Podoctidae — version 1€¦ · 1 Podoctidae — version 1.6 Adriano B. Kury Departamento de Invertebrados, Museu Nacional/UFRJ Quinta da Boa Vista, São Cristóvão, 20.940-040,

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Podoctidae — version 1.6

Adriano B. Kury

Departamento de Invertebrados, Museu Nacional/UFRJ

Quinta da Boa Vista, São Cristóvão, 20.940-040, Rio de Janeiro - RJ – BRAZIL

1. Introduction

Podoctidae is a medium-sized family with 120 species of medium-sized Laniatores.

They are usually heavily spined and variedly colored, with contrasting dark brown/pale

yellow, reddish brown/white or entirely intense green. Typical features are the ventral and

dorsal rows of spines on femur I of both sexes, the pairs of tubercles linking eyes, eye

mound and/or scutal areas and the extremely varied shaped of eye (or interocular) mound.

They typically occur in SE Asia and Australasia, but there are also representatives in other

continents. Descriptions in literature typically fail to correctly recognize sex of individuals,

because sexual dimorphism (which includes length and built of pedipalps, armature of

chelicerae and structure of interocular mound) is not yet understood. Boundaries of the

three subfamilies are meaningless. It is one of the most undersampled families of

Laniatores. Number of known species is likely to double in the next decades. The number

of genera, in contrast, will surely drop as soon a review starts. If it appears to the reader

that 60 genera are too much for 120 species (average only two species for genus), a

noteworthy remark is that there are nothing less than 32 generic names currently held as

junior synonyms. Nothing remotely similar to a phylogenetic analysis has ever been

published.

1.1. Subtaxa included.

3 subfamilies; 58 genera and 120 species. Subfamilies are traditionally separated by

meaningless features such as number of segments of distitarsus I and absence or presence

of scopula in posterior tarsi.

Included genera

Erecananinae: Erecanana Strand, 1911; Iyonus Suzuki, 1964; Lomanius Roewer, 1923.

Ibaloniinae: Asproleria Roewer, 1949; Austribalonius Forster, 1955; Bonea Roewer, 1913;

Eusitalces Roewer, 1915; Gargenna Roewer, 1949; Heteroibalonius G&G, 1947;

Heteropodoctis Roewer, 1911; Holozoster Loman, 1902; Ibalonianus Roewer, 1923;

Ibalonius Karsch, 1880; Ibantila Šilhavý, 1969; Leytpodoctis Martens, 1993; Mesoceratula

Roewer, 1949; Metibalonius Roewer, 1912; Orobunus G&G, 1947; Paramesoceras Roewer,

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1915; Pentacros Roewer, 1949; Podoctinus Roewer, 1923; Proholozoster Roewer, 1915;

Santobius Roewer, 1949 and Sitalcicus Roewer, 1923.

Podoctinae: Baramella Roewer, 1949, Baramia Hirst, 1912, Baso Roewer, 1923, Basoides

Roewer, 1949, Bistota Roewer, 1927, Centrobunus Loman, 1902, Dino Loman, 1892,

Dongmoa Roewer, 1927, Eupodoctis Roewer, 1923, Eurytromma Roewer, 1919, Gaditusa

Roewer, 1949, Hoplodino Roewer, 1915, Idjena Roewer, 1927, Idzubius Roewer, 1949,

Japetus Roewer, 1949, Laponcea Roewer, 1936, Lejokus Roewer, 1949, Lundulla Roewer,

1927, Metapodoctis Roewer, 1915, Neopodoctis Roewer, 1912, Oppodoctis Roewer, 1927,

Peromona Roewer, 1949, Podoctellus Roewer, 1949, Podoctis Thorell, 1890, Podoctomma

Roewer, 1949, Podoctops Roewer, 1949, Pumbaraius Roewer, 1927, Sibolgia Roewer,

1923, Stobitus Roewer, 1949, Tandikudius Roewer, 1929, Trencona Roewer, 1949,

Trigonobunus Loman, 1894, Tryssetus Roewer, 1936 and Vandaravua Roewer, 1929.

