Neophyllobius cibyci sp. nov.

urn:lsid:zoobank.org:act: 66ED492F-1C66-4CD7-8FF1-5340094F49A4

Figs 1–4; Table 2

Diagnosis

This species is unique due to a combination of following characters: female with dorsal setae reaching setae immediately behind, femur II with three setae (d, v’ and v”), femur III with two setae (d and v’), setae d on genua I–IV reaching tibiae, short setae (about 1⁄6 of podomere length) on femur I, setae l’ and d on femur I nearly located horizontally at same level, dorsal setae c1 and d1 distinctly longer than distance between setae c1–d1 and d1–e1 respectively, idiosomal setae d1 longest, and setae d and l’ on palpal femur heavily spinose.

Etymology

The specific name incorporates the acronym of the “ Centro de Investigación en Biodiversidad y Conservación” (CIByC) at the “Universidad Autónoma del Estado de Morelos ” in recognition of its work on biological conservation.

Type material

Holotype

MEXICO: ♀, CNAC009229, ex Tillandsia hubertiana at 2.5 meters (m) on Quercus obtusata . Collected from the type locality on 12 Mar. 2014, O. Cortés and R. Paredes coll. (RPL1219).

Paratypes

MEXICO: 1 ♀, CNAC 009230, ex T. schiedeana at 3 m on Q. obtusata (RPL 1220); 1 ♀, CNAC 009231, ex T. schiedeana at 3.05 m on Q. obtusata (RPL 1223); 1 ♀, CNAC 009232, 1 L, CNAC 009243, ex T. hubertiana at 2.1 m on Q. obtusata (RPL 1225); 1 ♀, CNAC 009233, ex T. hubertiana at 2.7 m on Sapium macrocarpum (RPL 1237); 1 ♀, CNAC 009234, ex T. schiedeana at 2.7 m on S. macrocarpum (RPL 1238); 1 ♀, CIUM, ex T. schiedeana at 2.7 m on S. macrocarpum (RPL 1241); 1 ♀, CNAC 009235, ex T. schiedeana at 2.9 m on Q. obtusata, 22 Aug. 2014, S. Gómez and R. Paredes coll. (RPL 1295); 1 ♀, CNAC 009236, ex T. schiedeana at 2.6 m on S. macrocarpum, 22 Aug. 2014, S. Gómez and R. Paredes coll. (RPL 1304); 1 ♀, CNAC 009237, on soil litter of S. macrocarpum, 23 Oct. 2014, R. Reyes and R. Paredes coll. (RPL 1341); 1 ♁, CNAC 009238, ex T. hubertiana at 3.6 m on Q. obtusata (RPL 1222); 1 ♁, CNAC 009239, ex T. hubertiana at 2.7 m on S. macrocarpum, 30 Apr. 2014 (RPL 1276); 1 ♁, CNAC 009240, ex T. schiedeana at 2.4 m on S. macrocarpum, 22 Aug. 2014, S. Gómez and R. Paredes coll. (RPL 1301); 1 PN, CNAC 009241, ex T. hubertiana at 3.4 m on Q. obtusata, 30Apr. 2014 (RPL 1255); 1 L, CNAC 009242, same data as holotype (RPL 1219); 1 L, CNAC 009244, ex T. hubertiana at 2.6 m on S. macrocarpum, 30 Apr. 2014 (RPL 1279). All collected from the type locality, with same data as holotype except where noted.

Type locality

MEXICO: Morelos, Tepoztlán, 1 km S of San Andrés de la Cal, 18.94305° N, 99.11786° W, 1490 m a.s.l.

Description

Female (n = 10) (Figs 1, 3 A–D, 4A–D)

Holotype female (followed in parentheses by range of holotype and nine paratype females).

GNATHOSOMA. 87 (81–98) long and 93 (77–93) wide. Subcapitulum with subcapitular setae m 28 (28–31) and two pairs of adoral setae Or1 11 (9–14) and Or2 9 (8–13); these three pairs are nude and slender, m longest (Fig. 1B); distance m–m 25 (21–25). Chelicerae 38 (30–44) long. Palp, five-segmented with following setal distribution: trochanter without setae; femur with two serrate setae, d 20 (17–20) and l’ 39 (31–39); genu with one long, slender, nude dorsal (d) seta 39 (33–39); tibia with three tactile setae (l’, l” and d) and one claw (sword-like seta); tarsus with two eupathidia (acmζ and sulζ), two simple setae (ba and va) and one small solenidion (ω) (Fig. 1A). Setae elcp present.

