A NEW REMARKABLE TROGLOMORPHIC GONYLEPTID
BRAZIL (ARACHNIDA, OPILIONES, LANIATORES)
Abel Pérez G. & Adriano B. Kury
The new genus Giupponia is described for the new species Giupponia chagasi from two
limestone caves in Serra do Ramalho, Carinhanha, Bahia State, Brazil. This species
shows remarkable troglomorphisms, such as complete depigmentation, very elongate
appendages, very high tarsal segmentation and complete loss of the ocular structures.
This is the first anophthalmic cave dwelling harvestman from Brazil. The affinities of
Giupponia are discussed on the grounds of external and genital morphology. Placement
in the family Gonyleptidae, subfamily Pachylinae is supported by external morphology
and genital features. The new genus is compared to the other two described troglomorp-
hic gonyleptid species, Pachylospeleus strinatii Šilhavý, 1974 and Iandumoema uai
Key words: anophthalmy, Bahia state, biospeleology, Gonyleptidae, harvestmen,
Iandumoema, Neotropics, Pachylinae, Pachylospeleus, troglobites.
Giupponia gen. nov.
Giupponia chagasi sp. nov.
Un nuevo y singular Goniléptido troglomorfo de Brasil (Arachnida, Opiliones,
Se describe un nuevo género y especie, Giupponia chagasi, proveniente de una cueva
calcárea localizada en la Serra do Ramalho, Carinhanha, Estado de Bahía, Brasil. La
especie presenta notables troglomorfismos tales como una fuerte despigmentación,
apéndices muy alargados, alta segmentación tarsal y pérdida completa de las
estructuras oculares. Este es el primer troglobio anoftalmo de Brasil. Se discuten las
afinidades de Giupponia sobre la base de su morfología externa y genital. Los
resultados obtenidos ubican a la nueva especie en la família Gonyleptidae, subfamilia
Pachylinae. Giupponia chagasi es comparada con las otras dos especies conocidas de
goniléptidos troglobios: Pachylospeleus strinatii Šilhavý, 1974 e Iandumoema uai Pinto-
Palabras clave: anoftalmia, Bioespeleología, Gonyleptidae, Opiliones, Iandumoema,
Neotrópico, Pachylinae, Pachylospeleus, Troglobios, Brasil.
Giupponia gen. nov.
Giupponia chagasi sp. nov
In the family Gonyleptidae there are only a few cave-dwelling species which
developed troglomorphisms. The monotypic subfamily Pachylospeleinae was
created for the genus Pachylospeleus Šilhavý, 1974 with the single species
Pachylospeleus strinatii Šilhavý 1974 from caves in São Paulo state (Šilhavý,
1974). The genus Iandumoema Pinto-da-Rocha, 1996 was created for the species
Iandumoema uai Pinto-da-Rocha, 1996 from the karst of Minas Gerais (Pinto-da-
Rocha, 1996). Both species are somewhat depigmented and possess elongate
appendages, but still retain some of the pigmented layer and the lens of the eye. An
expedition of the Arachnology Laboratory of the Museu Nacional in the Bahia State
(eastern Brazil) yielded some specimens of an undescribed depigmented species
with extremely elongate appendages and which lacks eyes altogether. This is the
first anophthalm harvestman from Brazilian caves, although not the first anophthalm
from Brazil — there is a eyeless small species belonging to an unknown family
(traditionally included in the meaningless family Phalangodidae), which has been
described from termite nests in the state of Rio de Janeiro — Caecobunus
termitarum Roewer, 1927. The new blind cave dwelling gonyleptid is herein
described as a new species in a new genus, and its subfamilial inclusion is
A new remarkable troglomorphic
gonyleptid from Brazil
Abel Pérez G.
& Adriano B. Kury
Quinta da Boa Vista,
São Cristóvão 20940-040,
Rio de Janeiro, RJ, Brasil
Revista Ibérica de Aracnología
ISSN: 1576 - 9518.
Dep. Legal: Z-2656-2000.
Vol. 5, 31-VII-2002
Sección: Artículos y Notas.
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44 Abel Pérez G. & Adriano B. Kury
Acronyms of repositories are AMNH (American
Museum of Natural History, New York), CBK (Colec-
ción Biopeleológica de la Sociedad Espeleológica de
Cuba, Havana), CAS (California Academy of Sciences,
San Francisco), IBUT (Instituto Butantan, São Paulo),
MNRJ (Museu Nacional, UFRJ, Rio de Janeiro), MZSP
(Museu de Zoologia, USP, São Paulo) and SMF (Senc-
kenberg Museum, Frankfurt). All measurements are in
Additional material examined: Iandumoema uai
Pinto-da-Rocha, 1996, 1
1 (MNRJ), Pachylospeleus
strinatii Šilhavý, 1974, 2
1 (MNRJ 6790).
