Early Lineages of Gonyleptidae (Arachnida Opiliones Laniatores)

Tropical Zoology 7: 343-353, 1994

Early lineages of Gonyleptidae (Arachnida Opiliones Laniatores)

ADRIANo B. KuRY Escola de Ciencias Biol6gicas, Universidade do Rio de Janeiro, Rua Frei Caneca 94, Rio de Janeiro 20.211-040, RJ, Brazil

Received 8 June 1993, accepted 18 December 1993

As result of cladistic methodology applied to the study of Gonyleptidae, four precocious branches are newly recognized in the phylogeny of the family. Three new subfamilies are proposed and diagnosed and a new concept is given for the Bout guyiinae Mello-Leitao 1923. Metasarcinae, a new subfamily is proposed for the Bolivian Metasarcus Roewer 1913 (formerly in Mitobatinae) and an undescribed Argentinean genus. Chaconatus Roewer 1929 (formerly in Prostygninae) is herein considered to be a new junior subjective synonym of Metasarcus. Heteropachylinae, a new subfamily is proposed for nine genera formerly described as Gonyleptinae or Pachylinae, and distributed along the Brazilian coast. Cobaniinae, a new subfamily is proposed for the genus Cobania Roewer 1913. Bourguyiinae should be the correct spelling of a subfamily name based on the generic name Bourguyia Mello-Leitao 1923. In the present new concept, four of its five genera are removed, while eight others are transferred from the Gonyleptinae, Pachylinae and Progonyleptoidellinae. To recon struct the phylogeny of Gonyleptic,lae, 16 morphological characters were analysed through HENNIG 86, providing a single most parsimonious cladogram of CI = 0. 77. The branching pattern obtained is: (Metasarcinae (Heteropachylinae (Cobaniinae (Bourguyiinae + the other Gonyleptidae))). Piresa angulispinosis H . Soares 1966 is newly referred to Mangaratiba Mello-Leitao 1940.

KEY WORDS: Arachnida, Gonyleptidae, new subfamilies, cladistics, Neotropics.

Introduction 344 Phylogeny 344 Systematic accounts 349 Metasarcinae n. subfam. 349 Heteropachylinae n. subfam. 350 Cobaniinae n. subfam. 351 Bourguyiinae Mello-Leitao 1923 351 Conclusions 35~ Acknowledgements 352 References 352 344 A.B. Kury

INTRODUCTION

The increasing knowledge of male genitalia in the Gonyleptidae, interpreted in the light of cladistic methodology, will surely lead to major changes in the concept of this family. As a first step, the Cranainae and related subfamilies were considered not to form a monophyletic group with the rest of the Gonyleptidae, which have the Cosmetidae as a sister group (KURY 1992). The comparative study of genital morphology (mainly of Brazilian genera), usually available from modern descriptions, brings new strength to corroborate this view. The present discovery of

PHYLOGENY

Based on previous study of the phylogeny of Gonyleptidae and related families (KURY 1992), Gonyleptidae and Cosmetidae are here assumed to be sister groups. The monophyly of the Cosmetidae is clearly supported by many autapomorphies in relation to the groundplan Cosmetidae/Gonyleptidae. The typical cosmetid penis possesses a well defined rectangular ventr.al plate, with a-short and thick stylus bearing a spiny mat and a thumb-like dorsal process arising from the glans (a similar structure is found in the Stygnidae). The typical gonyleptid penis shows also a well defined ventral plate, either rectangular or piriform, the glans bears a slender sigmoid stylus and a fan-like ventral process; a dorsal process is absent. The four groups of «living-fossils» cited above show intermediate features on penial morphology. Sixteen morphological characters, listed in Table 1, were cladisti cally analysed to construct a most parsimonious hypothesis of relationship among the major groups of Gonyleptidae. The matrix has been analysed through H ENNIG 86 (FARRIS 1988), with Cosmeti dae as an immediate outgroup. Trees were rooted by a hypothetic;al ancestor plesio morphic for all character states (Table 2). The multistate character 1 was considered as unordered, because there was no obvious degree of similarity justifying any particular sequence of transformation. The exact algorithm used here - «implicit enumeration» - yielded a single completely resolved most parsimonious tree (Fig. 6) with length 22, CI (consistency index) = 0. 77 and RI (retention index) = 0. 73. The following hypotheses of character transformation (see characters mapped on cladogram of Fig. 6) better explain the data: the clade Cosmetidae/Gonyleptidae acquired as groundplan synapomorphies the saddle-shaped eye mound (char. 1, Fig. 2b) and spiny mat on stylus (char. 2, Fig. 5b). As an alternative optimization, the Early gonyleptid lineages 345

