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1 June 2009 A Review of the Fossil Record of Spiders (Araneae) with Special Reference to Africa, and Description of a New Specimen from the Triassic Molteno Formation of South Africa
Paul A. Selden, Heidi M. Anderson, John M. Anderson
Author Affiliations +
Abstract

The fossil record of spiders as currently known is briefly reviewed, with special reference to Africa. The second specimen of Triassaraneus andersonorum Selden in Selden et al. 1999 is described from a different locality in the Triassic (Carnian, c. 225 Ma) Molteno Formation of South Africa.

INTRODUCTION

Over 90 % of fossil spiders originate from ambers of Cainozoic age (less than 65 Ma old). Cainozoic ambers preserve representatives of the majority of the 109 spider families known today. Of greater interest for phylogeny, therefore, is the fossil record of Araneae from the Mesozoic and Palaeozoic eras, and what it can tell us about the origins and longevity of major clades. Fossil spiders have been known from the late Palaeozoic Coal Measures of Europe and North America for more than a century (although it must be mentioned that some of these specimens are certainly not spiders, Penney & Selden 2006), but the first genuine Mesozoic spider to be formally described was only in 1984 (Eskov 1984). Over the succeeding decades only a few new species have been reported. More recently, much new material is being discovered in, for example, lower Cretaceous deposits of north-east Brazil and China, which is being described at the moment.

It is because of their great rarity that almost every new specimen of spider from the Palaeozoic and Mesozoic eras is worthy of report, and each occurrence can drastically alter our perception of spider phylogeny. For example, the identification of a single record of a family earlier than previously known predicts the occurrence of its sister clade total group to the same date. Thus, Penney's (2002) description of Cretaceous Oonopidae predicted the occurrence of the family's sister group, Orsolobidae. As pointed out by Penney et al. (2003) and others, the true biodiversity at any time in the past is greater than estimated from raw fossil data because range extensions and ghost lineages need to be taken into account. Figure 1 illustrates the present state of knowledge of the fossil record of spiders.

For the last two decades, the oldest known spider in the fossil record was thought to be Attercopus fimbriunguis (Shear, Selden & Rolfe in Shear et al. 1987), from the Devonian of New York, USA. Attercopus was considered sister to all other spiders. Recently, however, Attercopus has been shown to belong to a new order of arachnids: the Uraraneida Selden & Shear in Selden et al. 2008, an order that also includes the Permian Permarachne Eskov & Selden, 2005, which was originally described as an unusual mesothele. Mesotheles are known from upper Carboniferous and Permian strata (Selden 1996a, b, 2000; Eskov & Selden 2005), but no opisthotheles until Triassic time; putative Palaeozoic opisthotheles have been shown to be other arachnids or unrecognizable (Penney & Selden 2006). The oldest opisthothele known is the mygalomorph Rosamygale grauvogeli Selden & Gall, 1992, from the Lower Triassic (Anisian) of France, which was the first spider to be described from rocks of this period. Selden in Selden et al. (1999) described the oldest araneomorphs, Triassaraneus andersonorum (Fig. 3) and Argyrarachne solitus, from the Carnian Molteno Formation of KwaZulu-Natal, South Africa, and the Carnian Cow Branch Formation of Virginia, USA, respectively; their presence in the Triassic Period having been predicted by the occurrence of their sister group in 1992 (Selden & Gall 1992). Single, monotypic specimens were described from each of these localities; additional, but poorly preserved, specimens were noted, but not described. In this paper an additional specimen of Triassaraneus, now confirmed as an araneomorph spider, is described from the Molteno Formation locality of Telemachus Spruit in Eastern Cape, South Africa.

Fig. 1.