Giupponia new genus
TYMOLOGY. Genus name honors the arachnologist
Alessandro Ponce de Leão Giupponi, who not only
collected the holotype of Giupponia chagasi, but also
has been seriously involved in building up the Arachno-
logical Section of the Museu Nacional through his
devotion and friendship.
YPE SPECIES. Giupponia chagasi new species, by
IAGNOSIS. Gonyleptidae without eyes, eye mound
projected in high erect spine. Coxa IV in male armed
with dorso-apical trifid small apophysis. Frontal hump
with paired spines. Area I undivided. All scutal areas,
free tergites and sternites unarmed. Tarsus III-IV
without tarsal process. Femur IV without apophysis and
spines. Ventral plate with parabolic cleft and four basal
flattened truncated setae and three distal cylindrical
pointed setae. Glans without dorsal and ventral proces-
ses, partially fused to ventral plate. Relative length of
legs: II-IV-III-I. Tarsal segmentation male 8-9(3)/34-
37(4)/7/8, female 8-9(3)/29-36(4-5)/7/8. Calcaneus I
more than twice longer than astragalus, calcanei II-III a
little shorter, calcaneus IV much shorter than astragalus.
Giupponia chagasi new species
TYMOLOGY. Species name honours the other collector
of the holotype of Giupponia chagasi, the myriapodolo-
gist Amazonas Chagas Junior who is another enthusias-
tic member of our team.
YPE MATERIAL. holotype (MNRJ 4640), Carinhanha,
Serra do Ramalho, Lapa do Boqueirão, 29.vi.2001, leg.
A. Giupponi & A. Chagas; 1
8 paratypes (MNRJ
4699), Carinhanha, Serra do Ramalho, Gruta do Zé
Bastos, 28.vi.2001, leg. A. Giupponi, A. Chagas & R.
paratype each, same data (AMNH, CAS,
IBUT, MZSP, CBK, SMF).
ISTRIBUTION. Species known from two caves in the
Serra do Ramalho, municipality of Carinhanha, Bahia
State, Brazil (see Fig 12).
ESCRIPTION. Male holotype measurements. Carapace
1.51 long 1.81 wide, abdominal scutum 2.22 long 2.44
wide. Pedipalp: Fe 3.62, Pa 1.69, Ti 2.04, Ta 1.42, claw
Dorsum. — Scutum outline attenuate pyriform, elonga-
te, widest at area II. Cheliceral sockets shallow, frontal
margin of carapace with one projections in each of the
lateral thirds. Frontal hump of carapace armed with a
pair of pointed spines. Eye mound narrow, situated far
from frontal margin of carapace, projected in a very
high erect process. Without any trace of eyes. Mesoter-
gum divided in four areas by four transverse deep wide
grooves. Area I undivided. Each area with a transverse
row of low pointed tubercles. Posterior margin and free
tergites I-III also with a tubercle transverse row each.
Lateral margins with a few small tubercles. Two ozopo-
res of similar size present, only the anterior visible in
Venter. — Coxae I-IV and stigmatic area densely
covered with tubercles. Stigmatic area partially fused
with coxa IV. Genital opercle rounded, subquadrangu-
lar. Coxa IV without ventral apophyses, connected to
posterior margin of stigmatic area by a lateral process.
Stigmata visible. Stigmatic area with posterior groove
and thickened posterior border. All free sternites with a
transverse tubercle row each.
Chelicera. — Basichelicerite (trochanter) as long as
carapace, unarmed, without swollen bulla. Hand slender
and unarmed. Fixed and movable finger with 4 low and
Pedipalp. — Measurements of segments: Tr 0.5, Fe 3.4,
Pa 1.3, Ti 2.2, Ta 1.4, claw 1.0. Twice longer than body,
all segments much elongate and slender, with spination
concentrated in tibia-tarsus. Coxa long, reaching half of
basichelicerite in situ. Trochanter with ventral setiferous
tubercle. Femur smooth and unarmed, except for ventro-
basal setiferous tubercle. Patella unarmed, much elonga-
te and slender, only slightly thickened distally. Tibia
cylindrical, with five ectal (IiiIi) and five mesal (IiiIi)
setiferous tubercles. Tarsus slightly convex ventrally,
armed with three ectal (iII) and three mesal (IiI) setife-
rous tubercles and many ventral bristles.
