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Spinnerets and Silk-Producing System of Segestria Senoculata (Araneae, Araneomorphae, Segestriidae) S

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Spinnerets and Silk-Producing System of Segestria Senoculata (Araneae, Araneomorphae, Segestriidae) S Journal of Entomological and Acarological Research 2016; volume 48:5934 Spinnerets and silk-producing system of Segestria senoculata (Araneae, Araneomorphae, Segestriidae) S. Karschová, J. Hajer Department of Biology, J.E. Purkinje University, Ústí nad Labem, Czech Republic Abstract Introduction The spinning apparatus and silk of Segestria senoculata were stud- Spinning activity is a prominent feature of spiders, although there ied with the use of scanning electron microscopy and light microscopy, are also other groups of invertebrates capable of producing silk which confirmed the presence of four categories of spigots connected (Sutherland et al., 2010). The silk-spinning apparatus of spiders con- with four types of spinning glands (i.e., Glandulae ampullaceae major, sists of spinnerets and silk-spinning glands. Spinning glands, located Glandulae ampullaceae minor, Glandulae piriformes and Glandulae in the opisthosoma, are emptied through sets of outlets, known as pseudaciniformes). New data about the morphology of spinnerets and spigots or (less frequently) nozzles. Silk-producing glands originate spigots were obtained for both the adults and nymphal stages of both from ectodermal invaginations on the embryonic spinneret limb buds sexes. For the first time the silken threads of retreats, signal threads (Hilbrant & Damen, 2015). The spinning gland cells are capable of pro- and attachment discs of the members of Segestria were subjected to a ducing different types of silk onlysecretions. Spiders have a number of dif- detailed SEM study. The data resulting from studying the spinning ferent appendage types with radically different adult morphologies apparatus of S. senoculata was compared to current knowledge of the (Pechmann et al., 2010). Spinnerets are highly modified opisthosomal silk producing systems of families belonging to the Dysderoidea super- appendages (Kautzch, 1910; Yoshikura, 1955) that arise embryologi- family. Silks that are emitted from spigots in the course of retreat con- cally from the sameuse longitudinal series of limb buds (Dawydoff, 1949) struction are not (similarly to the other dysderoids) processed by the which give rise to the prosomal appendages, i.e. chelicerae, pedipalps spider’s legs during the subsequent process of hardening. Apart from and legs (Dawydoff, 1949; Whitehead & Rempel, 1959). the major ampullate glands/spigots, segestriids also possess developed Evolution of the order Araneae most likely occurred along three minor ampullate glands. Minor ampullate threads are used by S. senoc- major evolutionary lines, currently represented by the infraorders ulata spiders when making their signal threads. Mesothelae, Mygalomorphae, and Araneomorphae (Coddington & Levi, 1991). The silk-producing organs are the most remarkable organs exhibiting differences in their structures among the infraorders. The oldest known silk-producing spigots date back to the Middle Devonian of Gilboa, New York, USA (Shear et al., 1989) and their spinneret was very much like the posterior median spinnerets of the spider infra- Correspondence: Jaromír Hajer, Department of Biology, Faculty of Science, order Mesothelae (Selden et al., 2008). J.E. Purkinje University in Ústí nad Labem, eské mládeže 8, 400 96 Ústí Č The Mesothelae are the most archetypal of recent spiders known nad Labem, Czech Republic. Tel.: +420.475.283.614. (Haupt, 2003). The most thoroughly described genera are Liphistius E-mail: [email protected] Schiödte, 1849, from South and South-East Asia, and Heptathela Kishida, 1923, which includes 33 species from Japan, Vietnam and Key words: Spiders; Haplogynae; spinnerets; spigots; silk. China (World Spider Catalog, 2016). Liphistius has four pairs of spin- Non-commercialnerets: two pairs of multisegmental lateral and two pairs of monoseg- Acknowledgements: the authors gratefully acknowledge suggestions made mental median. In Heptathela kimurai the posterior and the median by the reviewers and the editor, which helped to improve the manuscript. pairs of spinnerets merge to form a single unpaired spinneret without This study was supported by the Grant Agency of J. E. Purkinje University in spinning capabilities (Haupt & Kovoor, 1993). Except for the Ústí nad Labem, Czech Republic. Mesothelae, four pairs of spinnerets are never present, since an ante- rior median pair has not developed. Received for publication: 14 April 2016. Evolution has led to great diversity in the spinning apparatus of Revision received: 15 November 2016. Accepted for publication: 16 November 2016. present day spiders seen from structural, functional, physical and chemical aspects of the silk secreted