Summary phylogenetic tree of Araneae, produced by combining the fossil record with the cladograms of Coddington and Levi (1991), Griswold (1993), Scharff and Coddington (1997), Griswold et al. (1998, 1999), and Ramírez (2000).

f01_105.jpg

Few spiders have been described from the Jurassic Period. The first to be described was Juraraneus Eskov, 1984, an orb-weaver from Transbaikalia, and the second, Jurarchaea, Eskov, 1987, is an archaeid (sensu lato) from Kazakhstan. One, from the Jurassic of Grimmen, Germany, has been figured but not yet formally described (Ansorge 2003; preliminary investigation by the senior author suggests this may be a palpimanoid), and additional specimens from Russian localities remain undescribed. More than 400 specimens recently become available for study from beds of Middle Jurassic age at Daohugou, Nincheng County, Inner Mongolia, China (Huang et al. 2006). Most belong to Uloboridae, and have yet to be studied, but some belonging to the superfamily Palpimanoidea (sensu Forster & Platnick 1984) were described recently (Selden et al. 2008).

Fig. 2.

Location map of the Telemachus Spruit locality (Eastern Cape, South Africa) in relation to the outcrop of the Late Triassic Molteno Formation (shown in black) (after Anderson et al. 1998).

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Few Cretaceous spiders had been formally described and named until recently, but work is continuing apace to rectify this situation. Cretaceous records include: indeterminate Araneae from Koonwarra, South Australia (Jell & Duncan 1986); mygalomorphs from Transbaikalia and Mongolia (Eskov & Zonshtein 1990); orbicularian araneomorphs from the Sierra de Montsech and Las Hoyas localities, Spain (Selden 1989, 1990, 1991; Selden & Penney 2003); an unnamed lycosoid from Orapa, Botswana (Rayner & Dippenaar-Schoeman 1995); a poorly preserved specimen from Mexico (Feldmann et al. 1998); many new spider specimens reported from the Crato Formation of Brazil, of which just a few have been described so far (Mesquita 1996; Selden et al. 2002, 2006); and many also are now being collected from the Lower Cretaceous of China (Chang 2004; Cheng et al. 2008). Spiders are known from Cretaceous amber from Canada (McAlpine & Martin 1969; Penney & Selden 2006), the Caucasus and Siberia (Eskov & Wunderlich 1994; Zherikhin & Eskov 1999), France (Schlüter 1978; Saupe & Selden 2009), Lebanon (Penney 2003a; Penney & Selden 2002), Myanmar (Cockerell 1920; Rasnitsyn & Ross 2000; Zherikhin & Ross 2000; Penney 2003b; Wunderlich 2008), New Jersey (Grimaldi et al. 2000; Penney 2002), Álava, Spain (Alonso et al. 2000; Penney 2006), Asturias, Spain (Arbizu et al. 1999), Jordan (Kaddumi 2007; Wunderlich 2008) and the Isle of Wight, England (Selden 2002).