Legs. — Dorso-basal apophyses of coxae I-II short,
simple. Coxa IV surpassing scutum in dorsal view, its
posterior border reaching groove IV and covered with
dorsal setiferous tubercles and dorso-apical trifid blunt
apophysis. Trochanter IV short with low tubercles.
Femur IV very long, only slightly sinuous, densely
covered with small sharp tubercles. Tarsus III-IV
without tarsal process (“pseudonychium”). Tarsal
counts 9(3)/34-37(4)/7/8. Ratio calcaneus/astragalus of
legs I-IV: 2.6/0.9/0.8/0.3. Leg measurements in Table I.
Genitalia. — Ventral plate subrectangular, a little
narrower in the middle, lateral margins much thinner
and rugose, distal margin with parabolic cleft. Setae
arranged in two groups. Basal group with four flattened
large truncate setae. Distal group with three cylindrical
pointed smaller setae. Glans without dorsal or ventral
A new troglomorphic Gonyleptidae 45
Figs. 1-2. Giupponia chagasi species nova, male holotype (MNRJ 4640). 1. Habitus, dorsal with detail of scutum
and tergites; 2. Same, more general view. Scale bars: 1 mm.
Measurements of long podomeres of legs of the
holotype of Giupponia chagasi n. sp.
Total length of leg
19.3 37.9 20.9 27.7
4.9 9.3 6.0 7.8
3.6 8.1 3.4 5.1
6.5 8.8 6.1 8.3
46 Abel Pérez G. & Adriano B. Kury
Figs. 3-4. Giupponia chagasi species nova, male holotype (MNRJ 4640). 3. Body, lateral view; 4. Coxae, stigmatic
area and sternites, ventral view. Scale bars: 1 mm.
A new troglomorphic Gonyleptidae 47
Figs 5-9. Giupponia chagasi species nova, male holotype (MNRJ 4640). 5. Right chelicera, ectal view, 6. Same,
frontal view; 7. Same, mesal view; 8. Right pedipalp, ectal view; 9. Same, mesal view. Scale bars: 1 mm.
48 Abel Pérez G. & Adriano B. Kury
Figs. 10-11. Giupponia chagasi species nova, male holotype (MNRJ 4640). 10. Distal part of Penis, dorsal view;
11. Same, lateral view. Scale bar: 0.1 mm.
processes, fused with ventral plate until mid-length.
Stylus thick and with apex swollen.
Female. — Much similar to male, excepted dorso-apical
apophysis of coxa IV, reduced to a little spiniform
tubercle. Tarsal counts (n=14) 8-9(3)/29-36(4-5)/7/8.
Ratio calcaneus/astragalus of legs I-IV: 2.0/0.8/0.8/0.2.
ATURAL HISTORY. The specimens of Giupponia
chagasi n. sp. have been collected in the deeper part of
both caves, in absolute darkness, high relative humidity
and far from the entrances. This results in high stability
of conditions of humidity and temperature of the
environment. The holotype has been collected more
than 800 m from the entrance of the Cave of Boqueirão
and the paratypes from 200 m to the deepest explored
zones of the Cave of Zé Bastos (700 m). The specimens
were found alone, usually hidden in small crevices on
the walls, a few were caught on the floor, under rocks
or fallen secondary formations (stalactites and stalagmi-
tes). In the Cave of Zé Bastos some specimens inhabi-
ted semi-flooded galleries, and placed themselves on the
mud-covered walls, near the waterline. The galleries
inhabited by individuals of this species show low
availability of food, namely, the absence of bat guano
and of the associated fauna, and the great distance from
the entrance (little influence from the external environ-
ment). Some specimens of Giupponia chagasi n. sp.
have been collected together with a new troglomorphic
species of Charinus (Arachnida, Amblypygi, Charini-
dae). Other associated fauna were roaches and spiders
of the family Ctenidae (R. Baptista & A Giupponi, pers.
Discussion — relationships
The relationships of Giupponia are discussed next. The
position of glans, free, based on an unfoldable sac, not
encapsulated in a concavity in the truncus define its
pertinence to the superfamily Gonyleptoidea (Kury,
Let us first review the absence of features typical
of each Gonyleptoid family and then examine the
positive evidence to include it the Gonyleptidae. Among
the families of Gonyleptoidea, the Stygnopsidae and
Agoristenidae are easily dismissed respectively because
lack of a spiny sac and trifid spines of ventral plate plus
keeled stylus. Giupponia also lacks the typical flattened
pedipalps which are a remarkable synapomorphy of the
Cosmetidae. The Assamiidae, besides the frontal
processes of carapace and the specialized depressed
pedipalps possess the introverted glans (Stacheltrichter).
