Four new species of Cosmetus from Panama , with comments on the systematics of the genus ( Opiliones : Cosmetidae )

Four new sympatric species of Cosmetus Perty, 1833 are described from “Reserva Natural Privada Burbayar, Provinciar Panamá, Panamá” (male holotypes deposited in MIUP). Cosmetus balboa sp. nov. can be distinguished by the combinations of following features: smooth ocularium, larger distal tubercle on pedipalpal femur, coxa I with large ventral tubercle directed upwards, coxa IV with one large dorsoproximal tubercle, two geminate (from base) dorsoapical tubercles with blunt apex. Cosmetus burbayar sp. nov., can be distinguished from other species of the genus by the irregular and discontinuous shape of its yellow spot, extending from lateral anterior to posterior margins and invading prosoma, areas I-III and free tergites. Cosmetus pollera sp. nov. can be distinguished from congeners by two small yellow and two large pairs of spots on prosoma and two other spots on posterior margin of dorsal scutum. Cosmetus tamboritos sp. nov. is distinguished from other species of the genus by the following combinations of characters: having two retrolateral apical tubercles on bulla, one being double size of other; coxa IV lacks patches of a cluster of four tubercles on dorsolateral proximal region and two pointed tubercles fused at their apices; and femur IV with bifid retrolateral apical tubercle. The penis of Cosmetus arietinus (Mello-Leitão, 1940) and C. variolosus Mello-Leitão, 1942 are described for the first time. A table with the main diagnostic features of Cosmetus species is given. We suggest that the spine of area III, sexually dimorphic chelicerae and posterior legs, and pigmentation of dorsal scutum are good diagnostic features at species level.

Among harvestmen, Cosmetidae is the second most diverse family of the suborder Laniatores, with more than 700 species (KURY 2003(KURY , 2011(KURY , 2013)).They occur from southern United States of America to southern South America (with exception of Chile) (KURY 2003(KURY , 2009)).However, the greatest species richness is recorded from tropical forests of Central and South America and from the Andes (PINTO-DA-ROCHA & YAMAGUTI 2013).Adult Cosmetidae can be easily recognized by the spoonshaped pedipalps, in which the lateral compression of femoratibiae covers the chelicerae (KURY & PINTO-DA-ROCHA 2007).ROEWER (1912) divided the family in two subfamilies, based on the armature of claws III and IV: Cosmetinae, with smooth claws, composed of 680 species and 116 genera, and Discosomaticinae, with pectinate claws, including 29 species and 10 genera (KURY 2003, 2009, KURY & PINTO-DA-ROCHA 2007, PINTO-DA-ROCHA & HARA 2011).Using the morphology of the claws for dividing cosmetids into two subfamilies has been rejected since RINGUELET (1959).The smooth claws seem to be a plesiomorphic character, however there is no other character to support the monophyly of Cosmetinae and no hypothesis on how the pectinated claws evolved within the family (FERREIRA & KURY 2010).Since no cladistics hypothesis has been advanced for the internal relationships of cosmetids, it is impossible to understand the evolution of this feature, and the validity and composition of Discosomaticinae (FERREIRA & KURY 2010).
