Taxonomic review of the Neotropical genus Neopachylus ( Arachnida , Opiliones , Gonyleptidae )

A taxonomic review of the genus Neopachylus Roewer, 1913 together with keys to the species for both males and females are presented. Gephyropachylus marginatus Mello-Leitão, 1931 is considered a junior subjective synonym of Neopachylus serrinha Soares & Soares, 1947, and Huralvius incertus Mello-Leitão, 1935 is considered a synonym of Neopachylus nebulosus (Mello-Leitão, 1936). This genus is restricted to southern Brazil, occurring in states of Paraná, Santa Catarina and Rio Grande do Sul.

Neopachylus was proposed by roewer (1913) to include N. bellicosus Roewer, 1913.The same author in 1915, described N. mamillosus Roewer, 1915. Later, Mello-leitão (1931) described three species in monotypic genera: Gephyropachylus marginatus Mello-Leitão, 1931 (in Tricommatinae), Huralvius incertus Mello-Leitão, 1935 and Nunduavius nebulosus Mello-Leitão, 1936 (both in Pachylinae).The descriptions of G. marginatus and H. incertus, were based on females only.SoareS & SoareS (1945) described N. herteli Soares & Soares, 1945, and two years later they described N. serrinha Soares &Soares, 1947 andN. imaguirei Soares &Soares, 1947.In the same year, a key for the species of Neopachylus was published by SoareS & SoareS (1947b).The last species to be described in the genus was Neopachylus taioensis Soares, 1966. Kury (2003), in his catalogue of the New World Laniatores, synonymized Gephyropachylus, Huralvius and Nunduavius with Neopachylus, and the genus is currently composed of nine species.Kury (2003) pointed out that Neopachylus species are very similar to each other and suggested a revision of the genus.The purpose of this paper is the taxonomic review of the genus Neopachylus.
Descriptions and illustrations.The descriptions are presented in alphabetical order and, with the exception of N. bellicosus, were based on the type material.Terminology follows acoSta et al. (2007), with the following exceptions proposed by Bragagnolo & Pinto-da-rocha (2012): granules are considered tegumentary microstructures smaller than 0.01 mm.The term armature refers to large regions of body/legs and are classified as: (1) apophyseshigh projections (normally higher than wide) with thinner or blunt tips, with no clear boundary with the surrounding cuticle, 2) setae-processes with thin apex and inserted in a socket (occur on pedipalps and penis), (3) tubercles-small projections without clear limit with the surrounding cuticle, normally as wide as high, apically thinner or cylindrical in all its extension and (4) a region is considered armed when one or two projections are higher/larger than any other in the same region.Terminology for the macrosetae of the ventral plate follows Kury & villarreal (2015).
A new term, rippled elevation, is created to describe the structure, present on scutal areas III-IV and free tergites in some Neopachylus species, that is higher than a tubercle, but lower than an apophysis.The terminology for pedipalpal setation length follows Pinto-da-rocha (1997).The females are described only insofar as they differ from the males.
Illustrations of external morphology were made using a Leica Leo 440 stereomicroscope with a camera lucida.The method of preparing the penis follows Pintoda-rocha (1997).Type measurements are in millimeters.Abbreviations in synonymy listing: cat, catalogue; cit, citation; descr, description; rdesc, redescription; syst, systematic.Abbreviations in descriptions: SL, scutum length; SMW, maximum scutum width; HOA, height of the ocularium apophysis; RPT, retrolateral apophysis bifid with prolateral stem short and truncated.
Distribution maps of the species: we used the computer software DIVA-GIS version 7.1.1.1 (hijManS et al., 2004) to make a map of where each species occurs.The records of geographical distribution were taken from the labels of the examined materials.The coordinates of geographical records were obtained from data banks available on the Internet: http://geonames.nga.mil/ and http://www.fallingrain.com/world/index.html(Directory of Cities and Towns in World -Global Gazetteer -version 2.2).The females have the same aspect as males regarding the coloration, ocularium apophysis and quantity and disposition of tubercles.However, there is sexual dimorphism in the armature of areas III-IV of dorsal scutum, posterior margin and free tergites, which is reduced.The armature of legs IV is also reduced in females, coxa with a small apical prolateral spiniform apophysis, remaining segments unarmed (except N. taioensis) Ocularium with a single spiniform apophysis, directed frontwards.Scutal areas I and IV divided by longitudinal groove; II and III undivided; areas III/IV with a par of strong paramedian rounded tubercles or oval rippled elevations.Posterior margins of dorsal scutum and free tergites with one row of small rounded tubercles or oval rippled elevations.Coxa IV with one prolateral apical apophysis blunt and bifid, one median subapical apophysis rounded, one retrolateral apophysis bifid with prolateral stem short and truncated (type RPT).Trochanter IV rectangular, with two prodorsal, two retrodorsal and 3-5 retrolateral apophyses (Fig. 36).Penis: ventral plate with distal and lateral margins concave, three pairs C1-C3 of latero-distal macrosetae, three pairs A1-A3 of latero-basal macrosetae, one pair on reduced macrosetae B clustered with A, more ventrally inserted, one pair D1 of lateral macrosetae between A and C; two pairs of reduced macrosetae E1-E2 on latero-distal flange; stylus with long stem, straight, subdistal filaments and ventral process almost parallel and smaller than stylus, thin, with apex curved downwards, serrate or smooth.
