Arrhopalites karabiensis, Vargovitsh, Robert S., 2009

Vargovitsh, Robert S., 2009, New Cave Arrhopalitidae (Collembola: Symphypleona) from the Crimea (Ukraine), Zootaxa 2047, pp. 1-47 : 10-20

Arrhopalites karabiensis sp. nov.

Figs 2 –33, Table 2

Diagnosis. Unpigmented; 1 + 1 eyes; trichobothria ABC form right angle and AB=2BC; Ant/head = 1.5–1.6; Ant III somewhat swollen; Ant IV with 5 (pseudo)subsegments usually indistinctly separated from each other by 2 or more annuli; 13 spine-like setae on head; all claws with inner tooth and all empodia with corner tooth, empodial filaments overtopping tip of corresponding claw; tenaculum with 1 setula; manubrium with 5 + 5 setae; dens with 3, 2, 1, 1, 1 anterior setae and anterodistal seta as a strong spine; Abd VI with broadened circumanal setae, rod-like apically and laterally serrated appendices anales, and with 4 + 4 or less cuticular spines.

Material examined. Holotype on slide (C-408-1): female, Ukraine, Crimea, Karabi Massif, Bol'shoy Buzluk Cave, 5.viii.2003. Paratypes on slides: female, collected together with holotype; male, 32 females, Karabi Massif, Kastere Cave, 4.viii.1998. Holotype and 31 paratypes are preserved in SIZNAS, 2 paratypes in SNHML.

Other material (on slides). Karabi Massif: male, 8 females, Kastere Cave, 12.viii.1993; 12 males, 11 females, 4 juv., 200 Let Simferopolya Cave, 5.08.1998; 6 females, juv., same cave, 8.vii.1994; female, Soldatskaya Cave, 8.viii.1993; male, Gvozdetskogo Cave, 9.viii.2003; male, 2 females, Mamontovaya Cave, 15.viii.2003; male, 3 females, Nakhimovskaya Cave, 14.viii.2003. Besides slides, part of material is preserved in alcohol.

Description. Female: body 1–1.2 mm length, without pigmentation ( Fig. 2 ).

Head ( Fig. 3 ): eyes 1 + 1, unpigmented. Labral setal pattern: a: 4, m: 5, p: 5; prelabral: 6. Clypeal area: 6 rows (from a to f); row a with axial seta. Interantennal area: 2 rows (α and β) with axial seta in row β. Dorsal area: 4 rows (from A to D) with 3 axial setae in rows A, B and C. 13 setae of head dorsum are spine-like ( Fig. 14 ) (in row A: 1 + 1; B: 1 + 1; C: 2 + axial + 2; D: 2 + 2) and 4 setae are not modified (in row A: 1 + axial + 1; in row B: axial).

Antennae: 1.5–1.6 times as long as head. Length ratio of antennal segments I: II: III: IV = 1: 2–2.2: 2.9–3.2: 5.9–6.4. Antennal segment I with 7 slightly spiny setae, subapical posterior one is very small; Ant II with 14 slightly spiny and simple setae ( Fig. 20 ). Ant III ( Fig. 21 ): generally swollen in subbasal and median part; with 18 setae and 2-rods sense organ; setae Api and Ape shorter and thinner than others, seta Aai very small and blunt, seta Ae curved and lies on one level with seta Ape, sense rods and seta Aa. Ant IV ( Fig. 22 ) indistinctly subdivided into 5 subsegments or pseudosubsegments, often separated each from other by weakly developed annuli. (Pseudo)subsegmental formula: 1 + 3 + 1 = (A + M1–M2) + (M3–M5) + (B). Ant IV bears 13 whorls of setae: 5 on apical subsegment (AI–AIII, M1–M2), 3 on median subsegments and 5 on basal subsegment (BA, BM1–BM3, BB).

Foreleg: precoxae 1, 2 and coxa with 1, 0, 1 setae respectively ( Fig. 4 ). Trochanter with 3 anterior and 1 posterior setae; femur with 12 setae, a4 turned perpendicularly to the longitudinal axis of the segment (Fig. 25). Tibiotarsus (Fig. 26): with 3 setae FP (FPe, FPae, FPpe); secondary seta FS missing; whorl I with 9 setae among which Ja curved and thickened; each of whorls II–V with 8 setae, interior setae thicker than others. Pretarsus with 1 anterior and 1 posterior setulae. Claw (Fig. 26): with inner tooth and 2 pairs of lateral teeth visible in dorsoventral position (Fig. 31); without tunica. Empodium (Fig. 26): thin; with corner tooth in subbasal half; with long apical filament exceeding tip of claw. Claw 4.2–4.6 times shorter than tibiotarsus.