1.2. Systematic historical background

The three subfamilies currently included in Podoctidae were created by Roewer

(1912) as subfamilies of his huge family Phalangodidae. The first author to sort them out

forming a separate family was Mello-Leitão (1938). The vast majority of genera and

species have been created by Roewer (e. g. 1912; 1923 and 1949), and that caused

systematics of this family to be in a very bad shape. Suzuki (e. g. 1964a-b; 1969; 1972;

1977) finely described and redescribed some species from SE Asia, and started to

synonymize some of those countless monotypic genera and Rambla (1984) did the same

with the Podoctidae of the Seychelles, but this is not enough to overturn the balance.

There is still much to be done on the Podoctidae.

The family Podoctidae as currently understood began to be widely accepted around

mid-seventies (e. g. Šilhavý, 1973 maybe the turning point; Suzuki 1977; Shear, 1982;

Martens, 1986) but in intermediate period papers such as Kratochvíl (1958) Briggs (1969),

Šilhavý (1961) and Suzuki (1964a-b; 1972) for example it is still considered to be part of

Phalangodidae. Staręga (1992) considered Erecananinae a separate family, but he still

owes us some explanation for that. I started to build a matrix for a phylogenetic analysis in

spite of the inadequacy of data available from literature. It is yet early to understand the

basic branching of the subgroups, but none of Roewer’s subfamilies ever emerges as a

clade from preliminary runs.

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1.3. Natural history

Little has been published on the natural history of this family. A few species from

India and the Philippines have been reported to carry a layer of small earth particles

adhered to the cuticle (Roewer, 1929; Martens, 1993). Collection labels and a few

literature reports (e. g. Suzuki, 1977) indicate that some species were collected by beating

shrubs and low trees, while others were found under logs as usual in Laniatores.

An unique feature among Opiliones is that males carry the eggs on their femora. It

has been reported for one single species from the Philippines, Leytpodoctis oviger

(Martens, 1993), but I observed at least one more species with this habit. This is an

undescribed species from Melanesia, and it is not specially closely related to Leytpodoctis.

This suggests that the strategy of carrying eggs on femora of male is much more

widespread within the family than we now know.

Species have been reported from caves, but without troglomorphisms and without

exclusivity in the hypogean environment.

2. Characterization

SIZE. Medium-sized Laniatores, body length 2.5 to 5 mm. Legs I-IV much variable

3-10/10-30/7-21/9-28 mm long. DORSUM. Dorsal scute (Figs 1-3) outline can be ovoid or

subtrapezoidal, in general without strong constrictions and carapace is not much narrower

than abdominal scute. Mesotergum usually clearly divided into areas by grooves, the areas

may be connected by tubercular bridges. No areas fused. Armature of areas and tergites

highly variable, maybe either completely smooth and unarmed or armed with strong

spines, which bear subapical setae. In some species the scutum is profusely covered with

warts, wrinkles, granules and tubercles. Common eye mound (or interocular mound) not

always present, it may be very high and densely granulous, bearing a diversity of spines

and with strong sexual dimorphism (Figs 2-3). Sometimes eyes are wide apart in separate

individual mounds, always linked to the frontal part of carapace by tubercular bridges

(which may be a synapomorphy for the family). CHELICERA. Cheliceral hands usually not

swollen, but basichelicerite often shows rich ornamentation and sexual dimorphism in

length and armature (Figs 4-6). PEDIPALP. Pedipalps with ventral row of spines in femur

and ventro-mesal and ventro-ectal spines in patella-tibia-tarsus. Tibia and tarsus do not

form a subchela. In many genera they are sexually dimorphic, basally clavate in male

and/or with clusters of divergent spines (Fig 9-10), in another group males have

immensely elongate segments (Figs 7-8). LEGS. Legs usually long and straight, often

covered with rows of pointed tubercles, specially leg I which in most species has powerful

ventral and dorsal rows of setiferous spines in both sexes (Fig 11). Distitarsus I 1-2

jointed. Distitarsus II 1-4 jointed. COLOR. Color background usually brown to yellow, some