Carl F. Roewer proposed a system of classification for cosmetid genera based mainly on a combination of armature of dorsal scutum (e.g.areas, anal operculum) and number of tarsal segments (ROEWER 1916(ROEWER , 1923(ROEWER , 1927(ROEWER , 1947)).However, the number of articles is highly variable, even within the same species (see MELLO-LEITÃO 1933, KURY et al. 2007, PINTO-DA-ROCHA & YAMAGUTI 2013).The "Roewerian system" led to the inclusion of several species in unrelated genera, and the erection of dozens of monotypic genera and several similar genera (see KURY 2003).An attempt to review this system was made by GOODNIGHT & GOODNIGHT (1953), who studied intraspecific variation in Mexican Cosmetinae, and concluded that tarsus I is less variable than the tarsi of other legs and that it can be used to define genera.However, evidence in support of their classification is not consistent and for this reason it has not been accepted by other researchers (KURY 2009).In the last 25 years, a few new contributions on the taxonomy of cosmetids have revealed new features that could be useful, such as the morphology of the penis, a structure that can provide characters   (1905) designated Cosmetus varius Perty, 1833 as its type species.After that, a few species were added to Cosmetus by authors such as GERVAIS (1842,1844), SØRENSEN (1884), KOCH (1839), andSIMON (1880).The original concept of the genus, by PERTY (1833), was similar to that of the family until 1839, when KOCH (1839) described the following new genera: Cynorta, Flirtea, Gnidia and Paecilaema.The author included in them species previously assigned to Cosmetus (see PICKARD-CAMBRIDGE 1904, KURY 2003, KURY et al. 2007). Finally, KURY (2003) synonymized, within Cosmetus, all cosmetid genera with unpaired armature on area III (single or geminated spines), lateral margin of scutum without constriction, ocularium flattened and legs unarmed.The list of synonyms includes seven genera: Cosmetigryne Roewer, 1916;Cosmetellus Roewer, 1927;Belemnometus Mello-Leitão, 1940;Procosmetus Mello-Leitão, 1942;Vervloetia Soares & Soares, 1946;Cosmetiplus Roewer, 1947;andOrthogryne Avram &Soares, 1983 (KURY 2003).
Currently, Cosmetus includes 17 species that are distributed from Panama to northern South America (Venezuela, Ecuador, Colombia, Peru and Brazil) (KURY 2003).
The main goal of this paper is describe four sympatric new species of Cosmetus, from central Panama, the northern border of its distribution.Additionally we make some comparisons with other species of the genus in order to contribute towards the classification of the group.
All measurements are given in millimeters.Measurements of appendages are from the right side of specimens.Illustrations were made using a Leica MZ APO stereomicroscope with a camera lucida and microscope ZEISS Axioskop 2 Plus.The description of the dorsal armature followed DASILVA & GNASPINI (2009).
The male genitalia were prepared according to PINTO-DA-ROCHA (1997).Nomenclature of genitalic description followed KURY & VILLARREAL (2015).Descriptions of the external morphology of females only contain information on features that differ from the male.
Table 1 shows a set of characters that are useful to identify Cosmetus species.It is based on the material described herein, as well as literature and other museum specimens examined.The illustrations of the male genitalia of C. arietinus (Mello-Leitão, 1940)
Venter.Coxa I with one row of 14 tubercles and 13 sparse tubercles.Coxa II, III and IV with sparse setiferous tubercles.Genital operculum and stigmatic area with few sparse small tubercles.
Chelicera.Dimorphic.Bulla with one large tubercle on proximal retrolateral region, eight small retrolateral tubercles, one tubercle on median region and two-three tubercles on prolateral 1 Coloration.Dorsal scutum dark brown with lighter margins.A pair of small yellow spots close to anterior margin, two large yellow spots on prosoma, from lateral margin to area I, two large yellow spots on area III, from posterior margin to near base of spine.Chelicera, pedipalp, legs and free tergites light brown.
Venter.Coxa I with two rows of wide tubercles, one row near to coxa II with 9-11 tubercles, other row near pedipalpal coxa with four tubercles, short tubercles on posterior region, one distal tubercle directed towards coxa II.Coxa II with small tubercles, two small tubercles directed towards coxa I, four tubercles united to coxa III.Coxa III with five-seven tubercles united to coxa IV.Genital operculum with setiferous small tubercles, stigmatic area smooth.
Chelicera.Bulla covered by small tubercles, with four larger basal retrolateral tubercles, one prolateral basal tubercle, three prolateral apical tubercles.Segment II with three small teeth on fixed finger.Segment III with one basal tooth, with intracheliceral space.
Pedipalp (Figs 6,10).Trocanter with one ventro-distal tubercle.Femur compressed, with one row of 10 dorsal tubercles, one ventral row of 21 tubercles and one distal tubercle.Patella with one distal prolateral tubercle and one ventral tubercle.Tibia elongate, with two lateral distal spines.Tarsus with setae on prolateral and retrolateral regions.
