A Harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie Cherts, Aberdeenshire, Scotland Jason A
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Transactions of the Royal Society of Edinburgh: Earth Sciences, 94, 341–354, 2004 (for 2003) A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland Jason A. Dunlop, Lyall I. Anderson, Hans Kerp and Hagen Hass ABSTRACT: A harvestman (Arachnida: Opiliones) is described from the Early Devonian (Pragian) Rhynie cherts, Aberdeenshire, Scotland. Eophalangium sheari gen. et sp. nov. is the oldest known harvestman. The material includes both males and a female preserving, respectively, a cuticle-lined penis and ovipositor within the opisthosoma. Both these structures are of essentially modern appearance. The Rhynie fossils also show tracheae which are, again, very similar to those of living harvestmen. This is the oldest unequivocal record of arachnid tracheal respiration and indicates that E. sheari was terrestrial. An annulate, setose ovipositor in the female suggests that it can be excluded from the clades Dyspnoi and Laniatores, in which the ovipositor lacks such annulations. However, the penis shows evidence of two muscles, a feature of uncertain polarity seen in modern Troguloidea (Dyspnoi). The presence of median eyes and long legs excludes Cyphoph- thalmi, and thus, E. sheari is tentatively referred to the suborder Eupnoi. Therefore, this remarkable material is implicitly a crown-group harvestman and is one of the oldest known crown-group chelicerates. It also suggests an extraordinary degree of morphological stasis within the eupnoid line, with the Devonian forms differing little in gross morphology – and perhaps in reproductive behaviour – from their modern counterparts. KEY WORDS: Arthropoda, Eupnoi, ovipositor, penis, phylogeny, tracheae, terrestrialisation. Harvestmen (Arachnida: Opiliones) are arachnids in which the small, cryptic, rather mite-like animals with a wide, but prosoma and opisthosoma are characteristically fused together somewhat localised, distribution (for an overview, see Giribet with little visible tagmosis of the body. Other apomorphic 2000). Laniatores is the most diverse group and includes larger, features include prosomal repugnatorial glands, an arrange- often quite bizarre-looking, brightly coloured and spiny ani- ment of the sclerites such that the genital opening is thrust mals with characteristically massive, sub-raptorial pedipalps forward between the leg coxae, a penis (in the male) and (for a recent summary, see Kury 2003). Extant Laniatores ovipositor (in the female), and a single pair of tracheae occur predominantly in tropical zones of the southern hemi- opening on the genital segment (Martens 1978; Shultz 1990, sphere. Palpatores are the most widespread group in northern, 1998). In some ingroup harvestmen, the second pair of legs is temperate regions (e.g. Martens 1978), although recent analy- longest and has a tactile, almost antenna-like function. While ses disagree about whether they are monophyletic (Shultz principally predatory on other arthropods, living harvestmen 1998) or paraphyletic (Giribet et al. 1999, 2002) (see Section 4). are – unusually for arachnids – to some extent omnivorous Nevertheless, all these models recognised two monophyletic (Ru¨hm 1926), and their diet can include dead arthropods, bird palpatore taxa, traditionally assigned the rank of tribe, but droppings, rotting fruit and other types of plant material. raised to suborders by Giribet et al. (2002). Dyspnoi include Some species specialise on snails. Most arachnids digest their the often cryptic troguloid harvestmen, some of which cover prey preorally and need to filter the liquified food through the themselves with soil particles. The remaining suborder, mouthparts. This implicitly terrestrial mode of feeding was Eupnoi, are the most familiar and commonly encountered developed early in arachnid history and was used by, for living harvestmen in temperate regions, easily recognisable by example, the Rhynie chert trigonotarbid arachnids (Dunlop their compact, oval bodies and long, slender legs. The present 1994). By contrast, harvestmen can ingest solid food particles, authors argue that their Rhynie fossils are probably eupnoids. a putative plesiomorphy which they share with xiphosurans This paper describes three exceptionally preserved harvest- (horseshoe crabs) and opilioacariform mites. Other plesiomor- men from the Early Devonian Rhynie cherts of Aberdeenshire, phic characters of harvestmen include chelate chelicerae Scotland. Two of these fossils were briefly described by formed from three articles and, perhaps, the absence of Dunlop et al. (2003) and represent the oldest record of Malpighian tubules, a type of excretory organ adapted to life Opiliones. Like many Rhynie arthropods (Hirst 1923; Hirst & on