European Arachnology 2005 - European Society of Arachnology

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Discussion 1. Dermal glands 46 EUROPEAN ARACHNOLOGY 2005 Numerous dermal glands are known from a number of Opiliones. They may help to modify integumental properties. Sometimes these secretions serve for camoufl aging (e.g., Trogulidae). In Cyphophthalmi these glands are very frequent but rather inconspicuous because their openings are very tiny and usually covered by a thin fi lm of secretion. Hence they have not been often recognised (e.g., MARTENS 1979, HAMMEN 1989). In Sironidae it seems likely that the secretions help to keep the surface hydrophobic. The dermal glands correspond to the class 3 glands according to the classifi cation of NOIROT, QUENNEDY (1974). This type of glands is composed by different cells. The secretory cell is extruding its secretion products into a microvilli bordered cavity from where the secretion passes within the cuticle-lined duct towards the exterior. The similarity of the ultrastructure of these dermal glands with the glands of the male adenostyle (tarsal gland) described by MARTENS (1979) from the same species is remarkable. The occurrence of secretion in the secretory and collar cells demonstrate that both cell types have a secretory activity. Their products are released into a funnel-like cavity lined by microvilli, which are formed by the secretory and collar cells. In the secretory cells, the formation of the granules runs through different stages. Before extrusion into the secretory reservoir, the granules merge into larger granules, which are electron-lucent. The proximal beginning of the duct is provided with a distinct probably sealing structure which may be involved in the control of the expulsion of the secretions. 2. Defence glands The defence, repugnatory or scent glands are present in all Opiliones. In the Cyphophthalmi these glands are located between the second and third pair of legs on the dorsolateral side of the body on specifi c elevations, called the ozophores (JUBERTHIE 1970, GIRIBET et al. 2002). The defence glands are considered fi rst to provide chemical defence against putative predators. It is possible that their secretion is released as fi ne sprays or as a droplet (JUBERTHIE 1976, MARTENS 1978, HOLMBERG 1986). In some studies the secretions have been suggested to be used for territorial marking (JUBERTHIE 1976) or as alarm pheromones (MACHADO et al. 2002). The chemical composition of the defence secretion was known for the Laniatores and Palpatores (EKPA et al. 1985, JONES et al. 1976). In a recent study the composition of the gland secretion of Cyphophthalmus duricorius was reported by RASPOTNIG et al. (2005) for the fi rst time of a species of Cyphophthalmi. It seems evident that the defence glands are derived from dermal glands: The lid-like structure covering the opening is simply an enlarged tubercle of the surface cuticle, the body of the gland is provided with a (cuticular) intima, and the glandular units found in the secretory part correspond largely in structure with the dermal glands. Discharge of the secretion may be achieved by an increase of haemolymphic pressure. Alternatively, it could be that the gland is kept constantly under pressure and the muscles attaching to the non-secretory part may serve as dilators of this part releasing the secretions when stimulated. No sphincter muscles have been seen. Thus cuticle properties may be responsible for keeping the opening closed when undisturbed. The appearance of a huge number of mitochondria is probably evident for the high activity of the defence glands. Together with the presence of the microvilli bordered cavities in the secretory part, an effective transport of secretions may be assumed. The defence glands of Cyphophthalmi, described by JANCZYK (1956) and JUBERTHIE (1961) by light microscopy, are ultrastructurally rather similar to those of Phalangiidae (CLAWSON 1988), a family belonging to the Palpatores, a taxon regarded by GIRIBET et al. (1999, 2002) as paraphyletic.