Legs.Coxa I with one dorsal tubercle on each side, one directed towards pedipalp, other towards coxa II; ventral tubercle directed upwards, its apex almost reaching the tubercle near pedipalp, one retrolateral apical tubercle.Coxa II with one tubercle directed towards coxa I, one prolateral tubercle.Coxa III with one dorsal tubercle near coxa II.Coxa IV with one dorsal proximal and two tubercles with blunt apex united since their base on dorsal distal region.Trochanters I-IV tuberculate.Femora I-III with one basal tubercle, IV with one ventro-distal tubercle.Patellae I, II, III and IV with two dorsal distal tubercles.Tibiae smooth.Tarsal segmentation: 7(3), 16 (3), 13, 15.
Penis (MZSP OP 1565) .Ventral plate wide and subretangular, wider at apical region, lateral margins straight, distal corners rounded, distal margin concave.Densely covered with thin microsetae on lateral margin, from median to apical region.Two pairs of small setae on basal region (group A); two pairs of large and curved distal setae and one pair of cylindrical setae with pointed apex (only distally tapering) on the median lateral margin (group C); one pair of basal ventral short setae (group E).Glans elongate, dorsal process well developed.Serrate caruncle of stylus with long ventral projections almost until apex.
Coloration.Light yellow, free tergites, legs and distal half of area III spine darker.Patellae and apex of tibiae lighter.
Distribution.Known only from the type locality.Etymology.The specific name "balboa" (in apposition) is a patronym in honour of Vasco Nuñez de Balboa, the first European to explore Panama.It is also the name of the Panamanian currency, first coined in the 1940's.
Venter.Coxa I with anterior row of 10 large tubercles and several small, sparsely distributed.Coxae II, III and IV densely covered by sparse tubercles.Genital operculum with small tubercles, stigmatic area smooth.
Chelicera.Dimorphic.Bulla with one-two distal tubercles, four retrolateral tubercles, two tubercles on proximal dorsal region, three proximal prolateral tubercles.Tuberculate dorsally.Segment II with setae on distal region, with one basal tooth on fixed finger.Segment III with one basal tooth, with intracheliceral space.
Pedipalp (Figs. 7,11).Trochanter with two ventro-distal tubercles.Femur compressed, with one dorsal row of six tubercles, one ventral row of 16 tubercles.Patella with one distal retrolateral tubercle, two retrolateral distal ventral setae.Tibia with retrolateral margin depressed, with setae on lateral margin directed downwards, two wide setae on distal region.Tarsus with lateral margin covered with short setae, one ventral retrolateral setae larger.
Penis (MZSP 67286) .Ventral plate with shallow concavity on distal margin, lateral margins parallel on basal 2/3, apical 1/3 slightly wider.Ventro-median to distal margin with small triangle-like projection.Three pairs of curved and flattened dorso-distal setae (group C); three pairs of cylindrical setae on median region, distal most pair smaller than others (group D), without setae on basal region.Glans elongated, dorsal process well developed.Stylus swollen apically, serrate caruncle with short ventral projections until subapical region.
Coloration. Brown orange, reticulate with dark brown on dorsum, chelicerae and legs.Two large yellow and rounded spots on posterior margin.Yellow stripe on lateral margin from coxa I to posterior margin, projecting to prosoma near leg I. Yellow spot on anterior to center of area I and surrounding areas I, II and III.One yellow spot anterior to spine of area III, one behind it.One yellow spot from margin of area II to lateral margin.
Distribution.Known only from the type locality.Etymology.The specific name is a noun in apposition and refers to the type locality of this species, an astonishing private reserve in Burbayar.The name is taken from the Kuna language (spoken in southwestern Panama and northwestern Colombia) and means "spirit of the mountains".
Venter.Region between coxae II-III ahead of genital operculum strongly depressed.Coxa I with one anterior row of 12 large tubercles, 15 posterior sparse, four on apical region.Coxae II-IV densely covered by sparse tubercles.Genital operculum with six small tubercles.Posterior margin and free sternites with one row of small tubercles.Stigmatic area with small setiferous sparse tubercles.