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species are deep green, legs may be ringed in black and yellow, scutal spines may be black

sharply contrasting with background. GENITALIA. Genitalia is unique (Fig 12), dorsal plate

is deeply cleft and there is an inflatable sac which inflates and exposes the stylus, flanked

by two powerful prongs (Figs 13-15). SEXUAL DIMORPHISM. Sexual dimorphism is shown

in basichelicerite which is very long and armed in male, in pedipalp, which in male may be

thickened at base with a cluster of divergent spines or extremely elongate while the female

pedipalp is short. Also shown in eye mound which in male can be wider, or much more

elaborate in ornamentation and leaned back against the scute (Fig 1).

3. Distribution

The peak diversity of the family is in Southeast Asia, specially in Papua New Guinea.

They also occur in Melanesia, Micronesia, Japan, India, Sri Lanka, Madagascar, Seychelles,

Mauritius and Central Africa. A single species is known from Australia. There is a sole

species – Ibantila cubana Šilhavý, 1969 – reported from the New World (Šilhavý, 1969),

introduced (information not given in the original description) in a botanical garden in Cuba

(A. Pérez, pers. comm.). Brasiloctis bucki, the Brazilian “Podoctidae” of Mello-Leitão (1938)

was later referred to the Triaenonychidae (see Soares & Soares, 1979).

4. Relationships

This is still obscure. They belong without doubt to the Grassatores, and it is possible

that they are most closely related to the Biantoidea, although there is no obvious

relationship with any other family.

5. References

Briggs, T. S., 1969. A new holarctic family of laniatorid Phalangids (Opiliones). Pan-Pacif.

Ent., 45: 35-50.

Goodnight, J. C. & M. L. Goodnight, 1957. Opiliones. Insects of Micronesia. B. P. Bishop

Mus., 3: 71-83.

Kratochvíl, J., 1958. Die Höhlenweberknechte Bulgariens (Cyphophthalmi und Laniatores).

Práce Brn. zákl. cesk. Akad. Véd., 30(375): 372-396.

Martens, J., 1986. Die Grossgliederung der Opiliones und die Evolution der Ordnung

(Arachnida). Actas 10 Congr. Aracnol. Jaca España, 1986,1: 289-310.

Martens, J. 1993. Further cases of paternal care in Opiliones (Arachnida). Tropical Zoology

6: 97-107.

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Mello-Leitão, C. F. DE, 1938. Considerações sobre os Phalangodoidea Soer. com descrição

de novas formas. Anais Acad. bras. Cienc. 10(2): 135-145, 1 pl.

Rambla, M., 1984. Contributions a l'étude de la faune terrestre des îles granitiques de

l'archipel des Sechelles (Mission P.L.G. Benoit - J.J van Mol 1972). Opiliones

(Arachnida). Ann. K. Mus. Midd-Afrika Zool. Wet., 242: 1-86, 3 lam.

Roewer, C. F., 1912. Die Familien der Assamiiden und Phalangodiden der Opiliones-

Laniatores (= Assamiden, Dampetriden, Phalangodiden, Epedaniden, Biantiden,

Zalmoxiden, Samoiden, Palpipediden anderer Autoren). Arch. Naturgesch., 78A(3):

1-242.

Roewer, C. F., 1923. Die Weberknechte der Erde. Sistematische Bearbeitung der bisher

bekannten Opiliones. 1116 pp., figs 1-1212, Gustav Fischer, Jena.

Roewer, C. F., 1929. Süd-indische Skorpione, Cheloneti und Opilioniden. Rev. suisse Zool.,

36(21): 609-639.