IDIOSOMA. Longer than wide, 280 (270–350) long (gnathosoma excluded), 235 (195–320) wide. Cuticle striated, except on coxae and attenuated between setal pairs c1, d1, e1 and f1 (Fig. 1C). Dorsum. With 15 pairs of serrated setae set on small tubercles; all setae are long, d1 longest and h2 shortest; all setal pairs are longer than distance to setal pairs immediately behind. Two pairs of eyes are positioned between setae sci and sce. Length of setae: vi 58 (47–58), ve 50 (42–56), sci 46 (40– 53), sce 46 (42–52), pdx 67 (55–71), c1 80 (66–83), c2 70 (53–70), d1 82 (64–82), d2 52 (43–54), e1 77 (60–77), e2 52 (45–57), f1 67 (54–72), f2 41 (36–43), h1 35 (34–39), h2 34 (29–34). Distances between setae: vi–vi 66 (45–66), ve–ve 96 (70–100), vi–ve 26 (26–32), sci–sci 115 (91–120), sce–sce 145 (125–150), sci–sce 45 (35–45), c1–d1 45 (34–58), d1–d1 16 (10–16), d1–d2 83 (82–94), d1– e1 42 (39–46), ve–sci 22 (15–22), pdx–pdx 23 (13–24), pdx–c1 27 (22–37), c1–c1 16 (11–17), c1–c2 98 (86–100), c2–c2 185 (155–190), e1– e1 14 (11–16), e1–f1 46 (37–59), f1–h1 50 (39–57), e1–e2 66 (56–70), d2– e2 42 (39–45), f1–f1 14 (9–14), f1–f2 66 (46–66), f2–f2 115 (105–125), e2–f2 42 (33–71), h1–h1 10 (6–11), h1–h2 30 (21–30), h2–h2 61 (50–68), f2–h2 33 (29–40). Venter. Coxal setae thinner than dorsal setae. Setal pairs 1a, 3a, 4a, ag, g, ps1, ps2 and ps3 nude and short. Setae 1b, 1c, 2c, 3b, 3c, 4b and 4c serrated. Setae 1c longest and ps1–3 shortest. Setal pairs 3a and 4a located on individual platelets. Genito-anal valves with one pair of genital setae (g). Coxa I grouped with coxa II, and coxa III with IV but not completely fused (Fig. 1D). Length of setae: 1a 26 (25–29), 1b 29 (22–30), 1c 69 (53–69), 2c 54 (33–56), 3a 34 (33–38), 3b 37 (25–40), 3c 49 (34–50), 4a 18 (18–26), 4b 21 (17–22), 4c 30 (22–31), ag 15 (15–19), g 10 (10–15), ps1 12 (10–14), ps2 12 (10–14), ps3 12 (9–12).

LEGS. Slender and long, leg I and IV longest. Lengths (excluding ambulacra): leg I 565 (480–565), leg II 495 (440–500), leg III 475 (465–530), leg IV 550 (510–590). Podomere lengths: femur I 180 (155–180), femur II 145 (125–145), femur III 145 (130–145), femur IV 165 (135–165), tibia I 185 (165–185), tibia II 160 (135–160), tibia III 185 (165–190), tibia IV 195 (180–200), tarsus I 62 (60–70), tarsus II 69 (60–70), tarsus III 71 (59–71), tarsus IV 71 (64–76). Leg setae as indicated in Figs 1 E–F, 3A–D and 4A–D. Tarsus I with five setal pairs: vs serrated, tc, a, p and u slender and nude, u bifurcated at tip. Setae tc longest (Fig. 1F). Setae lengths: v’ on femur I 26 (21–27), v” on femur I 27 (20–30), l’ on femur I 28 (21–30), d on femur I 24 (19–24), dorsal seta on genu I 250 (210–265), dorsal seta on genu II 275 (210–280), dorsal seta on genu III 335 (230–340), dorsal seta on genu IV 360 (215–360), solenidion ω on tarsus I 6 (6), ω on tarsus II 5 (4–5). All tarsi with ambulacrum bearing a pair of claws and an empodium with two rows of tenent hairs. Counts of setae on legs I–IV, indicating additional solenidia in parenthesis: coxae: 3–1–2–2, trochanters 1–1–1–1, femora 4–3–2–2, genua 1(κ)–1(κ)–1–1, tibiae 9(φ)–8(φ)–8(φ)–7(φ), tarsi 10(ω)–10(ω)–8–8.