Mid leg: precoxae 1, 2 and coxa with 1, 1, 3 setae respectively ( Fig. 4 ). Trochanter with 3 simple setae and anterior trochanteral organ; femur with 13 setae, 2 posterior ones are very small (Fig. 28). Tibiotarsus (Fig. 27): 3 setae FP present, seta FS missing; whorl I with 9 setae, whorls II–IV with 8 setae, whorl V with 7 setae (Vp absent). Claw (Fig. 27): broader than in foreleg; with inner tooth and 2 pairs of lateral teeth; in some specimens tunica was observed. Empodium (Fig. 27): broader than in foreleg; with corner tooth in subbasal part; apical filament exceeding tip of claw. Claw 4.3–4.9 times shorter than tibiotarsus.

Hind leg: precoxae and coxa with 1, 1, 3 setae ( Fig. 4 ); trochanter with anterior trochanteral organ, 3 anterior and 1 posterior simple setae; femur with 12 setae, among them 2 very small posterior setulae (Fig.

Arrhopalites karabiensis : 25–26, chaetotaxy of foreleg, anterior view: trochanter and femur (25), tibiotarsus and foot complex (26); 27–28, chaetotaxy of mid leg, anterior view: tibiotarsus and foot complex (27), trochanter and femur (28); 29–30, chaetotaxy of hind leg, anterior view: tibiotarsus and foot complex (29), trochanter and femur (30); 31, fore claw with lateral teeth, dorsal view; 32, chaetotaxy of furca, lateral view; 33, chaetotaxy of dens with missing seta (IIIpi), posterior view.

30). Tibiotarsus (Fig. 29): 3 setae FP, seta FSa present; whorls I–V as in mid tibiotarsus. Claw (Fig. 29): broader than in fore and mid leg; with inner tooth and 2 pairs of lateral teeth; tunica usually developed. Empodium (Fig. 29): the broadest; with corner tooth in the middle; apical filament exceeding tip of claw. Claw 5.4–6 times shorter than tibiotarsus.

Lengths ratio of tibiotarsi I: II: III = 1: 1: 1.3.

Ventral tube with 1 + 1 subapical curved setulae. Tenaculum ( Fig. 17 ): each ramus with 3 sclerotic teeth and a soft basal process; anterior lobe with 1 apical setula; tip of posterior lobe not exceeding tip of anterior lobe.

Furca (Fig. 32): manubrium with 5 + 5 posterior setae. Dens: anterior side with 5 groups of thick setae (3, 2, 1, 1, 1); Ia as heavy spine ( Fig. 12 ); posterior side with 9 spines or spiny setae (Ie, Ipe–Vpe, IIIpi, IVpi, Ii) and 7 simple setae (Ip–IVp, Bpe, Bp, Bpi) ( Figs 13 , 32; Table 2 ). Mucro: constricted on anterior side; edges gutter-like serrated; tip globular ( Figs 18 , 32). Dens 1.6–1.7 times as long as mucro.

Great abdomen ( Fig. 4 ): meso- and metathorax with thickened or slightly spiny dorsal setae ( Fig. 11 ). Trichobothria ABC form an angle about 90o and AB distinctly longer than BC so that AB=2BC. Single p seta of p-row of Abd I is located above the level of trichobothrium B (marked with arrow); seta b1 lies far behind the line BC; seta c1 lies above and seta c2 – below trichobothrium C. Posterior lateral complex with 2 + 3 and furca base complex with 8 setae. Posterior dorsal complex with about 30–35 setae distinctly shorter than hind claw ( Figs 8, 9 ), among them many extra setae, so that each of rows dI and dII consists of 10–11 setae and, in general, 3 rows arrangement is hardly seen. Ventral complex with 3 setae.

Fifth abdominal segment with two setae and trichobothrium D in row a, and 2 setae in row p ( Fig. 4 ).