At the time of publication of Eskov and Zonshtein's (1990) paper on Cretaceous mygalomorphs, there were hardly any spiders known at all from the Mesozoic era. It appeared that mygalomorphs, perhaps already an archaic group, may have flourished at a time of biotic crisis, when araneomorphs became reduced in number (Eskov & Zonshtein 1990). Since then, however, many more Mesozoic spiders, and particularly araneomorphs, have been described. Figure 1 shows that by Cretaceous time, many of the modern families were in existence or predicted by sister-group relationships. By the time of formation of the mid-Cainozoic ambers, nearly all extant families have fossil representatives or are predicted to occur. Interestingly, there are very few families known only from fossils; some new families from Cretaceous ambers recently described by Wunderlich (2008) have yet to be critically assessed. Some families described initially from fossils were later found to be extant, e.g. Archaeidae were first described from Baltic amber and later found alive in Africa and Madagascar. The Cretaceous extinct family, Lagonomegopidae Eskov & Wunderlich, 1994, is interesting on account of (a) its enormous posterior median eyes, which suggest a possible relationship (homology or analogy) with Salticidae (the most speciose modern family, and unknown in preCainozoic strata), and (b) the characteristic peg-teeth, which place it within the Palpimanoidea sensu Forster and Platnick (1984). Few arachnologists concur with the composition of this superfamily; in particular Schütt (2002) has provided evidence, mainly from spinneret morphology, that the palpimanoid families formerly allied with Araneoidea should be returned to that superfamily. These include Mimetidae and the archaeid families, which are araneophages characterised by elongated chelicerae bearing long peg-teeth. Of the described Jurassic spiders, several (Jurarchaea, Patarchaea Selden, Huang & Ren, 2008, Metaxyostraca Selden, Huang & Ren, 2008, and an undescribed specimen from Germany) belong in the Palpimanoidea, and the other (Juraraneus) is an early orb-weaver. The earliest known araneomorphs, Triassaraneus and Argyrarachne, resemble araneoids more than any other group. Conversely, cursorial spiders such as the Dionycha, including such speciose families as Salticidae, Thomisidae, Gnaphosidae and the clubionoids, which represent some 35 % of genera and 37 % of species at the present day (data from Platnick 2009), are as yet unknown in pre-Cainozoic strata. The fossil record thus appears to support the hypothesis, well argued by Dippenaar-Schoeman and Jocqué (1997), that the more derived hunting spiders arose from ancestors which used webs for prey capture (Vollrath & Selden 2007).

A study by Penney et al. (2003) examined how spiders fared across well-known mass extinctions, specifically the mid-Cretaceous Cenomanian—Turonian (C/T, 93.5 Ma), and the better-known Cretaceous—Tertiary (K/T, 65 Ma) events. Using quantitative techniques combining taxonomic and numerical data on amber fossils only, these authors showed that spiders suffered no decline at the family level during these mass extinctions. On the contrary, they increased in relative numbers through the Cretaceous and beyond the K/T boundary. They suggested that this extinction resistance of spiders is due to the majority of them being generalist predators. Those feeding predominantly on plantspecific herbivores that became extinct could easily have switched to a new primary food source, such as many of the non-herbivorous insects or polyphagous herbivores, which appear to have been little affected by the K/T extinction (Labandeira & Sepkoski 1993; Labandeira et al. 2002). Thus, the mass extinction that killed off the dinosaurs seems to have had little effect on the Araneae, at least at family level.

African fossil spiders

Spiders are common in African copal, which is a kind of sub-fossil amber; it chiefly occurs on the coasts of Kenya, Tanzania and Mozambique, and on the north-west coast of Madagascar. Softer and paler yellow than true amber, it dates from about Miocene and younger in age, and so its inclusions are generally of less interest for phylogenetic studies. It can be of great interest, however, for research on palaeogeography and palaeoclimatology of more recent geological times. For example, interesting work has been done comparing the fossil and extant spider species in Dominican amber and alive on the island of Hispaniola respectively (Penney 2008). In spite of specimens having been available in gem shops relatively cheaply throughout the world, few descriptions of African copal spiders appeared until recently. There can be a problem, however, in that some amber dealers will heat copal to make it appear to be true amber and thus gain higher prices. For example, Wunderlich (2004) suggested that the specimen of Entomocephalus formicoides Holl, 1829 from Baltic amber was really a fake in Madagascan copal. Holl's one-sentence description and figure (pl. 8, fig. 68a) suggests it almost certainly belongs to an ant-mimicing salticid genus such as Myrmarachne (which occurs in Madagascan copal), although the figure and description mentioned only six eyes (Penney 2003b). It was placed (erroneously) in Archaeidae by Petrunkevitch (1958); the location of the holotype is unknown.