Male (Figs 2 A–B, 3E–H, 4E–H), range of three paratypes

GNATHOSOMA. 72–81 long and 65–72 wide. Subcapitular setae m 25–26, setae Or1 9, Or2 8–9, distance m–m 17–19. Chelicerae 20–24 long. Length of palpal setae: femur d 18–20 and l’ 33–34, and genu d 35.

IDIOSOMA. 195–210 long (gnathosoma excluded), 170–180 wide. Dorsum (Fig. 2A). Setae shorter than those of females, c1 longest and h1 shortest. Length of setae: vi 35–38, ve 42, sci 38–42, sce 37–38, pdx 42–44, c1 51–53, c2 47, d1 40, d2 40–41, e1 45 –46, e2 34 –35, f1 24–25, f2 30, h1 9–10, h2 22. Distances between setae: vi–vi 40–45, ve–ve 65–69, vi–ve 20–25, sci–sci 80–87, sce–sce 105–112, sci–sce 10–11, c1–d1 39–44, d1–d1 8–10, d1–d2 55–59, d1– e1 30–33, ve–sci 15–17, pdx–pdx 9–10, pdx–c1 25–31, c1–c1 9–10, c1–c2 65–75, c2–c2 124–133, e1– e1 8, e1–f1 32–35, f1–h1 22–25, e1– e2 48–50, d2– e2 26–31, f1–f1 8, f1–f2 35–42, f2–f2 63–66, e2–f2 40–43, h1–h1 8, h1–h2 14–15, h2–h2 34–38, f2–h2 15–16. Venter (Fig. 2B). Agenital setae (ag) absent. Genital seta (g) very close to ps3. Length of setae: 1a 16–18, 1b 20–23, 1c 44–47, 2c 35–37, 3a 17–18, 3b 22–24, 3c 25, 4a 12–13, 4b 12–13, 4c 16–21, ps1 9, ps2 9, ps3 9, g 10.

LEGS. Leg IV longest. Lengths (excluding ambulacra): leg I 465–475, leg II 405–410, leg III 440–450, leg IV 475–485. Podomere lengths: femur I 150–155, femur II 115–125, femur III 120, femur IV 130– 135, tibia I 155, tibia II 125–130, tibia III 155–160, tibia IV 155–160, tarsus I 58–60, tarsus II 58–60, tarsus III 60, tarsus IV 65. Tibia I with two solenidia (φ and φ 2), and all tarsi with one long basal solenidion (ω) (Fig. 4 E–H). Leg setae as indicated in Fig. 3 E–H and 4E–H. Setae lengths: v’ on femur I 21–22, v” on femur I 25–27, l’ on femur I 22, d on femur I 24–25, dorsal seta on genu I 110–125, dorsal seta on genu II 130, dorsal seta on genu III 185–195, dorsal seta on genu IV 215–230, solenidion ω on tarsus I 27–28, ω on tarsus II 30, ω on tarsus III 30, ω on tarsus IV 27–28. Counts of setae on legs I–IV, indicating additional solenidia in parenthesis: coxae: 3–1–2–2, trochanters 1–1–1–1, femora 4–3–2–2, genua 1(κ)–1(κ)–1–1, tibiae 9(φ, φ 2)–8(φ)–8(φ)–7(φ), tarsi 10(ω)–10(ω)–8(ω)–8(ω).

Protonymph (Figs 2 C–D, 3I–L, 4I–L)

GNATHOSOMA. 70 long and 65 wide. Subcapitular setae m 20, setae Or1 5, Or2 6, distance m–m 14. Chelicerae 24 long. Length of palpal setae: femur d 13 and l’ 28, and genu d 27.