Sixth abdominal segment ( Fig. 23 ): with maximum 4 + 4 cuticular spines; circumanal setae broadened, often with subbasal serration ( Figs 5, 6 ); setae ms2 and as2 much shorter than seta ms1 and comparable with seta ps1. Appendices anales (mi5): rod-like, gradually broadening to distal part, with apical and subapical serration ( Figs 15–16 ).

Male: 0.8 mm length. Antennae 1.7–1.9 times as long as head. Ant I: II: III: IV = 1: 2.3–2.6: 3–3.3: 6.8–7.9. Ant III less broadened than in female. Ant IV with 5–8 subsegments, usually with distinct and more numerous annuli between subsegments than in female ( Fig. 19 ). Ratio ‘tibiotarsus: claw’ of foreleg/ mid leg/ hind leg = 4.5–5/ 4.7–4.9/ 5.6–6.4. Dens 1.5–1.6 times as long as mucro. Chaetotaxy of posterior-external margin of dens (pe) often incomplete. Small abdomen without cuticular spines and broadened setae, and with less setae than in female ( Fig. 24 ).

Variability. Dens normally as in Fig. 32, Table 2 , but abnormal chaetotaxy (missing or (and) shifted setae) also occurs. Among examined specimens several variations of dens chaetotaxy have been noted: (1) one of spiny setae IIIpi or (and) IVpi missing (Fig. 33) and remained seta sometimes shifted on intermediate position between normal IIIpi and IVpi positions; (2) posterior-external row (pe) in males often incomplete or (and) with shifted positions of setae ( Table 2 ): only less than 20% of examined specimens with complete 6 setae in row pe (as normal in females); (3) two females and one male show normal anterior chaetotaxy on right dens and incomplete (3, 2, 1...1) on left dens. Incomplete number of cuticular spines (less than 4 + 4 up to absence) on female small abdomen is often observed. In some females circumanal seta mps2 on dorsal anal valve asymmetrically forked.

Bionomy and distribution. All specimens were collected from water surface, in dark zone of caves at a vertical depth of more than - 70 m from entrance. Findings of this species exclusively in caves of only one karstic massif inhabited by common troglobitic speleofauna indicate that this species is rather troglobiont. A. karabiensis sp. nov. occurs in association with other species of Arrhopalitidae ( Table 1 ). Single juvenile specimens similar to described species where also found in Inzhenernaya Cave of Chatyr-Dag Massif and Paniya Cave of Aj-Petri Massif but their belonging to A. karabiensis sp. nov. is doubtful.

Etymology. The new species is named after type locality—Karabi Jajla.

Remarks. A. karabiensis sp. nov. is very close to the widespread A. caecus ( Tullberg, 1871) but a little larger than it and differs from it by: long apical filament on empodia overtoping tip of claw; more or less developed annulations between subsegments of Ant IV; generally swollen Ant III.

From A. ulehlovae Rusek, 1970 from Czech the new species differs by: antenna/head ratio (1.3 in A. ulehlovae ); 5 subsegments on Ant IV (3–4 in A. ulehlovae ); presence of cuticular spines on female small abdomen.

From the troglophile A. anulifer Nayrolles, 1990 from Thailand the described species differs by: anterior dens chaetotaxy (formula: 3, 2, 2, 1, 1 in A. anulifer ) and presence of distinct anterodistal spine; number and shape of cuticular spines on anal valves (5 + 5 in A. anulifer ); 1 apical setula on tenaculum (2 in A. anulifer ).

Chaetotaxy of furca, tenaculum, tibiotarsi and presence of annulations between subsegments of Ant IV resemble A. peculiaris sp. nov. (described below), but body proportions, shape of foot complex, chaetotaxy of head and small abdomen distinctly separate these species.

Species Characters e ae a ai i pi p pe

A. whorls I s + S + s - + s

karabiensis I–V II - s s - - - + s (-) Diagnosis. Unpigmented; 1 + 1 eyes; trichobothria ABC form right angle and AB equidistant with BC; Ant/ head = 2–2.4; Ant IV with 6–7 subsegments (or pseudosubsegments) separated from each other by more or less developed annulations; 5 spine-like setae on head; claws thin, empodia shorter than claws; tenaculum with 1 setula; manubrium with 5 + 5 setae; dens with 3, 2, 1, 1, 1 anterior setae and anterodistal seta as a weak spine; small abdomen without any cuticular spines, with not broadened circumanal setae, and with rod-like apically and laterally weakly serrated appendices anales.