Penis .Ventral plate: distal and lateral margins concave; venter densely covered by simple microsetae on distal half, sparse setae on anterior half; three pairs of straight macrosetae C1-C3 aligned on latero-distal border of VP, C1-C2 subequal, inserted more closely to each other, C3 much shorter; macroseta D1 much reduced, lateral, midway between A and C; three pairs of subequal latero-basal straight macrosetae A1-A3 pointing proximally, close to each other in a triangle, median a little more ventrally placed.Stylus: stem long, straight, with subdistal filaments on lateral and ventral regions, frontwards directed, with swollen and smooth apex.Ventral process: almost parallel to stylus, thin, with apex curved downwards and serrate.
Coloration (in ethanol): Dorsal scutum, ocularium, coxa and tibia of legs IV, apophysis, tubercles, free tergites and anal operculum dark brown.Legs I, II, III and tarsus IV light brown.Chelicerae and venter light brown, dark brown reticulated.Pedipalps, scutal margins and scutal grooves, spots surrounding ocularium and frontal hump yellow.
Geographical distribution (Fig. 71): Restrict to the coast of Southernmost Brazilian States (Santa Catarina and Rio Grande do Sul).
Intraspecific variation: Males with 1-3 ripples on paramedian tubercles of areas III and IV.Males (n=21): SL Diagnosis.Neopachylus herteli is similar to N. imaguirei and N. mamillosus in that it presents areas III and IV of dorsal scutum with a pair of oval rippled elevations (5-7 ripples each) and all free tergites with oval rippled elevations.It differs from them in that it presents one additional smaller median pair on area IV (Figs 2; 9).Trochanter IV with a prodorsal median apophysis curved, directed downwards, differs from N. imaguirei and N. mamillosus, which present it rounded (Fig. 38).Male (Paratype MZSP 1632).Measurements: SL 4.55; SMW 4.05; femur I 1.95; femur II 2.90; femur III 2.40; femur IV 2.50; HOA: 0.60.Dorsum (Figs 2; 9): Ocularium with a single median long apophysis, slightly directed frontwards with six small tubercles on base.Prosoma with six small scattered tubercles.Lateral margins of dorsal scutum with 15-17 rounded tubercles, beginning at the anterior grove of area I and reaching posterior groove of area IV.Area I with a pair of slightly enlarged paramedian tubercles and 3-4 smaller tubercles each side.Area II with a pair of slightly enlarged paramedian tubercles and 10 smaller tubercles scattered.Area III with a pair of oval rippled elevations, with five ripples each, and 10 small tubercles uniformly distributed.Area IV with two pairs of oval rippled elevations, one with seven ripples and one smaller, with five ripples each and five small tubercles on each side.Posterior margin of dorsal scutum with one row of 10 small oval rippled elevations, with 4-5 ripples each, and seven small rounded tubercles.Free tergites: I with 10 small oval rippled elevations, with 3-4 ripples each, and five small rounded tubercles; II with eight small oval rippled elevations, with 3-4 ripples each, and seven small rounded tubercles; III with eight small oval rippled elevations, with three ripples each, and six small rounded tubercles.Anal operculum with 22 small rounded tubercles.