IDIOSOMA. 295 long (gnathosoma excluded), 230 wide. Dorsum (Fig. 2C). Setae shorter than those of females, c1 longest. Length of setae: vi 41, ve 40, sci 38, sce 43, pdx 46, c1 60, c2 46, d1 56, d2 39, e1 56, e2 40, f1 50, f2 35, h1 30, h2 22. Distances between setae: vi–vi 48, ve–ve 83, vi–ve 18, sci–sci 97, sce–sce 130, sci–sce 39, c1–d1 42, d1–d1 11, d1–d2 69, d1– e1 38, ve–sci 14, pdx–pdx 17, pdx–c1 26, c1–c1 10, c1–c2 78, c2–c2 150, e1– e1 11, e1–f1 45, f1–h1 38, e1– e2 54, d2– e2 36, f1–f1 11, f1–f2 44, f2–f2 86, e2–f2 42, h1–h1 5, h1–h2 18, h2–h2 35, f2–h2 23. Venter (Fig. 2D). Setal pairs 4b, 4c, ag and g absent. Length of setae: 1a 21, 1b 23, 1c 35, 2c 34, 3a 26, 3b 27, 3c 11, 4a 14, ps1 9, ps2 9, ps3 9.

LEGS. Leg I longest. Lengths (excluding ambulacra): leg I 375, leg II 340, leg III 315, leg IV 325. Podomere lengths: femur I 130, femur II 95, femur III 100, femur IV 100, tibia I 115, tibia II 95, tibia III 115, tibia IV 105, tarsus I 55, tarsus II 55, tarsus III 55, tarsus IV 55. Leg setae as shown in Figs 3 I–L and 4I–L. Setae lengths: v” on femur I 21, l’ on femur I 20, d on femur I 18, dorsal seta on genu I 185, dorsal seta on genu II 205, dorsal seta on genu III 240, dorsal seta on genu IV 210, solenidion ω on tarsus I 5, ω on tarsus II 4. Counts of setae on legs I–IV, indicating additional solenidia in parenthesis: coxae:

3–1–2–0, trochanters 1–1–1–0, femora 3–2–1–1, genua 1(κ)–1(κ)–1–1, tibiae 6(φ)–5(φ)–5(φ)–3(φ), tarsi 9(ω)–9(ω)–7–5.

Larva (Figs 2 E–F, 3M–O, 4M–O), range of three paratypes

GNATHOSOMA. 45–50 long and 60 wide. Subcapitular setae m and palpal femur seta d absent, setae Or1 88, Or2 8. Chelicerae 20–22 long. Length of palpal setae: femur l’ 20–21 and genu d 20.

IDIOSOMA. 165–180 long (gnathosoma excluded), 130–135 wide. Dorsum (Fig. 2E). Setae shorter than those of females, c2 longest. Setal pair pdx absent. Length of setae: vi 28–29, ve 30–34, sci 34, sce 34, c1 35–38, c2 42–43, d1 32–34, d2 28, e1 35 –36, e2 25 –27, f1 30–34, f2 23–24, h1 17–18, h2 15. Distances between setae: vi–vi 30–35, ve–ve 58–60, vi–ve 18–20, sci–sci 65–68, sce–sce 85, sci–sce 28–29, c1–d1 43–46, d1–d1 10, d1–d2 37–38, d1– e1 25–27, ve–sci 10–11, vi–c1 35–41, c1–c1 10, c1–c2 55–60, c2– c2 105–110, e1– e1 10, e1–f1 25–30, f1–h1 22–24, e1– e2 35–39, d2– e2 37–40, f1–f1 9–10, f1–f2 25–26, f2–f2 50–55, e2–f2 30–32, h1–h1 8, h1–h2 9, h2–h2 20–22, f2–h2 22–24. Venter (Fig. 2F). Setal pairs 1c, 2c, 3b, 3c, 4b, 4c, ag and g absent. Length of setae: 1a 20–24, 1b 22–23, 3a 25, ps1 7, ps2 7, ps3 8.

LEGS. Leg I longest. Lengths (excluding ambulacra): leg I 260–265, leg II 240, leg III 250–255. Podomere lengths: femur I 80, femur II 60, femur III 70–75, tibia I 60–65, tibia II 55, tibia III 65–70, tarsus I 45, tarsus II 45, tarsus III 45. Leg setae as indicated in Figs 3 M–O and 4M–O. Setae lengths: v” on femur I 32–34, d on femur I 22, dorsal seta on genu I 43–45, dorsal seta on genu II 33, dorsal seta on genu III 45, solenidion ω on tarsus I 3, ω on tarsus II 3. Counts of setae on legs I–III, indicating additional solenidia in parenthesis: coxae: 2–0–0, trochanters 0–0–0, femora 2–2–1, genua 1(κ)–1(κ)–1, tibiae 3(φ)–3(φ)–3(φ), tarsi 7(ω)–7(ω)–5.

