taxonID	type	description	language	source
038106260C6C717118C0FD90FAA54326.taxon	description	Figs 1 – 6	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C6C717118C0FD90FAA54326.taxon	materials_examined	Material examined. UKRAINE: SIZK K- 2369 (jointly with SIZK K- 2363 – 2368), SIZK K- 27632 and SIZK K- 2880. All from Rovno amber, Rovno Oblast (Region), Sarny District, Klesov, Pugach quarry; late Eocene. Syninclusions: SIZK K- 27632, stellate hairs / trichomes, Acari; SIZK K- 2363, female of Chironomidae (Orthocladiinae); SIZK K- 2364, two Collembola, Entomobryomorpha; SIZK K- 2365, female of Chironomidae (Orthocladiinae); SIZK K- 2366, Oribatida: Platyliodes ensigerus Sellnick; SIZK K- 2367, two Collembola, Entomobryomorpha; SIZK K- 2368, two Collembola, Entomobryomorpha.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C6C717118C0FD90FAA54326.taxon	description	Description. Within the darker resin, the body of SIZK K- 2369 (Fig. 1) is deformed and covered dorso-laterally with an air bubble. During processing of the amber, the body of the harvestman (L: 0.825) was sawn off ventro-laterally. This fossil is female, as shown by the presence of three large thorns (acute tubercles / megaspines) ventrally on the femur of the pedipalp. Femur distally with a large mesal setose lobe / apophysis. All pedipalpal segments armed with stout setae, mostly on mesal surfaces. Claw large: 0.084. We could measure correctly only the length of palpal segments (Fe: 0.7, Pa: 0.32 – 0.35, Ti: 0.28 – 0.3, Ta: 0.56), and the length of some of the leg femurs (Fe I: 1.08, Fe II: 0.77). SIZK K- 27632 (Fig. 2) has a body dorsally obscured by the presence of breaks / cracks in the amber just above it. This specimen (L: 1.25) is also a female. A protruding ovipositor (furcal length: 0.126 – 0.14) (Fig. 2: arrow) is visible, as well as the presence of the three typical femoral megaspines. Ocular tubercle: length: 0.5, width: 1.0; diameter of each lens 0.175. Appendage lengths: chelicera, basal segment equivocal, distal segment 0.52, movable digit 0.17. We could correctly measure only the length of the following palpal segments: Cx: 0.3, Tr: 0.2, Ti: 0.22; Ta: 0.475; length of the tarsal claw: 0.07. Legs femur length: Fe II: 1.12, Fe III: 0.63; other femurs could not be measured accurately. SIZK K- 2880 (Figs 3 – 6) is not easily visible dorsally due to the presence of an air-pocket, as well as many cracks in the amber (Fig. 3). Ventrally, the body is covered with a white emulsion, as well as with filamentous fungal hyphae (Figs 4 – 5). It is noteworthy that a similar arthropod-pathogenic fungal overgrowth is usually observed on dead harvestmen in very humid conditions (Cokendolpher & Mitov 2007: 348 – 349, fig. 9.2; Wijnhoven 2009: 13 – 14, fig. 25; Barbosa et al. 2016: 13, fig. 2 A – B; P. Mitov, pers. obs.). Three large thorns (acute tubercles) (Fig. 6) are clearly visible on the femur of the pedipalp, indicating that this specimen is again female (see Gruber 1974; Shear 1975). Its body has L: 1.37. The ocular tubercle (Fig. 3) is characteristic of the species — compared to the length of the body it is large, with length: 0.56, and width: 0.87, and diameter of each lens 0.238. The length of only a part of the palpal segments could be measured accurately: Fe: 0.35, Pa: 0.35, Ti: 0.28, Ta: 0.35; Legs femur length: Fe I: 0.76, Fe II: 1.12, Fe III: 0.74, Fe IV: 0.91.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C6C717118C0FD90FAA54326.taxon	discussion	Remarks. Caddo dentipalpus is easy to recognise among the European amber harvestmen through its combination of a very large and broad eye tubercle, very large eyes and large thorns (acute tubercles / megspines) on the palpal femur. It has also been recorded from Baltic and Bitterfeld amber (Table 1) and as discussed by Shear (1975) and Dunlop & Mitov (2009) it is of particular interest for being almost identical to a living species, Caddo agilis Banks, restricted today to North America and East Asia (Suzuki 1976). No species of Caddo, or even the family Caddidae, occur in the Recent European fauna and the Rovno records further