Material examined. Holotype on slide (C-271-2): female, Ukraine, Crimea, Aj-Petri Massif, Geophysicheskaya Cave, 16.viii.1998. Paratypes on slides: 4 females, collected together with holotype. Types are preserved in SIZNAS.

Other material (on slides). Aj-Petri Massif: female, juv., Skelskaya Cave, 14.viii.1998; 3 males, 5 females, 10 juv., branch of Aj-Petri Massif, Great Canyon of Crimea, Paniya Cave, 2.viii.2005.

Description. Female: body 1–1.2 mm length, without pigmentation ( Fig. 34 ).

Head ( Fig. 35 ): 1 + 1 very small eyes, often not observed, unpigmented. Clypeal area with axial setae in row a (in holotype also in row b). Interantennal area with axial seta in row β. Dorsal area with 3(2) axial setae in rows A, B and C. 5 setae of dorsal area ( Fig. 45 ) are spine-like and relatively long (in row A: 1 + 1; B: 1 + 1; C: axial); others are not modified.

Antennae: 2.2–2.4 times as long as head. Antennal segments I: II: III: IV = 1: 2–2.3: 3–3.2: 7–7.7. Ant I with 6 anterior and 1 very small posterior setae; Ant II with 14 setae, one of which in subbasal part of the segment is much longer than others ( Fig. 50 ). Ant III ( Fig. 51 ) with 18 setae; Ant III organ includes 2 sense rods located in separate pits, and one of them is situated a little higher than another; setae Api and Ape short and thin; seta Aai very short and rod-like; seta Ae somewhat curved and lies on one level with Ape and sense rods; seta Aa situated distantly under sense rods level. Ant IV subdivided into 6–7 subsegments, separated each from other by more or less developed annulations ( Fig. 52 ). Subsegmental formula: 1 + 4/5 + 1 = (A + M1, M2) + (M3–M6/M7) + (M7/0 + B). Basal subsegment of Ant IV is subequal to Ant III or a little longer than it if Ant IV has 6 subsegments (1: 1–1.1) and a little shorter than Ant III if Ant IV has 7 subsegments (1: 0.85–0.89). Ratio basal subsegment: apical subsegment = 1.8–2 (if 6 subsegments present) or 1.5–1.6 (if 7 subsegments present). Ant IV with 15 whorls of setae: 5 on apical subsegment, 4(5) on median subsegments and 6(5) on basal subsegment. Whorls M1 and M2 seeming homological to other whorls of median subsegments, but with one additional seta in basal part of whorl ( Fig. 52 ).

Foreleg: coxa with 1 seta ( Fig. 36 ). Trochanter with 3 anterior and 1 posterior setae; femur with 11–12 setae, a4 turned perpendicularly to the longitudinal axis of the segment ( Fig. 55 ). Tibiotarsus ( Fig. 56 ): 3 setae FP present; seta FS missing; 9 setae in whorl I among which seta Ja curved and thickened; 8 setae in each of whorls II–V. Claw slender, without tunica, with very small inner tooth or without it; empodium narrow, with minute corner tooth, not reaching tip of claw ( Fig. 56 ). Claw 4.1–4.82 times shorter than tibiotarsus.

Mid leg: precoxae and coxa with 1, 1, 3 setae respectively ( Fig. 36 ). Trochanter with anterior trochanteral organ, 2 anterior and 1 posterior simple setae; femur with 13 setae ( Fig. 58 ). Tibiotarsus ( Fig. 57 ): 3 setae FP present; seta FS missing; 9 setae in whorl I; 8 setae in each of whorls II–IV; 7 setae in whorl V. Claw similar to fore claw but a little shorter; empodium broadened in basis and inclinated in the middle, with very small inner tooth or without it, not reaching tip of claw ( Fig. 57 ). Claw 4.36–4.96 times shorter than tibiotarsus.

Hind leg: precoxae and coxa with 1, 1, 3 setae respectively ( Fig. 36 ). Trochanter with anterior trochanteral organ, 3 anterior and 1 posterior simple setae; femur with 12 setae among which 2 posterior ones are very small ( Fig. 59 ). Tibiotarsal chaetotaxy as in mid tibiotarsus but secondary seta FSa present ( Fig. 60 ). Claw broader and shorter than fore and mid claw, with pair of small lateral teeth and very small up to absence inner tooth; empodium broader and longer than in fore and mid leg, not exceeding claws tip, with small up to absence corner tooth ( Fig. 60 ). Claw 5.6–6.5 times shorter than tibiotarsus.

