

Furcula. Dens with 2 rows of finely ciliate spines, number of spines per row unclear on all specimens examined, but inner row with at least 36 spines. Mucro elongate and slender, with 3 teeth, basal inner tooth absent (Fig. 43): ratio mucro length/width of dens tip 2.3–2.8 (mode=2.4).

The species is known only from caves in southern Belize

Trogolaphysa jacobyi sp. n. is a troglobiont (sensu Sket 2008, Culver and Pipan 2009). Living specimens seem to have eye pigment (Fig. 24), but we were unable to identify corneas on specimens examined. The only structure resembling a cornea corresponds to the EOS.

It could be argued that the constriction of the fourth antennomere places this species in Troglopedetes. However, the presence of a well-developed ciliate labial seta L2, the incomplete nature of the constriction on Ant. 4, and the similarity with Trogolaphysa belizeana (presumably with complete, unconstricted Ant. 4, and therefore an uncontested Trogolaphysa) suggest that Trogolaphysa jacobyi sp. n. should be retained in Trogolaphysa. Additionally, the fact that all other Troglopedetes species are restricted to the Old World have prompted us to retain the new species in Trogolaphysa.

This species is dedicated to JoAnn Jacoby, the junior author’s wife, in gratitude for her enthusiasm and assistance in the planning and execution of field-work in the caves of Belize and in many earlier excursions.

This species is a troglobiont, and all 5 collections (11 individuals) were taken in the dark zone (0 lux) on the floor (Fig. 44), often (80% of collections) in wet conditions associated with flowstone or calcite and drip pools, sometimes with scattered cricket droppings.

Two paratypes; Belize: Cayo District, Actun Chapal cave, 7 km SE of Benque Viejo del Carmen, 10.XII.1992, W.R. Elliott.

Additions to the original description.

Head. Dorsal chaetotaxy of the head identical to that of Trogolaphysa jacobyi sp. n., with macrochaetae A0, A2, A3, M3, S3, S5, Pa5 and Pm3. Labral margin smooth. Sublobular plate of outer maxillary lobe without setae-like appendages. Labial papilla E with lateral appendage reaching tip of papilla; 5 proximal smooth labial setae present, seta z (Soto-Adames 2010) longest. Labial triangle formula as M1M2rEL1L2A1-5 (Fig. 45): M1 ciliate, shorter but thicker than M2; r short, stout, apically acuminate; A2 close to r. Postlabium with few scales; columns ICEL with 7732 setae (Fig. 46); seta L2 shortest; ventral cervical setae 6+6.

Body. Dorsal macrochaeta formula as 62/3–43/0343+0+11. Mesothorax with macrochaetae p2, p3 and a5, and microchaetae m4 and p5 clearly visible; setae p1, m2 and p6 obscured. Metathorax with 3 macro- and 1 microchaetae arranged as in Trogolaphysa giordanoae sp. n. (Fig. 13). Abd. 1 with at least three inner microchaetae, apparently without a6, but lateral field of segment not clearly visible. Abd. 2 chaetotaxy normal (Fig. 47): with bothriotricha m2 and a5, sensillum as, macrochaetae m3 and m5, setae a6, m6 and p5, and fan-shaped supplementary setae around bothriotrichal complexes. Abd. 3 (Fig. 48) with insertion of bothriotricha m2, a5 and m5, macrochaetae m3, a7, pm6 and p6, and sensillum d2 normally placed. Chaetotaxy of Abd. 4 as in Fig. 49: inner macrochaetae A4, A5, B5 and B6 present, B6 smallest; outer macrochaetae T7, D3, E2, E3, E4, F1, F2, F3, F4, one member of series Fe and one posterior setae of unclear homology present; relative position of bothriotricha normal; microchaeta B4 absent, microchaeta Te1 present. Posterior setae 7+7. Intersegmental membrane between Abd. 4–5 with at least 4 lenticular organs, actual number of organs unclear due to folding of membrane.

Legs. Claw complex of pro- and metathoracic legs as in Figs 50–51. Tenent hair acuminate. Outer and lateral unguis teeth small, inconspicuous; inner paired teeth with one tooth slightly, but clearly larger, unpaired teeth absent. Unguiculus lanceolate.

Ventral tube. With 2+2 distal macrochaetae on anterior face.

The paratypes examined differ from the original description of the species in having labial seta L2 smooth instead of ciliate, in having only 2 posterior mesothoraxic macrochaetae, in the claws having lateral teeth and in the number of bothriotricha on Abd. 2 and Abd. 4.

Variation in the number of mesothoraxic macrochaetae is also seen in Trogolaphysa jacobyi sp. n and may be related to post-embryonic development. The chaetotaxy of Abd. 2 in fig. 12 of Palacios-Vargas and Thibaud (1997) suggests a composite of the chaetotaxy of Abd. 2 and 3, whereas the bothriotrichal complex of Abd. 4 shown in Palacios-Vargas and Thibaud (1997, fig. 13) seems based on an aberrant specimen.