indicate that Caddo was originally more widely distributed across Eastern Europe as well. Its modern relatives (Caddo agilis, C. pepperella Shear) are usually found in very humid, densely shaded areas, such as ravines, and there is usually a coniferous element in the forest (Shear 1975), or it can be found in bamboo thickets and beech forests (Suzuki 1976). Caddo agilis tends to occupy exposed surfaces (e. g., tree trunks, logs, stones) and C. pepperella occurs on the ground in the leaf litter and under fallen objects (Suzuki 1976; Shultz & Regier 2009). Caddo dentipalpus is found quite often in the European ambers. Perhaps the fossil species also preferred tree trunks, where it would have easily come into contact with sticky resin.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FEB8FC5346C6.taxon	description	Figs 7 – 8	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FEB8FC5346C6.taxon	materials_examined	Material examined. UKRAINE: SIZK K- 1915 (jointly with SIZK K- 1914, and SIZK K- 1916 – 1921). All from Rovno amber, Rovno Oblast (Region), Sarny District, Klesov, Pugach quarry; late Eocene. Syninclusions: SIZK K- 1914, Acari, two Isopoda: Oniscoidea; SIZK K- 1915, stellate hairs / trichomes; Collembola, Entomobryomorpha; Diptera: Chironomidae: Orthocladiinae;? Sciaridae; SIZK K- 1916, two Collembola, Entomobryomorpha; SIZK K- 1917: Collembola, Entomobryomorpha; SIZK K- 1918, Collembola, Entomobryomorpha; Sciaridae; SIZK K- 1919, Collembola, Symphypleona; Collembola, Entomobryomorpha, spider web; SIZK K- 1920, worker ant Ctenobethylus goepperti (Mayr); SIZK K- 1921, three Acari (one Oribatida); Collembola, Entomobryomorpha; worker ant Ctenobethylus goepperti (Mayr); Diptera: Sciaridae and Chironomidae.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FEB8FC5346C6.taxon	description	Description. SIZK K- 1915 has only its ventral side clearly visible (Fig. 7), as a larger air pocket hides the posterior abdomen ventro-laterally. Additionally, numerous air bubbles scattered throughout the resin obscure the view dorsally. However, the short apophyses (bearing a number of stout spines) typical for this fossil species on the inner (mesal) surface of the palpal patella are clearly visible (Fig. 8: black arrow); the tibia is slightly inflated distally (Fig. 8: white arrow) and also bears stout mesal spines in this distal region (for details see Dunlop & Mitov 2009). The specimen measures L: 1.625; abdomen width: 0.925; appendage lengths: palpal Ta: 0.42, the remaining palpal segments cannot be measured accurately; leg segments: Fe I: 2.25, Pa I: 0.25, Ti I: 2.75; Fe II: 2.75; Tr III: 0.12, Fe III: 1.25, Pa III: 0.25, Ti III: 1.0; Tr IV: 0.12, Fe IV: 2.0, Pa IV: 0.25, Ti IV: 1.72; the remaining segments could not be measured.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FEB8FC5346C6.taxon	discussion	Remarks. Amilenus deltshevi Dunlop & Mitov was previously known from Bitterfeld and Baltic amber (Dunlop & Mitov 2009; Elsaka et al. 2019). The present specimen is the first record from Rovno amber and another example of a harvestman species common to all three faunas (Table 1).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FB24FB1C45DE.taxon	type_taxon	Type species. Phalangium propinquum Lucas [= Metaphalangium cirtanum (C. L. Koch)] (Roewer 1911; Staręga 1984), by original designation.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C61717F18C0FB24FB1C45DE.taxon	discussion	Remarks. The new fossil specimens described here show similarities to representatives of the extant genera Phalangium Linnaeus, Dasylobus Simon, and Metaphalangium Roewer; mainly the presence of a process on the palpal patella, the shape, armament and position of the ocular tubercle, the slender legs and its light color (in amber) with a silver shine. These new fossils are probably juveniles or immatures due to their small size and the underdeveloped armament on the ocular tubercle. Therefore, we compared these specimens with the juveniles of some extant species of the above-mentioned genera and found that the new fossil species is most similar to juveniles of the