Chelicera.Dimorphic.Bulla with three proximal prolateral tubercles, one proximal retrolateral tubercle, two retrolateral apical tubercles of different sizes, two large distal prolateral tubercles.Segment II with setae on distal region, fixed finger smooth.Segment III with one basal tooth, one median tooth enlarged, eight small distal teeth, with intracheliceral space.
Legs.Coxa I with two anterior tubercles fused at their base, one posterior tubercle directed towards coxa II.Coxa II with one anterior tubercle directed towards coxa I. Coxa III small apophysis directed towards coxa II.Coxa IV with a cluster of four well developed tubercles on dorsolateral proximal region, two pointed tubercles contiguous at their apices.Trochanter I with one enlarged retrolateral tubercle directed towards leg II, one distal retrolateral tubercle and four ventral tubercles; II with two prolateral basal tubercles directed towards leg I, one large retrolateral basal tubercle directed towards leg III, one tubercle on each lateral distal sideand four ventral tubercles; III with one enlarged tubercle on each lateral basal side, two smaller tubercles on each lateral distal surface, nine-ten ventral tubercles; IV with one enlarged tubercle on each basal lateral, two tubercles fused at their base on each lateral distal, nine ventral tubercles.Femora I and II with one row of dorsal small tubercles, one row of lateral small tubercles and two ventral rows of small tubercles; III with two dorsal rows of small tubercles, two ventral rows of large tubercles increasing in size to apex, proventral row of tubercles higher than the retroventral one; IV with two dorsal rows, one prolateral row of higher tubercles on anterior ¼, rows of tubercles increasing in size to apex, proventral row with higher tubercles than the retroventral one, with one apical retrolateral bifid tubercle.Patellae I and II smooth; III with small sparse tubercles; IV with two dorsal rows of eigth tubercles each (retrodorsal larger), one apical prodorsal tubercle longer than other of the segment, two retrodorsal longer than other of the same row.Tibia IV with two ventral rows of small tubercles, increasing in size to apex, retroventral apical larger than other, with distal apophysis.Basitarsus I swollen.Tarsal segmentation 7(3), 20(3), 11, 14.
Penis (MIUP) .Ventral plate quadrangular, lateral margins parallel and straight, distal margin straight; three pairs of distal long setae (group C), two distal most pairs falciform, two intermediary pairs of straight setae and shorter (half the length of the distal setae) (group D), two rounded areas on ventral region.Truncus with one ventral short setae (group E).Glans elongate and thin.Serrate caruncle of stylus with short projection from ventral region to almost apex.
Coloration.Dorsal scutum and coxae dark brown.Chelicera brown.Legs and pedipalps light yellowish brown.Yellow patch U-shaped covering irregularly anterior margin, lateral surface of prosoma and areas, reaching the posterior margin and forming a narrow stripe behind spine.Dorsal scutum dark brown with small irregular yellow patches.Free tergites I-III with one stripe each.
Anterior margin narrower than male.Ocularium with nine small tubercles near eyes.Area II with nine small tubercles.Genital operculum with nine small tubercles.Bulla densely tuberculate dorsally, three-four tubercles on lateral surface of distal prolateral large tubercle, two proximal central dorsal tubercles, retrolateral tubercles smaller than male; apical retrolateral and prolateral tubercles well developed, movable finger with small teeth.Pedipalpal femur with one row of 10-11 ventral tubercles, dorso-proximal row with four tubercles.Pedipalpal tibia with four small prolateral distal tubercles.Trochanter II with seven ventral tubercles.Femora III-IV with short tubercles, except on apical region of femur IV, with ventral tubercles slightly longer than the others.Patella IV with two dorsal rows of tubercles, three distal prolateral tubercles, several small tubercles on ventral region.Tibia IV small-tuberculate.Basitarsus I with similar diameter as distitarsus.Distribution.Known only from the type locality.Etymology.The specific name "tamboritos" is a noun in apposition taken from the popular culture of Panama that refers to a drum style of music and a typical dance.Tamborito is danced by men and women from the central provinces of the country, dressed with colorful clothes.