Table 1. Morphological characters used to reconstruct the phylogeny of Gonyleptidae. Zero is the plesiomorphic state, 1 is the apomorphic condition for a binary character, a-b are diferent apomorphic states of a multistate character.

1. Eye mound: (O) convex, without depression, (a) saddle-shaped, (b) flattened, with hemispherical median tubercle. 2. Stylus: (0) ventrally smooth, (1) with mat of spines. 3. Chelicerae of male: (0) much swollen, (1) as that of female. 4. Femur IV of male: (O) without dorso-basal apophysis, (1) with pointed dorso-hasal apophysis. 5. Lateral margin of scute: (0) without special features, (1) bordered by a crest of blunt tubercles. 6. Apophysis of male coxa IV: (0) single branched, (1) bifurcated. 7. Ventral process of glans penis: (0) absent, (1) present. 8. Dorsal process of glans penis: (0) present, (1) absent. 9. Free tergite III: (0) without remarkable armature, (1) with conic projection, immense in male, smaller in female. 10. Pedipalpal femur: (0) without ventral armature, (1) with ventral row of stout spines. 11. Pedipalpal patella: (0) unarmed, (1) with a stout mesal spine. 12. Femur IV of male: (0) short and armed with rows of spines, (1) elongate and weakly armed. 13 . Free tergites: (0) independent of scute and with rounded corners, (1) with projected corners and more or less fused with scute. 14. Area III of dorsal scute: (0) with two paramedian spines, (1) unarmed. 15 . Tegument: (0) coriaceous, yellow to brown, (1) glossy black. 16. Microsculpture of dorsal scute: (O) finely granular, (1) densely · grouped round tubercles.

Fig. 1. - Bourguyia a/biornata Mello-Leitao 1923 (Museu de Zoologia da Universidade de Sao Paulo 10.580): distal part of penis, dorso-lateral view. This structure marks the transition between the typical cosmetid glans to the typical gonyleptid one; d = dorsal process, s = stylus, v = ventral process. 346 A.B. Kury

Table 2. Matrix of distribution of character states for six terminal taxa of Gonyleptidae and the outgroup.

1 1 1234567890123456 Ancestor 0000000000000000 Cosmetidae a100000000000000 Metasarcus a-----0-----11110000 Undescribed genus a111000011110000 Heteropachylinae 0111100000001100 Cobania 0011100000000111 Bourguyiinae b010111000010100 Other Gonyleptidae 0011111100000000

.~... b

~- 1!...... ~ .. . c

Fig. 2. - Transformation series of the eye mound in Gonyleptidae: (a) is the condition present in the primitive Tricommatidae and cranaine-like groups; (b) occurs in the ground plan of the Cosmetidae and in the Metasarcinae (and convergently in the Agoristenidae and in highly derivative Gonyleptidae as the Caelopyginae); (c) is exclusive of the Bourguyiinae.

a b

e Fig. 3. - Transformation series of opistosomatic characters. Abdominal scute: (a) area III armed, lateral margins without special tuberculation as in Cosmetidae and Metasarcinae; (b) area III unarmed, lateral margin bordered by a crest of rounded tubercles as in most Gonyleptidae. Free tergites: (c) as in the Cosmetidae, Stygnidae and many Gonyleptidae; (d) as in the Metasarcinae; (e) as in the Heteropachylinae. Early gonyleptid lineages 347