M. Gutjahr et al.: Dermal and defence glands in C. duricorius The similarity of these glands in Cyphophthalmi and “Palpatores” may seem to support a taxon “Cyphopalpatores” as suggested by MARTENS et al. (1981) and MARTENS (1986). However, since the peculiarities of these glands of other Opiliones than Cyphophthalmi and Phalangiidae are not known, such a conclusion would be overhasty and further studies have to be awaited. Acknowledgements: We would like to thank Peter Michalik (Greifswald) and Hartmut Fischer (Greifswald) for their technical help with scanning electron microscopy. The helpful comments of the referees are also gratefully acknowledged. References Received: 02.12.2005 Accepted: 28.03.2006 CLAWSON R. C. 1988. Morphology of defence glands of the opilionids (daddy longlegs) Leiobunum vittatum and L. fl avum (Arachnida: Opiliones: Palpatores: Phalangiidae). - Journal of Morphology, 196: 363-381. EKPA O., J. W. WHEELER, J. C. COKENDOLPHER, R. M DUFFIELD 1985. Ketones and alcohols in the defensive secretion of Leiobunum townsendi Weed and a review of the known exocrine secretions of Palpatores (Arachnida: Opiliones). - Comparative Biochemistry and Physiology, 81B: 555-557. GIRIBET G., M. RAMBLA, S. CARRANZA, J. BAGUNA, M. RIUTCORT, C. RIBERA 1999. Phylogeny of the arachnid order Opiliones (Arthropoda) inferred from a combined approach of complete 18S and partial 28S ribosomal DNA sequences and morphology. - Molecular Phylogenetics and Evolution, 11: 296-307. GIRIBET G. 2000. Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). - Revista Ibérica de Aracnologíca, 2: 49-76. GIRIBET G., G. D. EDGECOMBE, W. C. WHEELER, C. BABITT 2002. Phylogeny and systematic position of Opiliones: A combined analysis of chelicerate relationships using morphological and molecular data. - Cladistics, 18: 5-70. GIRIBET G., S. L. BOYER 2002. A cladistic analysis of the cyphophthalmid genera (Opiliones; Cyphophthalmi). - Journal of Arachnology, 30: 110-128. HAMMEN L. VAN DER 1989. An Introduction to Comparative Arachnology. The Hague, SPB Publishing bv, 576 p. HOLMBERG R. G. 1986. The scent glands of Opiliones: a review of their function. - In: EBERHARD W. G., Y. D. LUBIN, B. C. ROBINSON (eds): Proceedings of the ninth international congress of arachnology, Panama 1983. Washington, Smithsonian Institution Press, 131-133. JANCZYK F. S. 1956. Anatomie von Siro duricorius im Vergleich mit anderen Opilioniden. - Sitzungsberichte der mathematisch-naturwissenschaftlichen Klasse, 165 (6-8): 475-522. JONES T. H., W. E. CONNER, A. F. KLUGE, T. EISNER, J. MEINWALD 1976. Defensive substances of opilionids. - Experientia, 32: 1234-1235. JUBERTHIE C. 1961. Structure des glandes odorantes et modalités dùtilisation de leur sécrétion chez deux Opilions Cyphophthalmes. - Bulletin de la Societé zoologique de France - Evolution et Zoologie, 86 (1): 106-116. JUBERTHIE C. 1970. Les genres dÒpiliones Sironinae. - Bulletin du Muséum national d`Histoire naturelle, 41 (6): 1371-1390. JUBERTHIE C. 1976. Chemical defense in soil Opiliones. - Revue dÈcologie et de la Biologie du Sol, 13: 155-160. KARAMAN I. M. 2005. Evidence of spermatophores in Cyphophthalmi (Arachnida, Opiliones). - Revue suisse de Zoologie, 112 (1): 3-11. MACHADO G., V. BONATO, P. S. OLIVEIRA 2002. Alarm communication: a new function for the scent-gland secretion in harvestmen (Arachnida: Opiliones). - Naturwissenschaften, 89: 357-360. MARTENS J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. - In: SENLAUB K. H.-J. HANNNEMANN, H. SCHUMANN (eds): Die Tierwelt Deutschlands, Teil 64. Jena, Gustav Fischer, 464 p. MARTENS J. 1979. Feinstruktur der Tarsal-Drüse von Siro duricorius (Joseph) (Opiliones, Sironidae). - Zoomorphology, 92: 77-93. 47
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