Description of the penis of two additional Cosmetus species
Cosmetus arietinus (Mello-Leitão, 1940) (MZSP 14038) .Ventral plate elongate, distal margin concave, two apical pairs of long and curved setae and one pair median apical of short setae (group C).Two basal pairs of setae (group A).One pair ventral basal of short setae (group E).Truncus narrow and with the same diameter throughout almost all its extension.Glans elongated, with dorsal process.Serrate caruncle of stylus with short projections on apical region.
Cosmetus variolosus Mello-Leitão, 1942 (MZSP 238) (Figs. 28-30).Ventral plate elongate, distal margin slightly concave, two groups of lateral setae, one basal pair (group A) and one median pair of short setae (group D) (basal most twice as long as median pair), three pairs of apical long and curved setae (group C) (basal most smaller than others).Truncus elongated and with same diameter throughout almost all its extension.Glans elongated, with dorsal process almost as high as stylus.Serrate caruncle of stylus with elongated projections from median region to apex.

DISCUSSION
Among Laniatores, Cosmetidae has an impressive number of species, being almost as rich as Gonyleptidae, the richest family in this suborder.However, it has received much less attention than gonyleptids (see references in PINTO- DA-ROCHA et al. 2014).The limited interest on cosmetids is likely due to the absence of modern opiliologists in regions where the group is most diverse (Amazonian and Andean regions, and Central America), and the fact that researchers that work on other Neotropical groups are not very interested on cosmetids.An exception is the impressive work of GONZALEZ-SPONGA (1992), who described and redescribed 92 cosmetid species from Venezuela.Thus, it was not a surprise to find four new species of Cosmetus in the northern part of its distribution, on Chocó-Darién moist forest ecoregion (WWF 2015).That region also harbors the species C. flavopictus Simon, 1880.Northern to Chocó-Darién is the Isthmian-Atlantic moist forests ecoregion, where the northernmost record of the genus, the Panamanian species C. pulcher (see KURY 2003 for a general view on distribution of the genus), is found.
Recently, cosmetids have received somewhat increasing attention and new sets of characters have been proposed to enhance its generic classification, such as: the shape of the dorsal scutum (KURY et al. 2007) However, a hypothesis on the morphological evolution of the groups is still lacking.
The morphology of the spine on area III of the male was the main character used by KURY (2003) to place several similar cosmetids in Cosmetus.However, this feature is variable, and two species-groups can be recognized based on it: a single spine is observed in C. arietinus (Mello-Leitão, 1940) ).There are differences in the shape of the distal margin of the penis (straight or concave) and the number of setae in the basal groups of the penial ventral plate.
The structure of the spine of area III, sexual dimorphism of chelicerae and armature of posterior legs, and pigmentation of dorsal scutum (see Table 1) are good diagnostic features at the species level.However, at present, without a phylogenetic hypothesis, it is difficult to recognize species clades within Cosmetus, or even to know if the genus is monophyletic.

1
Departamento de Zoologia, Instituto de Biociências, Universidadede São Paulo.Caixa Postal 11461, 05422-970 São  Paulo, SP, Brazil.E-mails: amandacoronato@hotmail.com; ricrocha@usp.brABSTRACT.Four new sympatric species of Cosmetus Perty, 1833 are described from "Reserva Natural Privada Burbayar, Provinciar Panamá, Panamá" (male holotypes deposited in MIUP).Cosmetus balboa sp.nov.can be distinguished by the combinations of following features: smooth ocularium, larger distal tubercle on pedipalpal femur, coxa I with large ventral tubercle directed upwards, coxa IV with one large dorsoproximal tubercle, two geminate (from base) dorsoapical (Mello-Leitão, 1940) apex.Cosmetus burbayar sp.nov., can be distinguished from other species of the genus by the irregular and discontinuous shape of its yellow spot, extending from lateral anterior to posterior margins and invading prosoma, areas I-III and free tergites.Cosmetus pollera sp.nov.canbedistinguishedfrom congeners by two small yellow and two large pairs of spots on prosoma and two other spots on posterior margin of dorsal scutum.Cosmetus tamboritos sp.nov. is distinguished from other species of the genus by the following combinations of characters: having two retrolateral apical tubercles on bulla, one being double size of other; coxa IV lacks patches of a cluster of four tubercles on dorsolateral proximal region and two pointed tubercles fused at their apices; and femur IV with bifid retrolateral apical tubercle.The penis of Cosmetus arietinus(Mello-Leitão, 1940)and C. variolosus Mello-Leitão, 1942 are described for the first time.A table with the main diagnostic features of Cosmetus species is given.We suggest that the spine of area III, sexually dimorphic chelicerae and posterior legs, and pigmentation of dorsal scutum are good diagnostic features at species level.KEY WORDS.Cosmetinae; harvestmen; systematics; taxonomy.ZOOLOGIA 32(5): 409-422, October 2015

Table 1 .