Fig. 4. -Transformation series of appendicular characters. Chelicerae: (a) swollen as in the groudplan of Stygnidae, Cosmetidae and cranaine-like groups; (b) as in the Gonyleptidae. Pedipalps: (c) femur and patella poorly armed as in most Gonyleptidae; (d) femur with a row of ventral spines and patella with stout mesal spine as in the Metasarcinae. Leg IV of male: (e) simple coxal apophysis as in the Cosmetidae, Metasarcinae, Heteropachylinae and Cobaniinae; (f-g) bifid apophysis as in the Bourguyiinae (/) and the other Gonyleptidae (g) ; (h) femur without dorso-basal apophysis as in the Cosmetidae and Metasarcinae; (i) femur with apophysis as in most Gonyleptidae; (j) femur short and curved as in the groundplan of Cosmetidae, the Heteropachylinae, Cobaniinae; (k) elongate femur as in the Metasarcinae and Bour guyiinae.

'f?fd- f(e Fig. 5. - Transformation series of genital characters. (a) Stylus smooth as in the Stygnidae, cranaine-like groups and most Gonyleptidae; (b) stylus with a spiny mat as in the Cosmetidae and early derivative Gonyleptidae; (c) glans only with thumb-like dorsal process, as in the Stygnidae, Cosmetidae and early derivative Gonyleptidae; (d) also with a lamellar ventral process as in the Bourguyiinae; (e) only with the ventral process as in the advanced Gonyleptidae. 348 A.B. Kury

Cosmetidae Metasarcus 1a 2 9 10 11 12 Unnamed genus 13 Heteropachylinae 15 16 Cobania Bourguyiinae ... Synapomorphy -{} Reversal Other Gony leptidae -ID- Convergence 8 R14

Fig. 6. - Cladogram showing the most parsimonious hypothesis of sister group relationships among the major groups of Gonyleptidae. Black rectangles are unique synapomorphies, empty rectangles are rever sals, hatched rectangles are convergences. There is another possible optimization for character (1) (convergent appearance in Cosmetidae and in Metasarcinae). See text for further details.

saddle shaped eye mound was independently acquired by the Cosmetidae and Meta sarcinae. The Gonyleptidae are characterized by chelicerae not swollen in male (char. 3, Fig. 4b) and dorsa-basal femoral apophysis in leg IV (char. 4, Fig. 4i). The Metasarcinae retained this condition of eye mound (char. 1), the paired spines in area III (char. 14, Fig. 3a), and the featureless lateral margin of scute (char. 5, Fig. 3a), the derived alternatives are therefore synapomorphic for the other Gonyleptidae. As synapomorphies for the two genera of the Metasarcinae we have the armature of free tergite III (char. 9, Fig. 3d), the structure of pedipalp (chars 10-11, Fig. 4d) and the elongate and slender femur IV of male (char. 12, paralleled in the Bourguyiinae, Fig. 4k). The Heteropachylinae are specialized miniature forms, showing tergites fused with scute, and bearing lateral projections (char. 13, Fig. 3e) . The Heteropachylinae and Metasarcinae still have the primitive condition of a short stylus with a spiny mat like the Cosmetidae, which separates them from all other Gonyleptidae (char. 2, Fig. 5b). Cobania, which constitutes here the monotypic subfamily Cobaniinae, represents one more step towards the typical Gonyleptidae. The glossy black colour and the rounded scutal tubercles (chars 15-16), and perhaps the strongly developed coxa IV in male, may be cited as autapomorphies for the genus. All remaining Gonyleptidae are separated from Cobania by the sharing of bifurcated coxal apophysis in leg IV (char. 6, Fig. 4g) and the presence of a ventral lamellar process in the stylus (char. 7, Fig. 5d e). The fourth precocious branch of Gonyleptidae is represented by the Bourguy1inae, a highly homogeneous group, supported by many autapomorphies not included in the present analysis, such as: pedipalpal tibia with three ectal spines (IIi), semicircular shape of fan of glans, distal setae of ventral plate elongated and convergent, black pigmentation aglomerated in an open reticle. The penis of the Bourguyiinae has frozen a peculiar state in the transformation series, in having both ventral and dorsal Early gonyleptid lineages 349