genus Metaphalangium, and especially to representatives of M. cirtanum (Fig. 12), which is also the most widespread extant species of the genus. The similarity (see Figs 9 – 14) between these is expressed in the position and structure of the ocular tubercle, the presence of a medio-distal prominence of the palpal patella, the proportions of the pedipalpal segments, the slender legs, and especially in the presence of stout spine-like setae on the medial surface of the palpal patella and tibia (Figs 11 – 12, 14). Metaphalangium Roewer so far comprises about 13 recent species, distributed in the Mediterranean region, the Balkans, the Near East, and the Caucasus. Only M. albounilineatum (Algeria, Libya, Morocco, Tunisia, and southern Italy) and M. cirtanum (circum-Mediterranean region) have a wider range (Mkheidze 1952; Martens 1978; Crawford 1992; Staręga 1984, 2004; Prieto 2003, 2008; Özdikmen et al. 2010; Barrientos et al. 2017; Kury et al. 2020). The present discovery in Rovno amber is a new genus record for the fossil harvestman fauna.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	description	Figs 9 – 11, 13 – 14	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	materials_examined	Type material. UKRAINE: Holotype. SIZK K- 3197, Rovno region, Sarny District, Klesov, Pugach quarry; late Eocene. Syninclusions: SIZK K- 3197, stellate hairs / trichomes; Diptera: Rhagionidae. Paratype: SIZK K- 1254, with same data as for holotype, but syninclusions absent. Both from Rovno amber.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	etymology	Etymology. In honour of Prof. Jochen Martens (Mainz, Germany) in recognition of his extensive contributions to arachnology / opilionology, and on the occasion of his 80 th birthday; name in the genitive case.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	diagnosis	Diagnosis. Easily distinguished from all other fossil phalangiid species by the unique combination of: (1) palpal patella with a small and blunt medio-distal prominence, (2) palpal patella and tibia mesially covered with not densely spaced stout spine-like setae, typically arranged brush-like in 4 – 5 rows, (3) absence of a distal apophysis on the palpal tibia, (4) the ocular tubercle (ellipsoidal, normal in size) is up to twice its length from the frontal margin of the carapace, and (5) having thin and relatively long legs.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	description	Description. The holotype, SIZK K- 3197, is an almost complete specimen (Figs 9 – 10), possibly hinting at a juvenile / immature, best seen in dorsal view, but missing the ventral surface of the body (together with the leg coxae, and parts of the pedipalpal coxae and femora). Body, partly dorsally hidden in a gas bubble, and by a white emulsion anterior to the ocular tubercle. Body ovoid, light colored (colouration in amber – pale gray-yellow) with a silverish shine; length ca. 1.75; width of prosoma 1.5; width of opisthosoma 1.65. Ocular tubercle broader than long: length 0.175, and width 0.30, with 2 – 3 tubercles on the eye rings; diameter of each lens 0.15. Distance from ocular tubercle to front of prosoma 0.325. Ocular tubercle up to two times of its length from the frontal margin of the carapace. Pedipalps (with sawn-off coxae and femurs; the latter have only their distal parts preserved, with a length of 0.25), with article lengths of: Pa: 0.2; Ti: 0.32; Ta: 0.65; the claw, 0.07. On the preserved distal parts of the femurs there are spine-like setae apically and ventrally, ordered roughly in two rows. Palpal patella with a small and blunt medio-distal prominence; palpal patella and tibia mesially covered with not densely spaced stout spine-like setae, specifically brush-like, ordered in 4 – 5 rows (Fig. 11: arrows). These spine-like setae are 0.05 – 0.07 high and the distance between them is 0.03 – 0.04. On the medial surface of the patella they number 16 – 18, while on the medial side of the tibia they are about 26 – 28. Similar spine-like setae are scattered on the surface of the palpal tarsus as well, along with thinner setae; tarsus ends in a claw (Fig. 11). Legs relatively complete (Figs 9 – 10), all elongate and slender, and when preserved at their full length ending in a single claw. Leg article length: Leg I (Fe: 1.85, Pa: 0.5, Ti: 1.6, Mt: 1.75, Ta: 2.85 – 3.0); Leg II (Fe: 3.7, Pa: 0.5 – 0.6, Ti: 3.5, Mt: 3.25, Ta: 7.75); Leg III (Fe: 1.9, Pa: 0.35; full length of tibia, metatarsus, and tarsus not preserved [i. e., sawn-off]); Leg IV (Fe: 2.75, Pa: 0.55, Ti: 2.0, Mt: 2.85; full length of tarsus not preserved [partially sawn-off]). Index of leg length (IL) = 1.23. The leg segments are cylindrical; femurs with longitudinal rows of acute inclined tubercles; thinnest are the femurs of the second pair of legs, thickest those of the first and third pairs. Their patellae bear dorsally and ventrally sparse setae and dorso-distally two tubercles each. Tibia distally slightly dorso-ventrally flattened, with sparse setae laterally and with more setae on the ventral side. Metatarsus with few setae on all sides. Tarsi with short, fine and numerous setae on their ventral sides. The paratype, SIZK K- 1254, is enclosed in very dark amber (Fig. 13). During processing of the amber piece, the ventral body surface was sawn off with parts of the legs. Body length 1.62; prosoma width 1.25, abdomen width 1.25, possibly also hinting at a juvenile / immature. The body dorsally, the ocular tubercle, the chelicerae, the pedipalps and parts of the legs are covered with a white emulsion. Therefore, only part of the pedipalpal and leg segments can be seen and measured. Pedipalps (Fig. 14), with article lengths of: Fe: 0.35 – 0.4, Pa: 0.21, Ti: 0.37, Ta: 0.62. Palpal femurs dorso-apically and ventrally with stout spine-like setae, palpal patella and tibia medially also with stout spine-like setae. Palpal patella and tibia dorsally with setae; palpal tarsus with many setae; tarsus ends with a claw. Palps covered with a white emulsion, thus features of the setae appear much thicker than they would have been in life – compare with the holotype (Fig. 11). Leg article length: Leg I (Fe: ca. 1.5, Pa: 0.37, Ti: 1.25, Mt: 1.75, Ta: 1.125); Leg II (Fe: 2.75, Pa: 0.37); Leg III (Fe: 1.3, Pa: 0.36); Leg IV: femur is sawn off, the rest of it 2.0 mm long. Index of leg length (IL) = 1.2. The remaining leg segments are sawn off or cannot be measured correctly.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C63717918C0FF40FD9F405A.taxon	discussion	Remarks. The key characters of this new fossil Metaphalangium species are the small mesal projection on the palpal patella, in combination with the characteristically arranged stout spine-like setae along the entire mesal surface of palpal patella and tibia (Figs 11, 14). However, these stout spine-like setae (in the holotype at least; Fig. 11) are much thicker than those in extant forms; see the juvenile Metaphalangium cirtanum (Fig. 12). When apophyses are present in phalangiid harvestmen, they can be more prominent in immature instars and become less well expressed in adults (Mitov et al. 2015: p. 41; PGM, pers. obs.) and representatives of Metaphalangium are no exception. In extant members of this genus only the adults (both sexes) of Metaphalangium corsicum (Roewer) and Metaphalangium lusitanicum (Roewer) have short palpal apophyses. Of the studied representatives of the genus (see Material and methods), in adults of M. cirtanum (females) and M. strandi (both sexes), only the palpal patella bears a mesially pronounced rounded protrusion at the distal end, while in juvenile M. cirtanum the pedipalpal patellae bear clearly formed apophyses (Fig. 12). Conversely, in immature specimens of Metaphalangium martensi sp. n. such apophyses are missing — the palpal patella bears only a small and blunt medio-distal prominence (Fig. 11: black arrow). Differences can also be seen in the morphology of the legs: in both immature fossil specimens they are somewhat more slender than in extant juvenile Metaphalangium, and the index of leg length (IL) in the smallest studied juvenile M. cirtanum (with L: 3.3) is 1.06., while in M. martensi sp. n. (with L: 1.62 – 1.75), IL varies between 1.2 and 1.23.