Lourenço (2000) described Archaea copalensis (Archaeidae), which was thought to be the first record of this genus from Madagascan copal; the species was later synonymized with the extant Archaea gracilicollis Millot, 1948 by Wunderlich (2004). Previously, Wunderlich (1998) had described Mysmena dominicana (Mysmenidae) and Grammonota deformans (Linyphiidae, later placed in Ceratinopsis by Wunderlich (2004)) from supposed Dominican amber. He later realized that the amber was actually Madagascan copal treated to make it appear to be older and from a different provenance (Wunderlich 2004). Similarly, Wunderlich's (1999) report of the family Archaeidae from Dominican Republic amber also proved to be a mistake for Madagascan copal. Species from the following spider families are currently known as sub-fossils in Madagascan copal: Araneidae, Archaeidae, Clubionidae, Corinnidae, Deinopidae, Dictynidae, Hahniidae, Hersiliidae, Linyphiidae, ?Migidae, ?Miturgidae, Mysmenidae, Oonopidae, Philodromidae, Pholcidae, Salticidae, Scytodidae, Segestriidae, Selenopidae, Tetragnathidae, Theridiidae, Thomisidae and Uloboridae (Wunderlich 2004, 2008).

Only two specimens of fossil spider have been described from the continent of Africa: Triassaraneus, as mentioned above, and a supposed lycosoid spider from the Orapa diamond mine in Botswana (Rayner & Dippenaar-Schoeman 1995). The Orapa mine exploits a kimberlite pipe in which a crater lake developed during the Cretaceous period; the matrix bearing the spider and abundant insect remains has been dated to 93 Ma ago (Late Cretaceous: Turonian) by radiometry (Rayner et al. 1997). The relatively soft volcanic breccias of kimberlite pipes commonly accommodate circular crater lakes (see Gernon et al. 2009 for a review of the sedimentology of the Orapa kimberlite). The Orapa lake was shallow and the fine lamination of the sediment suggests little disturbance, so it was probably anoxic bottom waters which prevented decay of the biota. Because there has been little appreciable erosion since Cretaceous times, the lake sediments are still preserved. More than half of the insects at Orapa are Coleoptera, followed by Blattodea (about 20 %), and less than 10 % each of Diptera, Hymenoptera, Hemiptera, Orthoptera, Dermaptera and the rest (Rayner et al. 1997). A diverse flora of mainly angiosperms (about 90 %) that was washed into the lake has been described by Bamford (1990); ferns account for the remainder of the flora. The only spider to have been described from Orapa is an unnamed specimen with a body length of approximately 25 mm. It was referred to superfamily Lycosoidea on the basis of its habitus being that of a hunting spider. This was the first fossil spider to be described from Africa.

The second, and most important, fossil spider to be described from Africa comes from the Molteno Formation of the Karoo (Fig. 2), which was deposited in an extensive, intracontinental foreland basin bounded by rising fold mountains to the south and traversed by a system of braided rivers (Cairncross et al. 1995). The Molteno Formation reaches a maximum thickness of about 600 m and the erosional remnant extends over an area roughly 400 km north to south and 200 km west to east. The age of the formation is not tightly established, but on the basis of global biostratigraphic correlations (Anderson & Anderson 1983, 1989) is considered to be Carnian (Late Triassic), 222– 229 Ma. No absolute radiometric ages are available. A 30-year collecting programme (Anderson 2001; Anderson & Anderson 1983, 1984, 1989; Anderson et al. 1998) has yielded 100 fossil plant assemblages from the Molteno Formation. The flora, the richest known globally from the Triassic, includes 56 genera with 204 vegetative species. It is particularly characterised by some 20 species of Dicroidium (Ginkgoopsida). There is a roughly equal diversity of gymnosperms, including conifers, cycads and ginkgos, along with several new orders, and ‘pteridophytes,’ primarily horsetails and ferns. Though rare, insects comprise by far the most frequently encountered element of the fauna. A remarkable diversity of 117 genera and 333 species in 18 orders is provisionally recognised in the over 2000 specimens from 43 of the 100 plant assemblages. Coleoptera, Blattodea and Hemiptera dominate. Conchostracan crustaceans, from 20 of the plant assemblages, are represented by some three genera and eight species. The remaining fauna is sparse: three species of fishes (impressions only) from three assemblages, two species of bivalve from one assemblage, and two spider specimens. Dinosaur trackways, but no skeletal remains, have been identified at a few (non-plant) sites. Anderson (2001) provided a review of the Molteno Formation flora and fauna.