Lengths ratio of tibiotarsi I: II: III = 1: 1: 1.2.

Ventral tube with 1 + 1 subapical setulae. Tenaculum ( Fig. 48 ): ramus 3-dentate, with basal appendage; anterior lobe with 1 apical setula; tip of posterior lobe not reaching tip of anterior lobe.

Furca ( Figs 61, 62 ): manubrium with 5 + 5 posterior setae; dens chaetotaxy resembles A. caecus ( Table 2 ). Anterior side bears heavy setae: 3, 2, 1, 1, 1; Ia looks rather like heavy seta than spine ( Fig. 44 ). Dens 1.9–2 times as long as mucro. Mucro ( Figs 49 , 61, 62 ) with serrated edges and globular apex.

Great abdomen ( Fig. 36 ): dorsal setae of mesothorax short and not modified, comparable with other anterior setae ( Figs 41, 42 ). Trichobothria ABC form an angle about 90o and AB subequal with BC (AB≥BC). Single p seta of p-row of Abd I is located above the level of trichobothrium B (marked with arrow); seta b1 lies far behind the line BC; seta c1 lies above and seta c2 – below trichobothrium C. Posterior lateral complex with 4 and furca base complex with 8 setae. Posterior dorsal complex with 3 longitudinal rows of long setae, 3.5 times as anterior ones, 1.4 times as hind claw and longer than circumanal setae. Ventral complex with 2 setae.

Fifth abdominal segment with trichobothrium D and 4 setae ( Fig. 36 ).

Sixth abdominal segment ( Fig. 53 ): no broadened setae; cuticular spines absent; circumanal row is reduced (unique for genus): setae mps3, mpi1 and ms5 absent in type specimens. Appendices anales ( Figs 46, 47 ): rod-like, with weak lateral and apical serration, 1.6–2 times shorter than hind claw.

Male (from Paniya Cave, not from the type locality): 0.8 mm length. Antennae about 2 times as long as head. Ant I: II: III: IV = 1: 2.5: 3.4: 8.1. Ant IV very indistinctly or not subsegmentated (as well as in females from Paniya Cave). Ratio ‘tibiotarsus: claw’ of foreleg/ mid leg/ hind leg = 4/ 4/ 5. Dens 1.8–1.9 times as long as mucro. Apex or mucro not globular, only slightly swelled. Chaetotaxy of small abdomen as in Fig. 54 .

Variability. Dorsal + antennal area of head with 4 or 3 axial setae. Ant IV with 6 or 7 subsegments. Femur of foreleg with 11 or 12 setae. Inner tooth of claws and corner tooth of empodia very small or absent. Two specimens from Geophysicheskaya Cave and three from Paniya Cave show normal anterior chaetotaxy (3, 2, 1, 1, 1) on one dens ( Fig. 61 ) and peculiar (3, 2, 1, 1, 1, 1) on another dens ( Fig. 62 ). Mucro usually with more or less globular apex, but one specimen (0.9 mm) without developed swelling was observed. The same specimen shows unusual appendices anales without distinct basal papilla and curved in distal part.

Specimens from Paniya Cave differ from others by: very indistinct or unnoticeable subsegmentation of Ant IV; 5 thickened setae on head dorsum instead of distinct spines; distinct corner tooth on empodia, stronger and complete row of circumanal setae of small abdomen. It is possible that population from caves of Aj-Petri and population from the Paniya Cave of Great Canyon of Crimea form different subspecies or even separate species but additional material from more localities must be studied.

Bionomy and distribution. The specimens were collected in caves mainly from water surface, also from humid stalagmite and wooden substance. This species seems to be rare troglobiont distributed only in the caves of Western biospeleological region of Crimea (Aj-Petri Massif and its branches. In different caves it occurs together with Pygmarrhopaliltes tauricus sp. nov. and P. principalis skelicus ssp. nov. described below. Population of Paniya Cave looks like allopatric variance of the species or even separate species.