The Stygnidae do not possess a common eye mound and
show specializations in ventral plate. The Manaosbiidae
have a spindle-like swollen basitarsus plus typical
armature of mesotergum. The Cranaidae possess patella
and tibia of pedipalpus covered with coarse granula-
tions. The absence of sexual dimorphism of chelicerae
is typical of Gonyleptidae and Manaosbiidae. The loss
of a dorsal process of the glans penis appear to have
occurred many times independently in the families of
Gonyleptoidea, and its absence per se is not enough to
place this genus in a given family. The short rectangular
ventral plate with a parabolic cleft, in the same plan as
the truncus, with two unequal groups of setae is typical
of Gonyleptidae. This construction plan is present also
A new troglomorphic Gonyleptidae 49
Fig. 12. South America, showing location of Serra do
Ramalho (open circle), type locality of Giupponia chagasi
species nova, on the left margin of São Francisco River.
in early derivative Cranaidae such as Prostygnus,
who still possess although a dorsal process in the
glans. This description could also apply to the
Cosmetidae, which do not have this cleft. A typical
feature of the Gonyleptidae, however, is absent —
the flabelliform dorsal process of glans. This
structure appeared only in the “advanced” Gonylep-
tidae (cf. Kury, 1994). Other positive evidence to
include this genus in the Gonyleptidae is the presen-
ce of two ozopores, feature exclusive (although not
universal) of this family and the bifid (or trifid)
apical apophysis of coxa IV (single branched, when
present, in the other families). Therefore, we have
at least three positive evidences favoring the inclu-
sion in Gonyleptidae, and lots of reasons not to
include it in any other family of Gonyleptoidea.
Having established our hypothesis of Giuppo-
nia as a member of the Gonyleptidae, now comes
the problem of its subfamilial inclusion. The outer
features are so much cave-stereotyped that become
of little use. The unfused free tergites discard at
once the Heteropachylinae for our sampling univer-
se. Mesotergal area II not invading area I could
serve to discard the Goniosomatinae. The Gonyas-
samiinae are minute specialized Pachylinae with
hooked eye mound and sturdy spiny stylus, features
that do not match Giupponia.
The eye mound removed from the anterior
border of carapace with a single huge spine is not
typical of most subfamilies of Gonyleptidae. The
eye mound in Gonyleptidae is very plastic, but such a
position and armature occur in many genera of Pachyli-
nae and Tricommatinae (who could be an offshoot of
the former). The Tricommatinae typically show a fusion
of left and right halves of mesotergal area I, swollen
base of ventral plate and a lamina parva.
Male genitalia is the best basis for speculating
subfamilial pertinence. The complete absence of spiny
sacs and slender spines in ventral plate discards the
pertinence to the Metasarcinae. The absence of helycoi-
dal setae of distal group in ventral plate and the arrange-
ment of the basal group in the same plan as ventral plate
speak against the inclusion in the Gonyleptinae-like
group (that is, Gonyleptinae, Hernandariinae, Progony-
leptoidellinae, Sodreaninae, Caelopyginae). The Mito-
batinae and Bourguyiinae among other things always
possess a flabelliform ventral process of glans, absent
in Giupponia. The Pachylospeleinae, a poorly defined
subfamily in respect to Pachylinae, possess a ventral
process of glans.
The Pachylinae is by far the largest subfamily of
Gonyleptidae, including half of the valid species (400
out of 821). They surely do not form a monophyletic
group, and actually assigning a given genus to the
Pachylinae is almost the same as stating “Gonyleptidae
incertae sedis”. In accordance to Giupponia, many
genera of Pachylinae possess a naked stylus, without
ventral or dorsal processes and a subrectangular ventral
Discussion — comparisons with the other
There are two other species of troglomorphic Gonylepti-
dae, namely Pachylospeleus strinatii, type of the
monotypic subfamily Pachylospeleinae and Iandumoe-
ma uai, placed in Pachylinae. Both are roughly similar
in habitus, to the point that Helia Soares labelled a
specimen of I. uai as a new species of Pachylospeleus.