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C67716718C0FCB1FBA443EE.taxon	materials_examined	Material examined. UKRAINE: SIZK K- 1151 (jointly with SIZK K- 1150, and SIZK K- 1152 – 1158), SIZK K- 9767. All from Rovno amber, Rovno Oblast (Region), Sarny District, Klesov, Pugach quarry; late Eocene. Syninclusions: SIZK K- 1150, female Theridiidae (Araneae); Auchenorrhyncha (Cicadinea); SIZK K- 9767: stellate hairs / trichomes; SIZK K- 9768: Сollembola, Entomobryomorpha; SIZK K- 9769, incertae sedis; SIZK K- 1151, Diptera: Dolichopodidae; SIZK K- 1152: Cecydomyiidae (Lеdomyia sp.); SIZK K- 11533, Araneae, juv.; SIZK K- 1154, Acari: Cunaxidae (Cunaxinae) (Sergeenko et al. 2013); SIZK K- 1155, Auchenorrhyncha: Fulgoroidea; Diptera: Nematocera; SIZK K- 1156, Hymenoptera: Scelioninae; SIZK K- 1157, Diptera: Tipuloidea; SIZK K- 1158: female of Culicoides (Ceratopogonidae) (Sontag & Szadziewski 2011).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C67716718C0FCB1FBA443EE.taxon	description	Description. SIZK K- 1151 (Figs 15 – 16) is near the edge of the amber piece: part of the opisthosoma is concealed by an air bubble. During the processing of the amber piece, the front part of the prosoma was sawn off together with part of the limbs. However, the specific shape of the ocular tubercle and the structure of the remaining patellar and tibial apophyses of the pedipalps (Fig. 16) clearly indicate the generic affiliation of this fossil. Body length: 1.125, abdomen width: 0.87. Ocular tubercle width: 0.25, diameter of each lens 0.11. Appendage lengths: palpal tarsus length: 0.52, tarsus ends with claw; tibial apophysis length: 0.1; tibial apophysis maximum width: 0.07. The other palpal / limbs segments cannot be measured correctly. SIZK K- 9767 (Fig. 17) with L: 1.12 – 1.25; abdomen width: 0.88; Appendage lengths: Palps with article lengths of: Fe: 0.56; Pa: 0.18; patellar apophysis length: 0.28; Ti: 0.42 – 0.43; tibial apophysis length: 0.1; Ta: 0.77 – 0.78. Legs femur length: Fe I: 2.15 – 2.25; Fe II: 4.85; Fe III: 2.35; Fe IV: 3.35.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C67716718C0FCB1FBA443EE.taxon	discussion	Remarks. In extant forms of this genus, the tibial apophysis / protuberances of the pedipalps are usually more prominent in juveniles and females than in males. An apophysis is present on the pedipalpal tibia in these fossil specimens too (Figs 16 – 17), see also Dunlop & Mitov (2009: fig. 19). This combined with their small size indicates that, as in extant Dicranopalpus, these are most likely either juveniles (Staręga 2002) — see also photos of juvenile Dicranopalpus in Duinen (2020) — or subadults (most likely female) of relatively small-bodied species (body length in the range of 1.1 – 3.6). Comparative modern examples would be, e. g., Dicranopalpus angolensis (Lawrence) from Angola in Africa, or D. larvatus (Canestrini) and D. brevipes Marcellino, both from Italy (Lawrence, 1951: 40 – 42, figs 3 – 4; Wijnhoven & Martens 2019: fig. 4; Wijnhoven et al. 2020: fig. 4). Dicranopalpus ramiger has been previously recorded from Baltic and Bitterfeld amber (Table 1) and was the only harvestman species previously reported from Rovno amber (Perkovsky et al. 2010 a). Interestingly, its modern relative Dicranopalpus ramosus Doleschall appears to be in the process of moving northwards from its Mediterranean origins (Wijnhoven & Prieto 2015, and literature therein), and is thus reclaiming areas where the fossil record indicates the genus formerly occurred. Noordijk et al. (2007) noted that the living species D. ramosus is primarily scansorial on trees and shrubs. If the amber species had a similar lifestyle to the modern counterpart this might explain the frequency with which these animals tended to become trapped in resin. In northern European amber collections it is usually the predominant harvestman species (see Mitov et al. 2021).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716718C0FD70FD5647F5.taxon	description	Figs 18 – 21	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716718C0FD70FD5647F5.taxon	materials_examined	Material examined. UKRAINE: SIZK K- 7898, from Rovno amber, Rovno Oblast (Region), Sarny District, Klesov, Pugach quarry; late Eocene. Syninclusions: stellate hairs / trichomes; Isopoda (Oniscoidea).