Fig. 3.

Holotype of Triassaraneus andersonorum from the Triassic (Carnian) Molteno Formation, Upper Umkomaas ‘Waterfall Locality’ (UMK III), KwaZulu-Natal, South Africa: (A) photograph in incident light under ethanol, (B) camera lucida drawing. Scale bar = 1 mm.

f03_105.jpg

At the Telemachus Spruit locality there is a 10 cm thick, buff mudstone that is interpreted as an abandoned channel-fill (Cairncross et al. 1995). The flora of 12 genera and 19 species is strongly dominated by the single coniferous species Heidiphyllum elongatum (Morris, 1845) Retallack, 1981. The assemblage most likely represents two distinct plant communities: a mono-dominant stand of reed-like conifers colonizing sand bars in the braided river and, from farther afield, a Dicroidium-dominated riparian forest occupying the river bank. The insect fauna at this site, is represented by only 17 fragmentary specimens, and is dominated, as at other Molteno localities, by Coleoptera, Blattodea and Hemiptera. Conchostracans have not been found. The rarity of the single spider specimen is emphasized by the fact that John and Heidi Anderson have spent 90 man-hours cleaving slabs at this site and that all 900 curated and catalogued slabs have been carefully scanned for insects or other faunal elements under the binocular microscope.

TAXONOMY

Infraorder Araneomorphae Smith, 1902
Superfamily ?Araneoidea Latreille, 1806
Triassaraneus Selden in Selden et al., 1999
Type and only species: Triassaraneus andersonorum Selden in Selden et al., 1999.
Triassaraneus andersonorum Selden in Selden et al., 1999
Figs 35

  • Material examined: SOUTH AFRICA: KwaZulu-Natal: Holotype PRE/F 18560a (part) and 18560b (counterpart), immature or mature female from Upper Umkomaas ‘Waterfall Locality’ (UMK III). Eastern Cape: PRE/F 17234 Tel 111, immature from Telemachus Spruit locality (Anderson & Anderson 1983, 1984). All material from from Member Z of the Late Triassic (Carnian) Molteno Formation and deposited in the palaeobotanical collection of the South African National Biodiversity Institute, Pretoria.

  • Description of new specimen: The specimen is preserved as brown, organic cuticle on a pale, grey-brown shale. The shale is splintery and pieces readily fall away. Scattered throughout the shale are abundant plant remains, including on the fossil slab a male cone of Ginkgoopsida. Parts of at least three legs are preserved, though it is not possible to determine which legs these are. No body parts are preserved. On the right in Figs 4, 5 is a nearly complete walking leg, showing most of the femur to the tarsus. On the left are some of the opposing walking legs, but less complete. Note that in Fig. 5A, there is a loose piece of matrix bearing parts of a leg; this became detached and is not present in Fig. 5B (under ethanol), which therefore reveals more of the tibia of on the left-hand walking legs. The walking legs bear setae on all podomeres, and larger, thin spines on all podomeres except the tarsus. These spines, for example on the right leg patella and tibia, are long, thin and erect. Although tarsal claws cannot be seen, it is certain from the shape of this podomere (only preserved on the right side) that its distal end is present, so the claws must be small. There is no evidence of tarsal scopulae or claw tufts. The legs appear to be rather short, although the only one which is nearly completely preserved, on the right, may be a third leg, which is commonly short in orbicularians (orb-web weavers).

  • Figs 4, 5.