We had opportunity to examine males and females
of both species. The most striking resemblance between
both species is the scutum outline, pedipalps, general
structure of ventral plate and the presence of a ventral
process in glans. There are many significant differences,
such as 1) the trochanter IV of male, which in P.
strinatii is short, and in I. uai is long, hourglass-shaped;
2) P. strinatii possesses apical inner apophysis in coxa
IV, absent in I. uai; 3) the apophysis of coxa IV of male
in P. strinatii is slightly sinuous with a secondary small
branch while in I. uai it is abruptly bent without secon-
dary branch; 4) tarsal process absent in I. uai, present in
P. strinatii. Minor differences are for example the eye
mound, which in P. strinatii is unarmed and in I. uai is
armed with an erect very high spine (see also Table II).
Giupponia is neither specially closely related to
Pachylospeleus nor to Iandumoema. A synoptic scheme
(Table II) will help separate those three species at a
50 Abel Pérez G. & Adriano B. Kury
Comparison among troglomorphic Gonyleptidae.
Giupponia Iandumoema Pachylospeleus
with pair of spines 2 small pointed tubercles unarmed
armed with huge erect spine. armed with huge erect spine,
with apex bent backwards.
2 paramedian small setiferous
tubercles located on the poste-
rior slope of eye mound.
completely absent cornea reduced to 50% size of
epigean Pachylinae (Eusarcus
sp.) of similar size. Pigment
cornea reduced to 30% size of
epigean Pachylinae. Pigment
much reduced (literature data
conflicting: Šilhavý, 1974: much
reduced, but Pinto-da-Rocha,
extremely elongate slender, but short slender, but short
apophysis of coxa IV
Blunt, very short and trifid. Sharp, very long, abruptly bent
Sharp, very long, slightly sinuous
Coxa IV ventral
with lateral-inner apophysis
fused to stigmatic area.
without apophysis with strong inner lateral pointed
short, unarmed long, hourglass-shaped, with
strong wide basal prolateral
short, square, armed with strong
dorso-apical pointed apophysis
Femur IV of
straight curved basally straight, with dorso-basal
absent absent present
8-9(3)/29-37(4-5)/7/8 7(3)/16-22(3-4)/6/6 6(3)/15-16(5)/6/6
Ventral process of
absent present present
subrectangular, with parabolic
trapezoid, entire trapezoid, entire
Two caves in Bahia State Cave in Minas Gerais State Cave in São Paulo State
The cave-dwelling species herein described — Giuppo-
nia chagasi — is the first entirely anophthalm member
of Gonyleptidae — the largest family of Laniatores.
Besides the eyelessness, G. chagasi shows the most
advanced troglomorphisms ever recorder for this family,
such as, high number of tarsal segments, extremely long
pedipalps and legs, and high degree of depigmentation.
Its inclusion in any of the 15 currently recognized
subfamilies of Gonyleptidae is not positively supported
by any evidence, although the typological combination
of characters offers grounds for the inclusion in the
meaningless subfamily Pachylinae. G. chagasi up to
now is known from only from two caves, and this high
local endemism of this species provides support for the
conservation of the karstic area of Carinhanha in Bahia
This study has been supported by grant # 300175/94-9
from the Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) to ABK and a scho-
larship from the program CAPES/CNPq – IEL Nacional
– Brazil to APG. Prof. José Antonio Alves de Azevedo
and Jelcy Corrêa the Secretary of Education and Sports
of the Municipality of Petrópolis made possible a
survey in the Bahia caves. We thank Simone Mesquita,
head of the Central Laboratory of the Museu Nacional
and Fundação Universitária José Bonifácio (FUJB) for
multiple facilities. We are immensely indebted to our
colleagues Dr Renner Baptista, Alessandro Giupponi
and Amazonas Chagas Jr, who are always contributing
immensely with the study of Arachnids in the Museu
KURY, A. B. 1994. Early lineages of Gonyleptidae (Arachnida, Opiliones, Laniatores). Tropical Zoology, 7(2): 343-
INTO-DA-ROCHA, R. 1996. Iandumoema uai, a new genus and species of troglobitic harvestman from Brazil
(Arachnida, Opiliones, Gonyleptidae). Revta bras. Zool., 13(4): 843-848.
ILHAVÝ, V. 1974. A new subfamily of Gonyleptidae from Brazilian caves, Pachylospeleinae subfam. n. (Op. Gon.).
Rev. suisse Zool., 81(4): 893-898.
KURY, A. B. 1993. Análise filogenética de Gonyleptoidea (Arachnida, Opiliones, Laniatores). Unpublished Ph.
D. Thesis. Universidade de São Paulo.