processes (Figs 1 and 5d). The remnant Gonyleptidae, which include the major part of the species, with many subfamilies, are thought to be a monophyletic group on the basis of the loss of the thumb-like dorsal process of stylus (char. 8, Fig. 5e).

SYSTEMATIC ACCOUNTS

Metasarcinae n. subfam.

Mitobatinae (part): RoEWER 1913: 284; 1923: 508; MELLO-LEITAO 1932: 390; SoARES & SoAREs 1949: 224.

Type genus. Metasarcus Roewer 1913. Included genera. There is undescribed material from Northwestern Argentina, which will probably be described as a new genus (E.A. MAURY pers. comm.). Descriptions and drawings of habitus and male genitalia, kindly provided by Dr Maury, allow the present proposal of a presume sister group relationship between this undescribed species and both Bolivian nominal species Metasarcus bolivianus Roewer 1913 and Metasarcus armatipalpus (Roewer 1929). Distribution. Bolivia and Argentina. Diagnosis. Gonyleptidae without ventral process in glans penis, with thumb-like dorsal process; stylus with spiny mat; ventral plate roughly rectangular, constricted in middle; distal setae short, not helicoidal and parallel. Area III of dorsal scute with a pair of spines, free tergite III with an immense conic projection, remaining areas and free tergites unarmed; eye mound wider than long, low, with median depression. Coxa IV of male hypertelic, armed with a short apical apophysis; trochanter IV of male armed with a small inner distal spiniform apophysis; femur IV of male elongate, sub straight, armed with very weak spines.

Metsssrcus Roewer 1913

Metasarcus RoEWER 1913: 304 (type species M. bolivianus); 1923: 517; MELLO-LEITAO 1926: 140; RoEWER 1931: 107; MELLO·LEITAO 1932: 408; 1935: 108; SoARES & SoARES 1949: 234. Chaconatus RoEWER 1929: 276 (type species C. armatipalpus); MELLO·LEITAO 1932: 108. New synonymy.

Metsssrcus srmstipslpus (Roewer 1929) n. comb.

Chaconatus armatipalpus ROEWER 1929: 276, fig 43. Remarks. Chaconatus armatipalpus was originally referred to the Prostygninae by ROEWER (1929), while Metasarcus bolivianus was placed in the Mitobatinae. There is no positive evidence for either of the assignations, furthermore both species are very closely related, if not the same. The second generic name created by RoEWER seems superfluous, since it adds no information beyond that supplied by different specific names. 350 A.B. Kury

Heteropachylinae n. subfam.

Gonyleptinae (part): RoEWER 1913: 167; 1923: 463; MELLO-LEITAO 1932: 231; SoARES & SoARES 1949: 151. Pachylinae (part): RoEWER 1913: 10; 1923: 396; MELLO-LEITAO 1932: 131; SoAREs & SoARES 1954: 225.