    T. andersonorum specimen PRE-F 17234 Tel 111 from Telemachus Spruit: (4) camera lucida illustration, (5) actual fossil dry (A) and under ethanol (B). Abbreviations: fe - femur, mt - metatarsus, pa - patella, ta - tarsus, ti - tibia. Scale bar = 0.5 mm.

    f04_105.jpg

    DISCUSSION

    The small claws and lack of tarsal scopulae and claw tufts indicate that this was a web-living rather than a ground spider. Short legs suggest that this is a juvenile spider; however, if the right leg is indeed a third leg of an orbicularian, then this may not be the case. If a third leg, then it is not a great deal smaller than that of T. andersonorum, so it is unlikely to be a juvenile of that species. The leg spination of the right leg matches that of leg 3 in T. andersonorum, with one distally on the femur, one on the patella, two in the proximal half and one in the distal half of the tibia (compare Fig. 4 with fig. 2 in Selden et al. 1999). Therefore, this fossil is considered to be a second specimen of T. andersonorum. Triassic spiders are extremely rare, and this is only the second spider to come out of the Molteno Formation, after many years of painstaking search. It comes from a different locality from T. andersonorum, but is of the same age and possibly occupied the same habitat (Dicroidium-dominated riparian forest).

    A general impression gained from studying fossil arachnids is that one spider fossil occurs for approximately every 1000 insects in the Mesozoic lacustrine or lagoonal fossil insect localities, e.g. the Cretaceous Crato Formation of Brazil (Mesquita 1996; Selden et al. 2002), the Cretaceous of Spain (Selden 1989, 1990; Selden & Penney 2003) and the Jurassic and Cretaceous of China (e.g. Selden et al. 2008). The ratio in the Lower Cretaceous locality of Baissa, Transbaikalia is 1:1623 (24,366 arthropod remains) and that in the Middle-Upper Jurassic locality of Karatau, Kazakhstan it is 1:904 (19,909 arthropod remains) (M. Mostovski pers. comm., 2009). With some 2000 insects collected from the Molteno Formation, and a new specimen described herein, this ratio of spider to insect fossils is in agreement, and a similar ratio seems to occur in the Cow Branch Formation of Virginia. However, when comparing other Triassic and Permian localities the ratios are rather different. For example, there are no spiders among over 22,000 insect remains in the Ladinian/Carnian locality of Madygen, Kyrgyzstan (Shcherbakov 2008), and, similarly, no arachnids have ever been found among the tens of thousands of insect specimens from the Artinskian Wellington Formation of Kansas and Oklahoma (Beckemeyer & Hall 2007). The ratio is 1:2129 in the Kazanian locality of Verkhnii Kaltan, Siberia (M. Mostovski pers. comm., 2009), and only one spider specimen, one trigonotarbid and one uraraneid are known out of more than 8000 insects in the Kungurian Koshelevka Formation (including the Chekarda locality) of the Urals (Eskov & Selden 2005). A quantitative study of the ratios of insect remains to those of spiders and other arachnids in localities with abundant insect fossils (Rasnitsyn & Zherikhin 2002) would be extremely interesting.

    ACKNOWLEDGEMENTS

    The senior author thanks David Penney (Manchester) for discussion on Mesozoic spiders in amber, Roger Smith (Iziko South African Museum, Cape Town) for hospitality and a field visit to the Molteno Formation, Mike Mostovski (Natal Museum, Pietermaritzburg) for help with insect and spider abundances, and Jason Dunlop and Kirill Eskov for constructive reviews.

    REFERENCES

    1.

    J. Alonso , A. Arillo , E. Barrón , J.C. Corral , J. Grimalt , J.F. López , R. López , X. Martínez-Delclòs , V. Ortuño , E. Peñalver & P.R. Trincão 2000. A new fossil resin with biological inclusions in lower Cretaceous deposits from Álava (northern Spain, Basque-Cantabrian Basin). Journal of Paleontology 74: 158–178. Google Scholar

    2.