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716718C0FD70FD5647F5.taxon	description	Description. SIZK K- 7898 (Figs 18 – 19) has a deformed body (L: 1.5), prosoma width: 1.0. Carapace smooth, with only a few setae. Opisthosomal dorsum ornamented by regular rows of black thorns (Fig. 19). Venter: surface of all coxae, genital operculum and sternites with a black spine-like setae; ocular tubercle: length 0.25, and width 0.375. Appendage article lengths: chelicerae: basal segment: 0.25, distal segment: 0.67, movable digit: 0.3; pedipalps: Fe: 0.62, Pa: 0.57, Ti: 0.92, Ta: 0.37; Leg I: Fe: 1.0, Pa: 0.5, Ti: 1.12, Mt: 1.45, Ta: 1.55; Leg II: Fe: 1.7, Pa: 0.62, Ti: 1.97, Mt: 2.32, Ta: 3.2; Leg III: Fe: 1.0, Pa: 0.42, Ti: 1.0, Mt: 1.62; Leg IV: Fe: 0.92, Pa: 0.45; part of the segments could not be measured correctly. Pedipalps inflated (Figs 20 – 21), trochanter, femur and patella with scattered spine-like setae; tibia and tarsus throughout with dense brush-like setae — probably plumose glandular as in modern species, see Wolff et al. (2016) — ventro-distally, tibia with a long seta (Fig. 20: arrow). All leg articles with black, scattered spine-like setae; patellae, tibiae, metatarsi and tarsi with short setae; tarsi ending with a single claw.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716718C0FD70FD5647F5.taxon	discussion	Remarks. The Eocene localities fit into the present range of Sabacon, namely SW Europe, Siberia, Russia (Primorye Province), the Himalayas, India, China, Japan, Korea and North America (see Staręga 2002; Martens 2015). It is noteworthy that the shape of the palpal tibia and tarsus in Sabacon claviger (Figs 20 – 21) is closest to that of males of European members of the genus (i. e., Sabacon viscayanum ramblaianum Martens and Sabacon altomontanum Martens; e. g., Martens 1983: figs 32, 48).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716518C0F9FBFD46415E.taxon	diagnosis	Diagnosis. Small, dark fossil nemastomatids (body length range 1.25 – 1.52, body width ca. 1.0) with ovoid to quadrangular body, dorsally with fine granulation, only anterior corners of scutum magnum with more prominent granules, and with four pairs of slender, pillar-like spines, each slightly bulbous at the top, standing perpendicular to the body surface. In males, apophyses resembling an ‘ ice-cream scoop’ (or in dorso / dorso-lateral outline, boxing gloves) present on first cheliceral articles; apophysis approx. low, from 0.11 to ca. 0.2 mm. Pedipalps and legs elongate and slender. Pedipalps without apophyses (spurs / thorns / denticles). Palpal tarsus relatively short, in lateral view machete-like – distally rounded and slightly widened; in ventral view – like a baseball bat. Palpal Ta / Ti between 0.44 and 0.55 mm. Microsculpture elements of the leg femurs include loose-textured denticles and granules.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716518C0F9FBFD46415E.taxon	type_taxon	Type species. Nemastoma tuberculatum C. L. Koch & Berendt, 1854, by original designation.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716518C0F9FBFD46415E.taxon	etymology	Etymology. From the Greek ‘ para’ (beside, near) and the modern harvestman genus Histricostoma which this material closely resembles. Gender neuter.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C79716518C0F9FBFD46415E.taxon	discussion	Remarks. The extinct genus Parahistricostoma is morphologically closest to the recent genus Histricostoma Kratochvíl, which can be found today from the Alps through to the Balkans, Turkey and the Caucasus, and includes eight species (Kratochvíl 1958; Staręga 1976 b; Martens 1978, 2006; Snegovaya & Marusik 2012; Schönhofer 2013; Iorio & Delfoss 2015). The traditional assignment of Koch and Berendt’s (1854) amber species Nemastoma tuberculatum to the genus Histricostoma was initially based on the presence of false articulations on the leg femora (see Staręga, 