and spun by members of the var- ©Copyright S. Karschová and J. Hajer, 2016 ious families (Kovoor, 1987). Licensee PAGEPress, Italy The Araneomorphae, known as true spiders and with the largest Journal of Entomological and Acarological Research 2016; 48:5934 number of species, possess three pairs of spinnerets, namely the ante- doi:10.4081/jear.2016.5934 rior lateral (ALS), posterior median (PMS), and posterior lateral spin- nerets (PLS). In some Araneomorphae, the spinning apparatus This article is distributed under the terms of the Creative Commons Attribution Noncommercial License (by-nc 4.0) which permits any noncom- includes a functional homolog of anterior median spinnerets called the mercial use, distribution, and reproduction in any medium, provided the orig- ‘cribellum’. In other true spiders, including the Segestriidae family, inal author(s) and source are credited. the alternative homolog of anterior lateral spinnerets is a non-func- tional colulus (Foelix, 2011). [page 388] [Journal of Entomological and Acarological Research 2016; 48:5934] Article Segestriidae belongs to the haplogyne Dysderoidea superfamily, ta are, similarly to other species of the Segestriidae, usually found in silk which comprises also the Dysderidae, Oonopidae and Orsolobidae fam- tubular retreats with signal threads radiating from their entrances ilies (Coddington & Levi, 1991). Spider families referred to as the (Figure 1A-C). In the Czech Republic, these spiders are abundant under Haplogynae possess a developed so called haplogyne type of genitalia, tree barks and under stones found in forests, under stones on scree with females presenting fused copulation and fertilization ducts and slopes and also inside rock fissures (Buchar & RůŽička, 2002). The spec- male palps presenting reduction and fusion of the sclerites of the bulb imens studied for this contribution (16 females, 11 males and 6 nymphs) (Coddington, 2005). The mutual relationship of all four above men- were found and collected in the vicinity of the town of Děčín (Northern tioned families has been previously confirmed in phylogenetic studies Bohemia, Czech Republic, 50.8122325N, 14.3256342E). The specimens (Coddington, 2005; Michalik & Ramírez, 2014). The silk of all dys- were kept in tubular vials (100 mm long and 15 mm in diameter), which deroids is produced by glands called Glandulae ampullaceae, Glandulae were closed using balls of cotton wool, and also in cubic glass containers piriformes and Glandulae aciniformes (Glatz, 1972; Hajer et al., 2013; measuring 7×7×7 cm; small pieces of the moistened moss Hypnum Murphy & Roberts, 2015). The spigots which serve to emit the ampul- cupressiforme Hedw. were placed inside these breeding boxes, The body late gland silk in Dysderidae, Oonopidae and Orsolobidae are located length of adult specimens of both sexes measured up to 8-9 mm, the body individually on the spinning fields of anterior lateral spinnerets. In ref- length of nymphs collected in parental tubes was 4 mm. All of the spiders erence to their size, these glands are called major ampullate glands and were fed once a week with suitably-sized insects. The observations were the silk which is produced therein is called major ampullate or also carried out between October 2013 and February 2015. Voucher speci- dragline silk (Kovoor, 1987). The spinning fields of ALSs also contain mens were deposited at the Department of Biology, Faculty of Science, the spigots of piriform glands. J.E. Purkinje University in Ústí nad Labem, Czech Republic. Spinnerets of Segestria senoculata (Linnaeus, 1758) were first described by Apstein (1889), which found out that the spinnerets of this Microscopic and imaging equipment species are equipped with the spigots of ampullate, piriform and acini- Live spiders and their behavior during spinning activity were form glands. The same glands were described in this species by Millot observed with a stereo microscope NOVEX, which was equipped with (1931). The spinning apparatus of Segestriidae, including the histo- an INFINITY LITE Digital Camera. For a detailed study of the spinning chemical characteristics of their spinning glands, was described by Glatz apparatus, spinnerets were observed by a light microscope LEICA (1972). Piriform spigots, with low, flat basal parts, were described by DMLB, following prior tissue digestion with 20% KOH solution at Bílek (1992). Until now there are no works focusing on the SEM study of ~100°C in a glass test–tube.only Dissection for further spinneret observa- segestriid silks. The Segestriidae family is also remarkable by the fact tion was carried out on 80% ethanol-fixed specimens. The spinnerets that has developed one other pair of ampullate glands which are not were mounted in Liquido de Swan (Hrbáček, 1969), observed and found in other dysderoids (Glatz, 1972). These glands, which supply the drawn (Figures 2B and 3B,D,F) using
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