land. The position of Opiliones within Arachnida remains Maulik 1926; Scourfield 1926; Claridge & Lyon 1961; Ander- contentious. The recent, and comprehensive, study of Giribet son & Trewin 2003), they include remarkable details of the et al. (2002) found most support for placing harvestmen with internal and external anatomy. The tracheae described here are scorpions, pseudoscorpions and solifuges in Shultz’s (1990) the oldest unequivocal evidence for arachnid tracheal respira- clade Dromopoda, but concluded that current datasets are not tion and imply that these harvestmen were fully terrestrial (see decisive in resolving their affinities. Section 5.2). When compared to extant taxa, we can sex the Approximately 5500 living species of harvestmen are fossils based on their preserved genitalia. These structures also known, traditionally divided among three quite distinct sub- allow behavioural inferences about these Early Devonian orders (Shultz 1998; Giribet et al. 1999). Cythophthalmi are arachnids. 342 J. A. DUNLOP ET AL. 1. Other fossil harvestmen elevated levels of dissolved silica. Through interaction with organic detritus, these pools, rich in filamentous algal and In spite of their Recent diversity, fossil harvestmen are rare. cyanobacterial growth, provided ideal sites of preservation for The oldest previous record is an unnamed, long-legged the aquatic and terrestrial floras and faunas in the immediate Scottish specimen from the Lower Carboniferous of East area of the hot springs (see also Anderson & Trewin 2003). Kirkton (Wood et al. 1985). Four Coal Measures species have Further details of the geological setting, dating (Pragian) and been described: two from Commentry in France (Thevenin the preservational environment at Rhynie can be found in Rice 1901) and two from Mazon Creek in the USA (Petrunkevitch et al. (2002) and references therein. 1913). As noted by Beall (1997), at least one Mazon Creek 2.1.1. Palaeoenvironment. All three specimens described in species is a poorly preserved phalangiotarbid (an extinct the present paper were found contained within a 5–6-cm-thick order), and fossils from this locality referred to another extinct bed of massive, white chert within a single block. The bed itself arachnid order, Kustarachnida, are in fact misidentified har- contains a monospecific, in situ flora of mature Aglaophyton vestmen. Redescriptions would be welcome, but most of these plants, bearing numerous terminal sporangia which are either Palaeozoic fossils provisionally resemble eupnoids, while one empty or partially filled with spores. The Aglaophyton axes of the Commentry species is probably a dyspnoid. A putative show excellent preservation. Typically, all cells and tissues are harvestman from the Lower Permian of Germany is a misiden- preserved, indicating massive and rapid silicification. All aerial tified plant fragment (Ro¨ßler et al. 2003). A Jurassic ‘harvest- axes of Aglaophyton are oriented parallel to each other at an man’ described from Solnhofen in Germany is a crinoid angle of about 50( to the vertical plane, suggesting a strong (Petrunkevitch 1955), however unequivocal, but unnamed, and directional (?flooding) event prior to silicification. The Mesozoic palpatore harvestmen have been reported from the white bed overlies a chert-cemented sandstone, rich in com- Lower Cretaceous of Australia (Jell & Duncan 1986) and the pressed and decomposed axes of land plants. Several of the Upper Cretaceous of Lebanon (Roger 1946). Tertiary harvest- Aglaophyton axes of the white chert bed can be traced into the men mostly comprise numerous Baltic and Dominican amber underlying sandstone, suggesting that the sandstone represents inclusions and shale fossils from Florissant in Colorado; for the former soil and the Aglaophyton of the chert bed its surface example, see Cokendolpher (1986 and references therein) for flora. The white chert is directly overlain by a second massive details and Stare˛ga (2002) for a recent revision of the Baltic chert bed, up to 10 cm thick. This second chert also contains amber fauna. Most of these Tertiary fossils are eupnoids and a monospecific, in situ flora of Aglaophyton, with excellent dyspnoids. Fossil Laniatores are somewhat rarer, but have preservation in parts. Here, the aerial axes of Aglaophyton been recorded from both Baltic and Dominican amber (e.g. occur in an irregular orientation in this second chert bed. No Cokendolpher & Poinar 1998). The first fossil cyphophthalmid organic connection was found between the Aglaophyton plants has recently been discovered in the German Bitterfeld amber of both beds, indicating that the lower chert bed was silicified (Dunlop & Giribet 2003) and most of the Tertiary amber prior to being colonised by a new Aglaophyton flora. harvestmen can be referred to living genera. 2.1.2. Organic content.