The Spider Chelicerae: Some Problems of Origin and Evolution
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EUROPEAN ARACHNOLOGY 2003 (LOGUNOV D.V. & PENNEY D. eds.), pp. 349366. © ARTHROPODA SELECTA (Special Issue No.1, 2004). ISSN 0136-006X (Proceedings of the 21st European Colloquium of Arachnology, St.-Petersburg, 49 August 2003) The spider chelicerae: some problems of origin and evolution Õåëèöåðû ïàóêîâ: ïðîáëåìû ïðîèñõîæäåíèÿ è ýâîëþöèè S.L. ZONSTEIN Ñ.Ë. ÇÎÍØÒÅÉÍ Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel-Aviv University, 69978 Tel-Aviv, Israel. email: [email protected] Îòäåë çîîëîãèè, Ôàêóëüòåò åñòåñòâåííûõ íàóê èì. Äæîðæ Ñ. Âèçå, Òåëü-Àâèâñêèé óíèâåðñèòåò, Òåëü- Àâèâ, Èçðàèëü. email: [email protected] ABSTRACT. Some problems of the origin and evolution of the chelicerae in spiders are discussed. The enlargement and bending of the basal cheliceral segments combined with the displacement of the fang insertion point ventrad is recognized as the principal cheliceral modification observed in the Araneae. The chelicerae of the Liphistiidae and the Mygalomorphae are regarded as plesiomor- phic in their position and axial orientation, but as apomorphic by their configuration; the latter seems to be an evolutionary compromise caused by burrowing activity. The possibility of an origin of the labidognathous cheliceral construction from the specialized orthognathous chelicerae of liphistiids and mygalomorphs is regarded as improbable. ÐÅÇÞÌÅ.  ñòàòüå îáñóæäàþòñÿ âîïðîñû ïðîèñõîæäåíèÿ è ýâîëþöèè õåëèöåð ó ïàóêîâ. Îñíîâíûìè ìîäèôèêàöèÿìè õåëèöåð, íàáëþäàåìûìè ó îðòîãíàòíûõ ïàóêîâ, ïðèçíàþòñÿ óêðóïíåíèå è âûãèáàíèå îñíîâíûõ ÷ëåíèêîâ õåëèöåð è ñìåùåíèå òî÷êè ïðèêðåïëåíèÿ êîãîòêà íà èõ âåíòðàëüíóþ ñòîðîíó. Õåëèöåðû Liphistiidae è Mygalomorphae ïðèíèìàþòñÿ ïëåçèîìîðôíûìè ïî çàíèìàåìîìó ïîëîæåíèþ è îñåâîé îðèåíòàöèè, íî àïîìîðôíûìè ïî ñâîåé êîíôèãóðàöèè; ïîñëåäíÿÿ ïðåäñòàâëÿåòñÿ ðåçóëüòàòîì ýâîëþöèîííîãî êîìïðîìèññà, âûçâàííîãî ïåðåõîäîì ê íîðíîìó îáðàçó æèçíè. Âîçìîæíîñòü ïðîèñõîæäåíèÿ õåëèöåð ëàáèäîãíàòíîãî òèïà èç ñïåöèàëèçèðîâàííûõ îðòîãíàòíûõ õåëèöåð ëèôèñòèèä è ìèãàëîìîð- ôîâ ïðèçíàåòñÿ ìàëîâåðîÿòíîé. KEY WORDS: Araneae, spiders, chelicerae, evolution. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Araneae, ïàóêè, õåëèöåðû, ýâîëþöèÿ. Introduction considered (explicitly or by implication) the orthognathous chelicerae in spiders as precur- Until recently the traditional standpoint treat- sors of the labidognathous type. Starobogatov ing the cheliceral construction in the orthogna- [1985] also used a third category, plagiaxiality, thous spiders as an initial state of the labidogna- a somewhat intermediate cheliceral orientation thous chelicerae was dominant and used in a occurring in the Hypochiloidea. Otherwise, his great number of arachnological publications reconstruction was given in the traditional style: [see Platnick & Gertsch, 1976]. All those works orthognathyplagiaxial statelabidognathy. 350 EUROPEAN ARACHNOLOGY 2003 Kraus & Kraus [1993] first arrived at the (North American mygalomorphs including repre- conclusion that not only labidognathous cheli- sentatives of the Antrodiaetidae, Microhexura, etc.). cerae, but also the orthognathous cheliceral con- Also examined were some taxa of the Uropygi and struction represented the apomorphic modifica- the Araneomorphae. The majority of whip spider tions of a primary type defined as the plagiaxial specimens used, a few females representing two species of Liphistius, and a specimen of Hypochilus variant. In their opinion, this model can be repre- coylei (all from the collection of the American Mu- sented in all main lineages of the order Araneae. seum of Natural History) were kindly lent to me for The cheliceral configurations known for the Li- study through the courtesy of Dr. Norman Platnick. phistiidae, for a number of mygalomorph taxa Most of the figures were prepared specially for possessing shorter chelicerae, and for the repre- this study. Additionally, I had to use some drawings sentatives of the family Hypochilidae, were con- taken from other sources, all these were scanned so sidered as the most similar to the mentioned type. that the output was changed minimally from their Dunlop [1997] supported this hypothesis original form. In a few extraordinary cases some and added a new state, palaeognathy, found in figures were redrawn from the original source. All members of the fossil Paleozoic order Trigono- credits and relevant references are provided. With minor exceptions, the terminology follows tarbida. He suggested that both orthognathy Kaestner [1956], Starobogatov [1985], Shultz and labidognathy can be derived from palaeog- [1990], Kraus & Kraus [1993] and Dunlop [1994, nathy through the plagiaxial type by simple 1996c, 1997]. The subbasal cheliceral segment cor- torsions of basal cheliceral