2002: 602), and later largely based on its dorsal ornament (Dunlop 2006; Dunlop & Mitov 2009). As in Histricostoma, four pairs of slender, pillar-like spines, each slightly bulbous at the top (“ Stabdorne ” sensu Martens 1978) are observed. The distal end of the pedipalp femur in both the fossils and living Histricostoma is similar, and the leg trochanters are spinulate. However, the question remains do these features alone justify placing the fossil in the modern genus? We could identify at least four subtle differences between the Histricostoma - like fossils and modern Histricostoma species. First, the form of the cheliceral apophysis in the fossils differs from that seen in recent species. Second, there is no evidence for a medial spur (thorn / denticle) on the distal end of the male palpal patella, as is seen in all modern Histricostoma. Third, the legs and pedipalps of the fossils are somewhat longer and more slender than the appendages observed in living species. Finally, the microsculptural elements of the leg femora are also different compared to those in the type species of Histricostoma (H. drenskii), as well to those of H. creticum and H. dentipalpe. We feel that this character combination does merit a separate genus, diagnosed as above. Furthermore, lacking any data on the structure of the penis in this fossil species, we cannot resolve its generic relationships through comparing its genital morphology with that of extant species.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	description	Figs 22 – 24	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	materials_examined	Holotype. MB. A. 1612; ex Berendt collection, repository number 7248. Baltic amber; Palaeogene (Eocene); precise locality unclear.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	distribution	Distribution. Baltic and Bitterfeld amber (Staręga 2002; Dunlop 2006; Dunlop & Mitov 2009; Elsaka et al. 2019).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	diagnosis	Diagnosis: As for the genus.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	materials_examined	Material examined. UKRAINE: SIZK K- 25396, SIZK K- 7460. Rovno region, Sarny District, Klesov, Pugach quarry; late Eocene; (Figs 22 – 24). Syninclusions: SIZK K- 25396, decaying plant remnants; insect larva; SIZK K- 7460, two Acari (one Oribatida).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	description	Description. SIZK K- 25396 is a male specimen, body and legs covered by a white emulsion dorsally; the prosoma and part of the opisthosoma obscured by an air-pocket. The fossil is clearly visible only ventrally (Fig. 22). Body and limbs (in amber) black. L: 1.25, abdomen width: 0.88. The armament of the scutum and the dorsal granulation / microsculpture of the body as well as the ocular tubercle are not visible. Only the tips of three of the four pairs of dorsal spines are visible. Chelicerae: basal segment equivocal, length of distal segment: 0.43, movable digit length: 0.17; basal article distally with apophysis; the cheliceral apophyses are shaped like an ‘ ice cream scoop’ (Fig. 23), directed slightly downwards and resemble boxing gloves in dorso-lateral outline; see also Dunlop & Mitov (2009). The length / height of the apophyses is 0.126. The cheliceral apophyses and the frontal distal segment of the chelicerae bear strong setae. One of the setae, located distally at the apex of the apophysis, is twice as long (ca. 0.04 mm) as the others. The opening of the cheliceral glands / secretion area is probably located on the medial depressions of the apophysis (Fig. 23: white arrow). Cheliceral basal segment laterally spinulate; Palps with article lengths of: Tr: 0.25, Fe: 0.84, Pa: 0.77, Ti: 0.53, Ta: 0.29; as Ta: Ti = 0.55. Pedipalps elongate and slender, without apophyses (spurs / thorns / denticles). Palpal trochanters more cylindrical than oval. Palpal coxa and trochanter ventrally covered with denticles (Fig. 23). Distal end of the patella slightly curved downwards. Palpal tibia slender, baseball bat-shaped – basally dilated; palpal tarsus slender, club-shaped in dorsal / ventral view, laterally