Intraspecific variation

In the case of Neophyllobius cibyci sp. nov., we observed that leg setation teratologies are relatively rare, but include the following: 1) adult specimens with certain setae in different locations; for example, female right femur I with v” distal to d instead of proximal (CNAC009236), female right femur I with l’ positioned before d (CNAC009234), female left femur I with l’ positioned before d (CNAC009233), female tibiae I with seta dm between v’ and v” (CNAC009231 and CNAC009235) instead of distal, female right tibia I with dm between v’ and v” (CNAC009232) instead of distal, and male right tibia I with l’p distal to l”p instead of proximal (CNAC009238); 2) female specimens without setae on left or right legs, for example left femur I without l’ (CIUM), left femur I without v” (CNAC009236), and left tibia I without v” (CNAC009236 and CIUM).

Remarks

Neophyllobius cibyci sp. nov. resembles N. farrieri De Leon, 1958 collected on “Spanish moss” (probably Tillandsia usneoides) in Oaxaca, Mexico. In both species, dorsal setae on genua I–IV are very long, extending beyond the tip of the tarsus, setae on femur I are short (about 1⁄6 of podomere length), and setae l’ and d on femur I are nearly located at the same level. These species can be differentiated by the following characters: in N. cibyci sp. nov., dorsal setae c1 and d1 are distinctly longer than the distance between setae c1–d1 and d1–e1 respectively, setae d1 are the longest of the dorsal setae, and setae d and l’ on palpal femur are heavily spinose. In N. farrieri, the lengths of dorsal setae c1 and d1 are the same as the distance between setae c1–d1 and d1–e1 respectively; setae e1 are the longest of the dorsal setae, and setae d and l’ on palpal femur are weakly spinose.

Neophyllobius cibyci sp. nov. includes at least three postembryonic instars (larva, protonymph and adult). We found only one nymphal instar (protonymph) which resembles the female type proposed by Bolland (2001).

Ecology

Neophyllobius cibyci sp. nov. was the most abundant camerobiid species with 17 specimens, most of which (76.5%) were collected in dry season (March and April). This species is deemed to be an inhabitant of the tree canopy because most specimens (94%) were collected on epiphytic bromeliads at 2.8 m (± 0.4) and only one female specimen (CNAC009237) was found on soil litter. Neophyllobius cibyci sp. nov. was found in low numbers, usually one specimen per sample, except for two samples which had two specimens each. The closely resembling species, N. farrieri, also inhabits epiphytic bromeliads of the genus Tillandsia .

Distribution

This species is only known from San Andrés de la Cal, Morelos, Mexico. The type locality of N. cibyci sp. nov. is about 440 km as the crow flies from the known distribution of N. farrieri (Puenta [sic.] de Nejapa, Oaxaca, Mexico).

Setal homology among Neophyllobius cibyci sp. nov. instars

The collection of different instars provides material for study and suggests the ontogenetic development pattern of gnathosoma, idiosoma and leg chaetotaxy.Although, Grandjean’s idiosomal and leg chaetotaxy systems were previously applied to camerobiid mites (Kethley 1990; Fan & Walter 2011), this is the first study of complete chaetotaxy (on gnathosoma, idiosoma and legs) involving three different instars (larva, protonymph and adult) of a camerobiid mite species. Swift’s (2001) study of leg chaetotaxy of Caligonellidae (Raphignathina: Raphignathoidea), included one larva and one adult female of an undetermined species of Neophyllobius . We found only one nymphal instar (protonymph); for this reason and because the presence of deutonymph is questionable in Neophyllobius, we prefer not to hypothesise the presence of setae in this particular instar (deutonymph).

There are different concepts about the leg chaetotaxy. Fundamental setae are those present when the appendages in question are first formed, that is on legs I–III in the larva or on leg IV in the protonymph. Accessory setae are any which are formed during subsequent moults (Grandjean 1941; Norton 1977; Swift 2001). Setal priority for each podomere refers to a list of setal organs that appear at the beginning of ontogeny (fundamental setae) and have greater priority, or force, than organs than develop later (accessory setae), which suggests that these setal organs are less susceptible to evolutionary regression (Norton 1977; Swift 2001). Setae of equal priority are listed together within parentheses as proposed by Norton (1977).