Puerto Rico: Isabela, Guajataca Commonwealth Forest, Rd. 446, 18.41263°N, 66.96887°W, top of mogote at crossroad between trails 6, 25 & 26, leaf litter, 15.V.2009, F. Soto (2 specimens); Cayey, Rd. 4471, Km 4.1, leaf litter, 18.VI.1998, F. Soto (1 specimen); Mayagüez, University of Puerto Rico, secondary forest east of Biology Building, 18.21350°N, 67.13774°W, royal palm (Roystonea borinquena O.F. Cook) leaf litter, III.2009, M. Ospina (3 specimens).

Additions to the original description.

Head. Dorsal chaetotaxy as in Fig. 52: macrochaetae A0, A2, M3, S3, S5, Pa5 and Pm3 present; 1+1 microchaetae inserted near A1. Postlabium with all setae ciliate; columns ICELO with 41232; posterior setae on column I detached from anterior group.

Body. Mesothorax (Fig. 53) with one anterior (a5) and six posterior macrochaeta; microchaetae m2, m4, p5 and p6 present; microchaetae p1 and p2 absent. Metathorax with 4 inner microchaetae, as in Trogolaphysa riopedrensis (Fig. 58). Abd. 1 with 4 posterior setae; seta a6 absent. Abd. 2 and 3 as in Trogolaphysa giordanoae sp. n. (Figs 15, 16); Abd. 2 seta p5 fusiform, with enlarged socket (Fig. 54). Abd. 4 as in Fig. 55: inner macrochaetae A4, A5, B5 and B6 present; macrochaetae Te2, D3, E2, E3, F1–3 present; 4 other lateral and posterior small macrochaetae present. Posterior setae 13–14+13–14. Intersegmental membrane between Abd. 4–5 with 4 lenticular organs.

Ventral tube. Anterior face with 3+3 distal macrochaetae; smaller individuals with 2+2 macrochaetae.

Puerto Rico: Maricao, Maricao Commonwealth Forest, near observation tower on Rd. 120, 18.14444°N, 66.97962°W, leaf litter, 8.VI.1998, F. Soto (1 specimen); Mayagüez, University of Puerto Rico, secondary forest east of Biology Building, 18.21350°N, 67.13774°W, royal palm (Roystonea borinquena) leaf litter, III.2009, M. Ospina (3 specimens).

Additions to the original description.

Head. Dorsal chaetotaxy as in Fig. 56: macrochaetae A0, A2, M3, S3, Pa5 and Pm3 present. Postlabium with all setae ciliate; columns ICELO with 41232; posterior setae on column I detached from anterior group.

Body. Mesothorax as in Trogolaphysa jataca (Fig. 53). Metathorax as in Trogolaphysa riopedrensis (Fig. 58). Abd. 1 as in Trogolaphysa riopedrensis (Fig. 59) with one anterior (a6) and 4 posterior setae. Abd. 2 and 3 as in Trogolaphysa giordanoae sp. n. (Figs 15, 16); Abd. 2 seta p5 fusiform as in Trogolaphysa jataca. Abd. 4 as in Trogolaphysa jataca (Fig. 55): inner macrochaetae A4, A5, B5 and B6 present; macrochaetae Te2, D3, E2, E3, F1, F2, F3 present; 4 other lateral and posterior small macrochaetae present. Posterior setae 13–14+13–14. Intersegmental membrane between Abd. 4–5 with 4–6 lenticular organs.

Ventral tube. Anterior face with 3+3 distal macrochaetae.

Puerto Rico, Aguadilla, Caimital Alto, Villa Grajales, 18.44058°N, 67.11840°W, moist mown lawn, 9.VII.1999, F. Soto (1 specimen); USA Virgin Islands, St. Thomas, 18.35348°N, 64.93520°W, wet leaf litter, patch of forest along Rd. 33 near intersection with Rd. 40, 28.VI.2000, F. Soto (1 specimen).

Additions to the original description.

Head. Dorsal chaetotaxy as in Fig. 57: macrochaetae A0, (A2), A3, S3, S5, Pa5 and Pm3 present. Postlabium with all setae ciliate; columns ICELO with 41232; posterior setae on column C detached from anterior group.

Body. Mesothorax as in Trogolaphysa jataca (Fig. 53). Metathorax as in (Fig. 58). Abd. 1 with 1 anterior (a6) and 4 posterior setae (Fig. 59). Abd. 2 and 3 as in Trogolaphysa giordanoae sp. n. (Figs 15, 16). Abd. 4 as in Fig. 60: inner macrochaetae A4, A5, B4, B5 and B6 present; outer macrochaetae D3, E2, E3, F1, F2, F3, Fe3 present; at least one other outer macrochaeta present. Posterior setae 17+17. Intersegmental membrane between Abd. 4–5 with 4–6 lenticular organs (Fig. 60).

Ventral tube. Anterior face with 4+4 distal macrochaetae.

Remarks: The individual from St. Thomas lacks head macrochaetae A2. In the individual from Aguadilla the dorsal and outer teeth of the unguis end on the basal fourth of the claw instead of the distal half.