Etymology. Latin " peculiaris " means peculiar, specific—this name shows unusual and rare for the genus combination of features and peculiar tendency to bear 6 rows of setae on anterior dens along with normal 5- row pattern.

Remarks. A. peculiaris sp. nov. resembles many characteristics of A. karabiensis described above, but they are different in several features concerning trichobothrial pattern, head chaetotaxy, antennae, foot complex and small abdomen.

A. gul Yosii, 1966 b from South Korean caves as well as the new species is characterized by long antennae with annulations on Ant IV, thin and long claws, 5 rows of setae on anterior dens, and absence of spines on small abdomen, but such characters as presence of broadened and serrated circumanal setae on small abdomen, presence of only 1 spiny seta on posterior dens are different from those of the new species. Presence of 5 spiny setae on head dorsum is also specific feature.

Figures

FIGURES 2 – 18. Arrhopalites karabiensis sp. nov.: 2, outline of habitus; 3, chaetotaxy of head, frontal view; 4, chaetotaxy of great abdomen, lateral view; 5 – 16, shape of setae: (mpi 1) of Abd VI (5), (mps 2) of Abd VI (6), (ms 1) of Abd VI (7), posterior dorsal seta of great abdomen (8), (dI- 1) of great abdomen (9), anterior dorsal seta of abdomen (10), dorsal seta of mesothorax (11), spine-like (Ia) seta of dens (12), spine-like (Ipe) seta of dens (13), spine-like seta of head vertex (14), appendices anales, lateral view (15) and dorsal view (16); 17, tenaculum; 18, apex of mucro.

FIGURES 19 – 24. Arrhopalites karabiensis sp. nov.: 19, outline of antenna of male; 20 – 22, chaetotaxy of antennal segments, anterior view: Ant I – II (20), Ant III (21), Ant IV (22); 23 – 24, chaetotaxy of Abd VI: female (23), male (24).

FIGURES 34 – 49. Arrhopalites peculiaris sp. nov.: 34, outline of habitus; 35, chaetotaxy of head, frontal view; 36, chaetotaxy of great abdomen of juv. specimen, lateral view; 37 – 47, shape of setae: (ms 1) of Abd VI (37), (mps 2) of Abd VI (38), (dI- 1) of great abdomen (39), posterior dorsal seta of great abdomen (40), anterior dorsal seta of abdomen (41), dorsal seta of mesothorax (42), (Ie) of dens (43), (Ia) of dens (44), spine-like seta of head vertex (45), appendices anales, dorsal view (46) and lateral view (47); 48, tenaculum; 49, apex of mucro.

FIGURES 50 – 54. Arrhopalites peculiaris sp. nov.: 50 – 52, chaetotaxy of antennal segments, anterior view: Ant I – II (50), Ant III (51), Ant IV, posterior view (52); 53 – 54, chaetotaxy of Abd VI: female (53), male (54).

FIGURES 55 – 62. Arrhopalites peculiaris sp. nov.: 55 – 56, chaetotaxy of foreleg, anterior view: trochanter and femur (55), tibiotarsus and foot complex (56); 57 - 58, chaetotaxy of mid leg, anterior view: tibiotarsus and foot complex (57), trochanter and femur (58); 59 – 60, chaetotaxy of hind leg: trochanter and femur, anterior view (59), tibiotarsus and foot complex, posterior view (60); 61, chaetotaxy of dens and mucro, posterior view; 62, chaetotaxy of furca, lateral view: dens with abnormal (3, 2, 1, 1, 1, 1) set of anterior setae.

Tables

TABLE 2. Dens chaetotaxy of Arrhopalites karabiensis sp. nov. and A. peculiaris sp. nov.; + = simple seta; S = spinelike; s = weakly spine-like; v. p. = varying position; in brackets = variability.

III - IV -	- -	s s (-)	- -	- -	s (-; v.p.) s (-; v.p.)	+ +	s (-; v.p) s (-; v.p)
V -	-	-	-	-	-	-	s (-; v.p)
whorl B - A. peculiaris whorls I s I–V II -	- + s	+ s s	- + -	- s -	+ - -	+ + +	+ s s
III - IV -	- -	s s	- -	- -	s s	+ +	s +
V -	-	- (s)	-	-	-	-	+
whorl B -	-	s	-	-	+	+	+
Arrhopalites peculiaris sp. nov. Figs 34–62, Table 2