machete-shaped – distally rounded and slightly dilated. Palpal articles (except trochanters) ventral with clavate setae (L: 0.042). Legs long and slender. Femur length (pseudoarticulations of leg femora in parentheses): Fe I: 1.33 (with three pseudoarticulations, middle); second pair of legs missing; Fe III: 1.44 (with 3 pseudoarticulations, central); Fe IV: 1.33 (with 5 pseudoarticulations, centro-basal). Trochanters of the legs with spinule-like microsculptural elements (Fig. 23: black arrow), similar to those in Histricostoma creticum and H. drenskii. Microsculpture elements of the leg femurs include loose-textured denticles and granules. Metatarsal and tarsal leg segments densely setose. Legs with 10 – 16 tarsal segments, ending with a single claw. Description. K- 7460 is a male specimen, ventrally covered by a white emulsion (Fig. 24), dorsally not well visible due to fractures in the amber and due to an air-pocket adjacent to the legs and body. Four pairs of slender, pillar-like spines, each slightly bulbous at the top, can be seen. The dorsal granulation / microsculpture of the body is not visible. Body and limb color (in amber) black. Body length: 1.3 – 1.4, abdomen width: 0.80. Ocular tubercle: length: 0.15 and width: 0.25; diameter of each lens: 0.06; Chelicerae: basal article distally with apophysis. Pedipalps elongate and slender, without apophyses. Palpal trochanter more cylindrical than oval. Palpal tibia slender, baseball bat-shaped – basally dilated; palpal tarsus slender, baseball bat-shaped in ventral view (Fig. 24). Palps with article lengths of: Ti: 0.69 – 0.73, Ta: 0.38; Ta: Ti = 0.52 – 0.55; the remaining palpal segments cannot be measured correctly. The distal end of the patella curves slightly downwards. Palpal segments (except trochanters) with clavate setae (L: 0.042) ventrad. Legs long and slender. Segments of Leg I with lengths of: Fe: 1.58, Pa: 0.32, Ti: 0.85; Leg III with article lengths of: Fe: 1.71, Pa: 0.32, Ti: 0.80; tarsus ends in a single claw. The pseudoarticulations and the microsculpture of the leg femora are obscured. The leg trochanters are spinulate – their microsculpture is similar to that in Histricostoma creticum and H. drenskii.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7B716318C0FC5DFAF94139.taxon	discussion	Remarks. For additional morphological data see Dunlop & Mitov (2009: 351 – 355, figs 1 – 2, 6 – 8).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7D716318C0FC79FB5F47FA.taxon	description	Figs 25 – 27	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7D716318C0FC79FB5F47FA.taxon	materials_examined	Material examined .. UKRAINE: SIZK UA- 1565, Rovno amber, precise locality unclear; late Eocene. Syninclusions: none.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7D716318C0FC79FB5F47FA.taxon	description	Description. A juvenile covered by a white emulsion concealing the dorsal granulation and the armature of the scutum (Fig. 25). With L: 0.50; prosoma width: 0.27; abdomen width: 0.28; ocular tubercle elliptic (Fig. 26), with length: 0.06 and width: 0.08; diameter of each lens 0.03; Palps with article lengths: Fe: 0.308; Pa: 0.238; Ti: 0.238; Ta: 0.14. Palpal articles (except trochanters) ventrally with clavate setae (L: 0.056) (see Figs 26 – 27). The tarsus of each leg is one-segmented (Fig. 25) with a length of: Ta I: 0.28; Ta II. – IV.: 0.25, respectively.	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
038106260C7D716318C0FC79FB5F47FA.taxon	discussion	Remarks. Given the shape of the ocular tubercle, the proportions of the pedipalps, and the shape of their segments, this juvenile most probably belongs to Parahistricostoma tuberculatum, too. The length of the body, the single tarsal segments and their length, indicate that this juvenile nemastomatid, by analogy with other extant nemastomatid species, could be the first or the second juvenile stage (cf. Immel 1954: figs 8, 10).	en	Mitov, Plamen G., Perkovsky, Evgeny E., Dunlop, Jason A. (2021): Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine). Zootaxa 4984 (1): 43-72, DOI: https://doi.org/10.11646/zootaxa.4984.1.6