Roewer, C. F., 1949. Weitere Weberknechte XIV. Über Phalangodidae II. Senckenbergiana,

30: 247-289.

Shear, W. A., 1982. Opiliones: 104-110. In: Parker, S.P. (ed.) Synopsis and classification

of living organisms. Vol 2. McGraw Hill Book Co, New York, pp 1232.

Šilhavý, V., 1961. Die Grundsätze der modernen Weberknechttaxonomie und Revision des

bisherigen Systems der Opilioniden. Verh. 11. intern. Kongr. Ent. Wien, 1960, 1:

262-267.

Šilhavý, V., 1969. Ibantila cubana gen. nov. spec. nov., the first representative of

subfamily Ibaloniinae Roewer (Arachn, Opilionoidea) from America. Acta soc. zool.

Bohemoslov., 33(4): 372-376.

Šilhavý, V., 1973. Two new systematic groups of gonyleptomorphid phalangids from the

Antillean-Caribbean Region. Agoristenidae fam. n. and Caribbiantinae subfam. n.

Vestnik Ceskosl. spolec. zool., 37(2): 110-143.

Soares, H. E. M. & B. A. M. Soares, 1979. Opera Opiliologica Varia XVI. Novo gênero de

Gonyleptidae e presença de Triaenonychidae no Brasil (Opiliones). Revta bras. Ent.,

23(3): 169-173.

Staręga, W., 1992. An annotated check-list of harvestmen, excluding Phalangiidae, of the

Afrotropical Region (Opiliones). Ann. Natal Mus., 33(2): 271-336.

Suzuki, S., 1964a. A remarkable new phalangodid Dongmoa oshimensis from Japan.

Annot. zool. jap., 37: 163-167.

Suzuki, S., 1964b. A new representative of the Erecananinae from Japan. Annot. zool. jap.,

37: 221-225, 7 figs.

Suzuki, S., 1969. On a collection of opilionids from South East Asia. J. Sci. Hiroshima Univ.,

(B1) 22 (2): 11-77.

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Suzuki, S., 1972. Opiliones of Semangkok Forest Reserve, Malaysia. J. Sci. Hiroshima Univ.

B1, 24(1): 1-37, 94 figs.

Suzuki, S., 1977. Report on a collection of opilionids from the Philippines. J. Sci. Hiroshima

Univ., B1, 27(1): 1-120.

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Captions for illustrations:

Figs 1-3. Lomanius longipalpus mindanaoensis Suzuki, 1977 from the Philippines. Habitus.

1. Male, lateral view. 2. Female, dorsal view. 3. Male, dorsal view. (From Suzuki,

1977).

Figs 4-6. Hoplodino hoogstraali Suzuki, 1977 from the Philippines. 4. Female chelicera,

ectal view. 5. Male chelicera, mesal view. 6. Same, ectal view. To the same scale.

(From Suzuki, 1977).

Figs 7-8. Lomanius longipalpus mindanaoensis Suzuki, 1977 from the Philippines. 7. Left

pedipalpus of male, ectal view. 8. Left pedipalpus of female, ectal view. (From

Suzuki, 1977).

Figs 9-10. Hoplodino hoogstraali Suzuki, 1977 from the Philippines. 9. Left pedipalpus of

male, mesal view. 10. Left pedipalpus of female, ectal view. (From Suzuki, 1977).

Fig 11. Lomanius longipalpus mindanaoensis Suzuki, 1977 from the Philippines. Left leg I

of male, lateral view. (From Suzuki, 1977).

Fig 12. Heteropodoctis quinquespinosus (Roewer 1911) from Papua New Guinea (CAS).

Distal part of penis, dorsal view. Scale bar = 150 µm. Photo Darrell Ubick.

Figs 13-15. Unspecified podoctid, schematic view of penis. 13. Unexpanded, lateral view.

14. Unexpanded, dorsal view. 15. Expanded, lateral view. (From Martens, 1986).

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