Diagnostic characters for the 21 species of Cosmetus .
Goodnight & Goodnight, 1942ight, 1942single present on both one row of TUB / one row of TUB --C.serrulatus (González-Sponga, 1992) geminate (distal half) present on both short basal TUB / one row of long TUB -none C. soerenseni (Mello-Leitão, C. unispinosus (Roewer, 1916) geminate (distal half) present on both ---C.variolosus Mello-Leitão, 1942 geminate (distal half) present on both -straight one C. varius Perty, 1833 geminate absent on both sparse TUB / one row of TUB --C.pollera sp.nov.geminate (distal half) *absent on both smooth / one row of long TUB *deep V-cleft *three C. balboa sp.nov.single present on both basal TUB / one row of long TUB *shallow concavity *two C. burbayar sp.nov.single present on both basal TUB / one row of long TUB *shallow concavity *none C. tamboritos sp.nov.geminate (distal half) present on both basal TUB / *one row of long TUB on median region *straight *none C. coxaepunctatus Roewer, 1927 irregular ST on LM and lateral sides of dorsal areas, small rounded ST smooth pointed sSmooth -C.flavopictus Simon, 1880 small rounded ST, narrow stripe on EM of AM and LM smooth blunt smooth absent C. mesacanthus Kollar in Koch, 1839 pale with small dark rounded ST smooth pointed smooth absent C. migdaliae (González-Sponga, 1992) irregular and discontinuous ST from PR to PM, small rounded ST smooth pointed one row of short TUB present C. peruvicus (Avram & Soares, 1983) C. burbayar sp.nov.*irregular discontinuous ST from PR to PM smooth pointed smooth present C. tamboritos sp.nov.irregular ST along all LM smooth pointed prolateral and ventral rows of long TUB, *bifid retrolateral apical TUB present (ROEWER 1927)metus balboa sp.nov.(Figs.2,10)issimilar to C. columnaris(ROEWER 1927)in having a high, tuberculate and blunt spine on area III and dorsal scutum without spots (see ROEWER 1927: 627, fig.50).It is also similar to C. biacutus Roewer, 1947 in having two tubercles on the apex of the spine on area III and one proximal dorsal tubercle on coxa IV (see ROEWER 1947: 31, fig.102).C. balboa sp.nov.can be distinguished by its smooth ocularium (ocularium with tubercles in C. columnaris, see ROEWER 1927), larger distal tubercle on pedipalpal femur (tubercles of same size in C. columnaris and C. biacutus), coxa I with large ventral tubercle directed upwards (C.columnaris and C. biacutus without ventral tubercle directed upwards, see ROEWER 1927 and ROEWER 1947: 31, pl.12, fig.102 respectively), coxa IV with one large dorsoproximal tubercle (C.columnaris possesses three dorsoproximal tubercles), two geminate (from base) dorsoapical tubercles with blunt apex (C.columnaris possess two apical tubercles not geminate, see ROEWER 1927: 627, fig.50 and C. biacutus have one distal apophysis, see ROEWER 1947: 31, pl.12, fig.102).