Trochanter IV with two prodorsal apophyses, one robust apical directed retrolaterally, one curved median directed downwards; two retrodorsal subapical apophyses geminate at base; five retrolateral apophyses, one large spiniform apical slightly directed backwards, one median conical, two small subbasal rounded and one basal conical, with a small rounded tubercle on its base.Femur IV curved, with one large, spiniform retrolateral basal apophysis, slightly directed backwards; one row of eight conical prolateral apophysis, median enlarged; ventrally with one row of six conical apophysis, from median to apical region, smaller than prolateral apophysis and one conical ventrobasal apophysis.Patella IV with a small spiniform prolateral apical apophysis.Tibia III with six small retrolateral conical apophyses.Tibia IV with prolateral and retrolateral rows of small conical setiferous apophyses in all extension.Diagnosis.Similar to Neopachylus herteli e N. mamillosus in that it presents areas III and IV, posterior margin of dorsal scutum and free tergites with oval rippled elevations and femur IV strongly curved.It differs from N. herteli in that it presents tubercles scattered on areas I-II (almost in a median line in N. herteli) (Fig. 3) and from N. mamillosus in that it has the prolateral apical apophysis of coxa IV with dorsal stem almost the same size as the ventral stem (Fig. 21) and lateral margins of dorsal scutum with one row of rounded tubercles of the same size, beginning at the anterior groove of area I and reaching the posterior margin of dorsal scutum (Figs 3; 10).Ocularium with a single median short apophysis slightly directed frontwards with six small setiferous tubercles on base.Prosoma with eight small scattered tubercles.Lateral margins of dorsal scutum with 22-20 rounded tubercles, beginning at anterior groove of area I and reaching the posterior margin of dorsal scutum.Area I with a pair of slightly enlarged paramedian tubercles, 6-7 small tubercles uniformly distributed on each side.Area II with a pair of slightly enlarged paramedian tubercles, 21 small tubercles uniformly distributed.Area III with a pair of oval rippled elevations, with six ripples each elevation and 25 small rounded tubercles uniformly distributed.Area IV with a pair of oval rippled elevations, with 7-8 ripples on each elevation, 11-15 small rounded tubercles on each side, uniformly distributed.Posterior margin of dorsal scutum with two pairs of oval rippled elevations on each side, with 3-8 ripples on each elevation, and row of 15 small rounded tubercles.Free tergites: I with a small oval rippled elevation (2-3 ripples by elevation) on each side, and a row of small rounded tubercles; II with one row of 14 tubercles; III with one row of nine tubercles.Anal operculum with 20 small rounded tubercles.
Penis  63): four small scattered tubercles on prosoma; areas I and II covered by small rounded tubercles, uniformly distributed, with the median pair enlarged.Armature of posterior margin, areas III and IV, not rippled as in male, represented by one pair of large rounded tubercles, with small tubercles uniformly distributed.
Geographical distribution (Fig. 71): It is known from highlands in the state of Paraná.

Neopachylus mamillosus
Male (Holotype SMF 1095).Measurements: SL 4.65;SMW 4.80;femur I 2.20;femur II 3.40;femur III 2.75;femur IV 3.00. Dorsum (Figs 4;11): ocularium with a single median apophysis (damaged) with six small tubercles on base.Prosoma with six small scattered tubercles.Lateral margins of dorsal scutum with 15-13 rounded tubercles, beginning at the anterior groove of area II and reaching the posterior groove of area IV, enlarged at area II/III.Area I with a pair of slightly enlarged paramedian tubercles and five small scattered tubercles on each side.Area II with a pair of slightly enlarged paramedian tubercles and 14 small tubercles uniformly distributed.Area III with a pair of robust oval rippled elevations, with six ripples each, and 27 small rounded tubercles uniformly distributed.Area IV with a pair of robust oval rippled elevations, with 7-9 ripples, and 9-12 small rounded tubercles uniformly distributed on each side.Posterior margin of dorsal scutum with 3-4 oval rippled elevations on each side (with 3-8 ripples each one), and a row of 13 small rounded tubercles.Free tergites: I with two small median oval rippled elevations (with 2-3 ripples each), on each side of tergite, and a row of 15 small rounded tubercles; II with one row of 13 small rounded tubercles; III with one row of eight small rounded tubercles.Anal operculum with 15 small rounded tubercles.