Type genus. Heteropachylus Roewer 1913. Included genera. Aesotrinoma H. Soares 1977, Canopilio H. Soares 1968, Chave sincola Soares & Soares 1946, Heterogonyleptes Roewer 1913 ( = Triaenosoma Roewer 1913 = Pseudotriaenosoma Mello-Leitao 1927), Heteropachylus Roewer 1913, Manga ratiba Mello-Leitao 1940, Me/loa Roewer 1930, Thaumatoleptes Roewer 1930, Tribu nosoma Roewer 1943. Distribution. Brazilian Atlantic Fares;:, from Parana state to Ceara state. Diagnosis. Gonyleptidae without ventral process in glans penis, with thumb-like dorsal process; stylus with spiny mat, ventral pL:te rectangular, with distal border slightly concave; distal setae short, not helicoidal and parallel. Area III of dorsal scute with variable armature, mostly unarmed, the free tergites, somewhat fused with scute, may bear median spines; eye mound wider than long, moderately high, with a pair of tubercles. Coxa IV of male hypertelic, armed with a short apical apophysis; trochanter IV of male armed with a small inner distal spiniform apophysis; femur IV of male short, curved, armed :Jlith small spines. Remarks. The Heteropachylinae include small gonyleptids, mostly described under monotypic genera. They occur along the Atlantic coast mainly in Northeastern Brazil, area rich in miniature forms of Stygnidae, Cosmetidae and «Phalangodidae Phalangodinae», but where Gonyleptidae are poorly represented. So far it is not possible to draw an accurate picture of relationships among its genera.

Msngaratiba Mello-Leitao 1940 Mangaratiba MELLO-LEITAO 1940: 2 (type Mangaratiba monstrosa Mello-Leitao 1940); SoARES & SoARES 1954: 272.

Diagnosis. Heteropachylinae with anterior margin of cephalothorax smooth; all scutal areas unarmed, scutal area IV discrete, entire; free tergites I-II with lateral teeth, free tergite III with median spine; tarsus I five-jointed, coxa IV of male with sharp spiniform dorsal and ventral apical apophyses; trochanter IV of male dorsally 'Inarmed.

Msngarstiba angulispinosa ~H. Soares 1966) n. comb. Piresa an&ulispinosis H. SoARES 1966: 95, Fig. 8.

Remarks. The genus Piresa Roewer 1927 includes mainly Brazilian species, which are typical Gonyleptidae. P. angulhpinpsis (from Parana, Southern 3razil) was put in this genus by the overall similarity, but there is no synapomorphic evidence to conserve this arrangement. It appears to be the closest relative of Mangaratiba monstrosa Mello-Leitao 1940 (from Rio de Janeiro, Southeastern Brazil), and there fore it is here removed to the hitherto monotypic genus Mangaratiba. Early gonyleptid lineages 351

Cobaniinae n. subfam.

Gonyleptinae (part): SoARES & SoARES 1949: 151. Pachylinae (part): RoEWER 1913: 10; 1923: 396; MELLO-LEITAO 1932: 131; SoAREs & SoARES 1954: 225.

Type genus. Cobania Roewer 1913. Included genus. Only the type. Diagnosis. Gonyleptidae without ventral process in glans penis, with thumb-like dorsal process; stylus without spiny mat, ventral plate rectangular, elongate, wider at base, with a slight median depression in the distal margin; distal setae short, not helicoidal and parallel. Areas of dorsal scute and free tergites I-III entirely unarmed; eye mound wider than long, moderately high and unarmed. Coxa IV of male hypertelic, armed with a stout outer single branched apical apophysis; trochanter IV of male armed with a stout inner distal spiniform apophysis; femur IV of male short, sigmoid, armed with robust spines. Remarks. Cobania picea (Bertkau 1880) is a robust gonyleptid, with remarkably developed coxae IV, endemic to the «campos rupestres» (open formation of Brazilian highlands, above 2,000 m high) of Itatiaia.

Bourguyiinae Mello-Leitao 1923 Bourguyinae MELLO-LEITAO 1923: 128; 1926: 348; ROEWER 1929: 265; MELLO-LEITAO 1932: 221; SoARES & SOARES 1948: 556. Gonyleptinae (part): ROEWER 1930: 342; SoAREs & SoAREs 1949: 151. Mitobatinae (part): SIMON 1879: 226; RoEWER 1913: 284; 1923: 508; MELLO-LEITAO 1932: 390; SoARES & SoARES 1949: 224.