GNATHOSOMA. Larvae have Or1 and Or2. Setal pattern on palps is consistent across larva, protonymph and adults, except that dorsal seta (d) on femur do not appear until the protonymphal instar; count of setae from palp coxa to tarsus: 1 (elcp), 0, 2, 1, 3 + 1 claw, 4 + ω. Setal pair m on subcapitulum do not appear until the protonymphal instar.

IDIOSOMA. The dorsal chaetotaxy in larvae includes 14 setal pairs: vi, ve, sci, sce, c1, c2, d1, d2, e1, e2, f1, f2, h1 and h2 (Fig. 2E). Setal pair pdx appears in protonymph (Fig. 2C). The ventral chaetotaxy in larvae includes six setal pairs: 1a, 1b, 3a, ps1, ps2 and ps3 (Fig. 2F). Setal pairs 1c, 2c, 3b, 3c and 4a are added in protonymph (Fig. 2D). Setal pairs 4b and 4c are added in adults (Figs 1D, 2B). Setal pair ag is present only in adult females (Fig. 1D).

LEGS (Table 2). Setae v are absent on trochanters I–III in larva (Fig. 3 M–O), and do not appear until the protonymphal instar (Fig. 3 I–K). Seta v on trochanter IV does not appear until adulthood (Fig. 3D, H). Femora I–II of the larva with setae v” and d (Fig. 3 M–N), femur III of the larva (Fig. 3O) and femur IV of the protonymph (Fig. 3L) only with seta d. Seta l’ is added on femur I of the protonymph (Fig. 3I). Setae v’ on femora I–III (Fig. 3 A–C, E–G), and l” on femur IV (Fig. 3D, H) are added in adults. The setal priorities on femora I and II for Neophyllobius cibyci sp. nov. are: (d, v”), l’, v’. Priorities on femora III and IV are: d, l”, v’. Genua I–II in all instars with seta d and κ (Fig. 3 A–B, E–F, I–J, M–N), and genu III only with d (Fig. 3C, G, K, O). Genu IV of the protonymph and adults only with seta d (Fig. 3D, H, L).

Tibiae I–III of larva with setae dm, l’, l” and φ (Fig. 3 M–O). Tibia IV of the protonymph with setae dp, v’m, l” and φ (Fig. 3L); dp and l”p are added to tibiae I–III (Fig. 3 I–K) of the protonymph. Seta v’m is added to tibia I (Fig. 3I) of the protonymph. Setae l’p, and d are added to tibiae I–II of adults (Fig. 3 A–B, E–F). Seta v”m appears only on tibiae I of adults (Fig. 3A, E). Setae l’p and v’m are added to tibia III of adults (Fig. 3C, G). Setae l’, dp, l”p and d are added to tibia IV of adults (Fig. 3D, H). Tibia I of males with φ 2 (Fig. 3E). The setal priorities for tibiae I and II in Neophyllobius cibyci sp. nov., are: (dm, l’, l”), (dp, l”p, v’m), (d, l’p, v”m). Priorities on tibiae III and IV are similar except v”m is not formed.

Tarsi I–II of the larva with vs’, (tc), (p), (u) and ω (Fig. 4 M–N), tarsus III with vs’, (tc), (u) and ω (Fig. 4O). In tarsi I–II of protonymph, setal pair a is added (Fig. 4 I–J). Seta vs’ is lost on tarsi I and III of protonymph but vs” appears (Fig. 4I, K). Tarsus IV of the protonymph with vs’, (tc) and (u) (Fig. 4L). Setae vs’ reappears in tarsi I and III of adults (Fig. 4A, C, E, G). Seta vs” appears on tarsi II and IV (Fig. 4B, D, F, H). Setal pair p is added in tarsus IV of the adults (Fig. 4D, H). Tarsi III–IV of males with ω (Fig. 4 G–H). The setal priorities for tarsi I and II are: (tc’, tc”, u’, u”, vs’, p’, p”), (a’, a”, vs”). Priorities on tarsi III and IV are: (tc’, tc”, u’, u”, vs’), (p’, p”, vs”).