Legs (Figs 4;(24)(25)(26)36;40;67;68): coxa IV with prolateral apical apophysis bifid, dorsal stem over twice as long as ventral stem.Trochanter IV with two prodorsal apophyses, one small median rounded, one robust apical curved frontwards, with eight small tubercles on its apex and three tubercles on its base; two small rounded prolateral apophyses; two retrodorsal apophyses, one spiniform apical with three small rounded tubercles on its base, one subapical single and conical with one small rounded tubercle on its base; three retrolateral spiniform apophyses, one apical, one median (smaller), and one basal.Femur IV strongly curved, with one large spiniform retrolateral basal apophysis, slightly directed backwards, one row of three rounded prolateral median apophyses close to each other, one small prolateral apical apophysis; three retrolateral spiniform apophyses, one large, subapical directed backwards, one median, small with three small tubercles on its base, and one apical directed backwards, with small rounded tubercle on its base.Ventrally with two subbasal apophyses: one conical and other rounded, and three small basal tubercles.Patella IV with three small apophyses (prolateral, ventral and retrolateral).Tibia III with six small spiniform retrolateral apophyses, from base to apex.Tibia IV with prolateral and retrolateral rows of small conical apophyses in all extension.Tarsal counts: 6/9/7/6.Penis (Figs 58; 59): Ventral plate.Distal and lateral margins concave; densely covered by simple microsetae on venter; macrosetae C1-C3 subequal, straight; macrosetae A1-A3 straight, subequal, A2 more ventral; macrosetae B and D indistinct.Stylus.Stem long, straight, with swollen apex with subdistal filaments directed upwards on lateral and ventral sides.Ventral process.Almost parallel to stylus, thin, with apex curved downwards and serrate.
Female unknown.Geographical distribution (Fig. 71): It is known only from type-locality.
Material examined.Only type material.Mello-Leitão, 1936 (Figs 5;12;27-29;41;50-52;64;71) Nunduavius nebulosus Mello-leitão, 1936:11,  Diagnosis.Similar to Neopachylus bellicosus and N. taioensis in that it presents areas III and IV with a pair of large rounded and robust tubercles.It differs in that it presents short single median apophysis on ocularium, lateral margins of dorsal scutum and free tergites with one row of small rounded tubercles of the same size (Figs 5;12).Male femur IV is similar to that of N. bellicosus, but differ in that it presents three retrolateral median pointed apophysis (basal smaller), slightly directed backwards With a single short apophysis slightly directed frontwards, with five small rounded tubercles.Prosoma with five small scattered tubercles.Lateral margins of dorsal scutum with one row of 9-13 small rounded tubercles, beginning on area II and reaching the posterior groove of area IV.Area I with one pair of slightly enlarged paramedian tubercles, and two smaller on each side.Area II with one pair of slightly enlarged paramedian tubercles, and four scattered small tubercles.Area III with a pair of robust rounded paramedian tubercles, and four scattered small tubercles.Area IV with a pair of robust rounded paramedian tubercles, and three small tubercles on each side.Posterior margin of dorsal scutum with one row of 12 rounded tubercles, five median enlarged.Free tergites with small rounded tubercles: I with one row of 14 tubercles; II with one row of 13 tubercles; III with one row of 13 tubercles.Anal operculum with 15 small rounded tubercles.
Penis Legs IV (Fig. 64): coxa shorter than in males, with a small pointed prolateral apical apophysis; trochanter, femur, patella and tibia only covered by small setiferous tubercles.Tarsal counts: 6/ 8/ 7/ 7. Synonymic note: Neopachylus incertus was described based on a female and it is here synonymized with N. nebulosus due to morphological similarities (areas I-II of dorsal scutum with one pair of slightly enlarged paramedian tubercles; areas III-IV with a pair of robust rounded paramedian tubercles; posterior margin and free tergites with one row of small rounded tubercles) and couples were found in the same localities.
Geographical distribution (Fig. 71): recorded from interior of states of Paraná and Santa Catarina, Brazil.
Diagnosis.It is similar to Neopachylus herteli, N. imaguirei and N. mamillosus in that it presents areas III and IV of dorsal scutum with one pair of oval rippled elevations.The lateral margins of dorsal scutum with one row of rounded tubercles of same size from area I to IV and apophyses of trochanter IV make it very similar to N. imaguirei, it differs from them in that it has less ripples on elevations (2-4) of areas III and IV and posterior margin of dorsal scutum and free tergites with one row of small rounded tubercles of same size, without rippled elevations (Figs 6;13).