Type genus. Bourguyia Mello-Leitao 1923. Included genera. Asarcus Koch 1839, Bogdana Mello-Leitao 1940, Bourguyia Mello-Leitao 1923, Cnemoleptes Mello-Leitao 1941, Caldasius Roewer 1930, Opistho plites Soerensen 1884, Stylopisthos Roewer 1930, Styloleptes Piza 1943. Distribution. Serra do Mar and Serra da Mantiqueira (mountain chains of Southeastern Brazil) from about 600 to 1600 m high. Diagnosis. Gonyleptidae with flabelliform ventral process in glans penis, in most genera also a thumb-like dorsal process; stylus without spiny mat, ventral plate rectangular, elongate, with a slight median depression in the distal margin; distal setae long, not helicoidal and convergent in the apex. Dorsal scute and free tergite I entirely unarmed; free tergites II-III armed with a median conic tubercle; eye mound much wider than long, very low and armed with a small round tubercle or unarmed. Coxa IV of male hypertelic, armed with an external apical apophysis formed by two parallel branches; trochanter IV of male in some genera armed with a stout internal distal hook; femur IV of male elongate, sub-straight, armed only with a few straight distal spines; femur IV of female sigmoid. Black reticule of dorsum concentrated in very small spots. Remarks. The correct spelling of a subfamily name based on the generic name Bourguyia should be Bourguyiinae. This is another monophyletic group never before recognized. The available name Bourguyiinae is currently applied to Bourguyia and four other genera that should be assigned to other subfamilies (see Table 3). On the 352 A.B. Kury

Table 3. Genera currently included in Bourguyiinae and which should be removed from this subfamily.

Camarana Mello-Leitao 1935 (---+ Tricommatidae). Discocyrtoides Mello-Leitao 1923 (---+ Mitobatinae; see KuRY 1990). Hypophyllonomus Giltay 1928 (---+ Pachylinae). Segadasius Mello-Leitao 1949 (junior synonym of Despirus Roewer 1929, [KuRY in prep.]; subfamily uncertain).

other hand, other genera, scattered in other subfamilies should be included: from Mitobatinae (Asarcus, Bogdana and Cnemoleptes), Progonyleptoidellinae (Opisthoplites and Stylopisthos) and Gonyleptinae (Caldasius and Styloleptes) . These eight genera include about 20 species from Brazil.

CONCLUSIONS

With the establishment of a sequence of basal sister groups, an important step was taken towards a phylogenetic analysis of the Gonyleptidae, the most important group of Neotropical Opiliones, which deserves more intensive study. It is noteworthy that the typical Gonyleptidae are by far most abundant in the Mata Atlantica, where the relict groups are poorly, if at all, represented. The marked asymmetry of the branching pattern is a common result, and may reflect modes of speciation. The next target is to dismantle the large subfamily Pachylinae into monophyletic units, and to check the monophyly of the extant subfamilies.

ACKNOWLEDGEMENTS

I wish to thank Dr E.A. Maury (Museo Argentino de Ciencias Naturales, Buenos Aires) for kindly sharing his unpublished results of research on Argentinean Gonvleptidae. The scanning electron micrograph was taken by Dr M. Attias in the lnstituto de Biofisic '" Carlos Chagas Filho (Universidade Federal do Rio de Janeiro). Many thanks are due to the following curators for loan of material: Prof A. Timothea da Costa (Museu Nacional da Universidade Federal do Rio de Janeiro), Dr J.L. Moreira Leme (Museu de Zoologia da Universidade de Sao Paulo), Dr M. Grasshoff (Senckenberg Museum Frankfurt am Main) and Dr H . Enghoff (Zoologisk Museum Copenhagen). The fellowship number 91/4769-6 from FAPESP (Funda~ao de Amparo a Pesquisa do Estado de Sao Paulo) is gratefully acknowledged.

REFERENCES

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