segments, and some responds to the cheliceral coxa of lower arachnids, particular states can be explained by reversals. absent in spiders and their Recent relatives [see However, the palaeognathplagiaxial ver- Dunlop, 1997]. sion appears to be only one of a few possible Some terms as well as the states denoted should ways of explaining the situation observed. A be specified. The term plagiaxial is used only in new attempt to restore the idea to separate the cases when the cheliceral fangs meet each other at approximately 90°, axial orientation of the basal orthognaths and labidognaths from each other cheliceral segments occupies an intermediate posi- was made by Eskov & Zonstein [1990a]. A tion between the forward-directed and downward- little later, I demonstrated another scheme show- directed types; these axes are divergent laterally. ing the independent origin of orthognathous The terms palaeognathy, plesiognathy and neog- and labidognathous chelicerae in spiders at the nathy as well as the derived forms (e.g., plesiogna- Third Eurasian Arachnological Conference. This thous) are used to designate the orthognathous states report was briefly reviewed by Mikhailov found in the Trigonotarbida, in the Pedipalpi (that [1992], but it has never been published. Despite includes the orders Amblypygi, Uropygi and Schi- the general formal resemblance of both hypoth- zomida) and in the Araneae, respectively. The first term (palaeognathy) was first used by Dunlop [1997], eses, they were based on quite different as- whereas the latter two are newly applied to the sumptions and a number of the significant dif- arachnids. The swollen neognathous chelicerae are ferences between them could be listed. The characterized by the claw insertion point displaced expanded and reworked variant of the 1992 ventrad [Dunlop, 1994: fig. 1]. The least modified version is given here. plesiognathous construction retains many ancestral characters including a trapezoidal basal cheliceral Material and methods segment provided with a toothed vestige of the former immovable finger [Dunlop, 1994: figs 2, 3]. Finally, A noticeable part of the material used for this the palaeognathous cheliceral configuration includes study was lent from the spider collection of the both the fangs more or less displaced, and the gener- Zoological Institute of the Russian Academy of al cheliceral orientation changed from forward- to Sciences, St. Petersburg, in the early 1990s. It in- downward-directed [Dunlop, 1994: fig. 4]. cluded numerous mygalomorph taxa belonging to The term phrynid type applies only to the gener- the families Atypidae, Idiopidae, Actinopodidae, alized subchelate construction occurring in the Re- Ctenizidae, Cyrtaucheniidae, Hexathelidae, Diplu- cent taxa represented by the Amblypygi, the Uropy- ridae, Nemesiidae, Barychelidae and Theraphosidae. gi, the Schizomida, and found to be applicable also In addition, material was kindly sent to me by Dr. for some members of the fossil order Trigonotarbida Kirill Eskov (Heptathele) and Dr. Frederick Coyle (for details see the corresponding parts of this study). S.L. Zonstein. Origin and evolution of spider chelicerae 351 This term reflects a certain evolutionary level only; Pedipalpi it does not convey any taxonomic context. Plesiomorphically, ancestors of the Tetra- pulmonata presumably possessed three-seg- Results and discussion mented, chelate chelicerae with the movable finger (the future tetrapulmonate cheliceral fang) Origin located retrolaterally; it is the cheliceral con- struction that is shared by the less advanced arachnids, including representatives of the or- The allied groups der Palpigradi adjacent to the tetrapulmonates Most phylogenetic studies agree in placing [Dunlop, 1997; Selden & Dunlop, 1998]. Araneae into the same group with Amblypygi, The orthognathous chelicerae occurring in Uropygi and Schizomida other arachnid or- the representatives of those orders were consid- ders with members also possessing two pairs of ered to be the most archaic cheliceral construc- book-lungs and the clasp-knife-shaped cheli- tion occurring within this pair of taxa. The main cerae1. Together they constitute the Tetrapul- constructive features of this type are as follows: monata Shultz, 1990. Some disagreement oc- (1) contrary to spiders, the chelicerae here are curs only in drawing spiders closer to one or relatively small (cf. Figs 18); (2) basal cheli- another subtaxon within this group. ceral segments are trapezoidal and flattened; Until recently Amblypygi were found to be (3) the fang is relatively short and joined to the the closest relatives of spiders [see Platnick & basal segment close to its foremost dorso-api- Gertsch, 1976]. Most often Uropygi s.lat. (The- cal vertex; (4) the toothed part of cheliceral lyphonida + Schizomida) has been considered a furrow representing