Male (Paratype MZSP 1769).Measurements: SL 4.50;SMW 4.30;femur I 2.25;femur II 3.25;femur III 2.80;femur IV 4.00;HOA: 0.40. Dorsum (Figs 6;13): ocularium with short single apophysis slightly directed frontwards, with eight small setiferous tubercles on base.Prosoma with five small scattered tubercles.Lateral margins of dorsal scutum with 14-15 small rounded tubercles, beginning at the anterior groove of area I and reaching the anterior groove of area IV.Area I with a pair of slightly enlarged paramedian tubercles and 8 tubercles on each side.Area II with a pair of slightly enlarged paramedian tubercles and 35 small rounded tubercles uniformly distributed.Area III with a pair of oval robust rippled elevations, with three ripples each, and 29 small rounded tubercles uniformly distributed.Area IV with a pair of oval robust rippled elevations, with 3-4 ripples each, and 12-13 small rounded tubercles uniformly distributed on each side.Posterior margin of dorsal scutum with one row of 14 small rounded tubercles.Free tergites with small rounded tubercles: I with one row of 19 tubercles; II with one row of 16 tubercles; III with one row of 13 tubercles.Anal operculum with 15 small rounded tubercles.
Coloration (in ethanol): dorsal scutum, ocularium, coxa-tibia of leg IV, apophyses, tubercles, free tergites and anal operculum dark brown.Legs I, II, III and tarsus Diagnosis.Similar to N. bellicosus and N. nebulosus in that it presents areas III and IV of dorsal scutum with a paramedian pair of rounded and robust tubercles.It differs in that it has lateral margins of dorsal scutum with one elevation near grove III (Fig. 7), trochanter IV with low apical prodorsal apophysis (smaller than other Neopachylus species) with apex directed frontwards (Fig. 43) and femur IV with two rounded dorsobasal apophyses geminate at base (exclusive of N. taioensis) (Fig. 33).
Male (paratype MZSP 1339).Measurements: SL 4.95; SMW 5.10; femur I 2.00; femur II 3.70; femur III 2.95; femur IV 3.75; HOA: 0.60 mm.Dorsum (Figs 7; 14): ocularium with single long apophysis, slightly directed frontwards, with six small tubercles on base.Prosoma with 5 small scattered tubercles.Lateral margins of dorsal scutum with one elevation on each side, near groove III, with a large rounded tubercle on each elevation and one row of 7-9 small rounded tubercles, beginning at posterior groove of area I and reaching the anterior groove of area III.Area I with a pair of slightly enlarged paramedian tubercles and two small tubercles on each side; Area II with a pair of slightly enlarged paramedian tubercles and 12 small tubercles uniformly distributed; Area III with pair of paramedian rounded robust tubercles and 15 small tubercles uniformly distributed; Area IV with a pair of paramedian rounded robust tubercles and 3-6 small tubercles on each side.Posterior margin of dorsal scutum with one row of 13 rounded tubercles, median enlarged.Free tergites with small rounded tubercles: I with one row of 15 tubercles; II with one row of 12 tubercles; III with one row of nine tubercles.Anal operculum with 12 small rounded tubercles.Legs (Figs 7;[33][34][35]43): coxa IV with prolateral apical apophysis bifid, dorsal stem larger than ventral stem.Trochanter IV with two prodorsal apophyses, one median rounded, one large apical with apex directed frontwards, with two small rounded tubercles between them; one subapical prolateral rounded apophysis; five rounded retrodorsal apophyses, one apical, four subapical; three pointed retrolateral apophyses, one basal, one median, one apical (directed backwards).Femur IV straight, with two rounded dorsobasal apophyses geminate at base; two dorsomedian curved retrolaterally apophyses; three small rounded prolateral subbasal apophyses; one small spiniform prolateral apical apophysis, with apex slightly directed backwards; one pointed retrolateral subapical apophysis; four ventral apophyses, two rounded basal close to each other, one subbasal conical, one median large rounded curved retrolaterally.Patella IV with two spiniform ventroapical apophyses.Tibia IV with one prolateral and one retrolateral rows of small conical setiferous apophyses, in all extension.Tarsal counts: 6/9/7/7.Penis (Figs 60; 61): Ventral plate.Distal and lateral margins concave; ventrally densely covered by simple microsetae, more sparse on basal region; macrosetae C1-C3 straight, C3 clearly smaller, C1-C2 longer and closer to each other; D1 much reduced, inserted between macrosetae A and C; macrosetae A1-A3 straight, clustered in triangle, A3 a bit smaller, A2 more dorsally inserted.Stylus.Long stem, straight, with subdistal filaments directed upwards, swollen and smooth apex.Ventral process.Almost parallel to stylus, thin, with apex curved downwards and serrate.
Geographical distribution (Fig. 71): It is known only from two localities east part of plateaeu in the Santa Catarina State.