    J.M. Anderson , ed . 2001. Towards Gondwana alive. Vol. 1. Pretoria: Gondwana Alive Society. Google Scholar

    3.

    J.M. Anderson & H.M. Anderson 1983. Palaeoflora of southern Africa. Molteno Formation (Triassic) Volume 1. Rotterdam: A .A. Balkema. Google Scholar

    4.

    J.M. Anderson & H.M. Anderson 1984. The fossil content of the Upper Triassic Molteno Formation, South Africa. Palaeontologia Africana 25: 39–59. Google Scholar

    5.

    J.M. Anderson & H.M. Anderson 1989. Palaeoflora of southern Africa. Molteno Formation (Triassic) Volume 2. Rotterdam: A .A. Balkema. Google Scholar

    6.

    J.M. Anderson , H.M. Anderson & A.R.I. Cruickshank 1998. Late Triassic ecosystems of the Molteno/Lower Elliot biome of southern Africa. Palaeontology 41: 387–421. Google Scholar

    7.

    M. Arbizu , E. Bernárdez , E. Peñalver & M.A. Prieto 1999. El ámbar de Asturias. In : J. Alonso , J.C. Corral & R. LÓpez , eds, Proceedings of the world congress on amber inclusions, Álava, Spain. Estudios de Museo de Ciencias Naturales de Álava 14 (Núm. Espec. 2 ): 245–254. Google Scholar

    8.

    J. Ansorge 2003. Insects from the Lower Toarcian of Middle Europe and England. Acta Zoologica Cracoviensia 46 (Suppl. - Fossil Insects): 291–310. Google Scholar

    9.

    M.K. Bamford 1990. The angiosperm palaeoflora from the Orapa Pipe, Botswana. Ph.D. dissertation. Johannesburg, South Africa: University of the Witwatersrand. (Unpubl.) Google Scholar

    10.

    R.J. Beckemeyer & J.D. Hall 2007. The entomofauna of the Lower Permian fossil insect beds of Kansas and Oklahoma, USA. African Invertebrates 48 (1): 23–39. Google Scholar

    11.

    B. Cairncross , J.M. Anderson & H.M. Anderson 1995. Palaeoecology of the Triassic Molteno Formation, Karoo Basin, South Africa sedimentological and palaeontological evidence. South African Journal of Geology 98: 452–478. Google Scholar

    12.

    Chang J.-P . 2004. Some new species of spider and Sacculinidae fossils in Jehol biota. Global Geology 23: 313–320. (in Chinese with English summary) Google Scholar

    13.

    X.-D. Cheng , Q.-J. Meng , X.-R. Wang & C.-L. Gao 2008. New discovery of Nephilidae in Jehol Biota(Araneae, Nephilidae). Acta zootaxonomica sinica 33: 330–334. (in Chinese, with English summary) Google Scholar

    14.

    T.D.A. Cockerell 1920. Fossil arthropods in the British Museum: IV. Annals & Magazine of Natural History, Series 9 6: 211–214. Google Scholar

    15.

    J.A. Coddington & H.W. Levi 1991. Systematics and the evolution of spiders (Araneae). Annual Reviews of Ecology and Systematics 22: 265–292. Google Scholar

    16.

    A.S. Dippenaar-Schoeman & R. Jocqué 1997. African spiders. An identification manual. Plant Protection Research Institute Handbook No. 9. Pretoria: Agricultural Research Council of South Africa. Google Scholar

    17.

    K.Y. Eskov 1984. A new fossil spider family from the Jurassic of Transbaikalia (Araneae: Chelicerata). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1984: 645–653. Google Scholar

    18.

    K.Y. Eskov 1987. A new archaeid spider (Chelicerata: Araneae) from the Jurassic of Kazakhstan, with notes on the so-called “Gondwanan” ranges of recent taxa. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 175: 81–106. Google Scholar

    19.

    K.Y. Eskov & P.A. Selden 2005. First record of spiders from the Permian period (Araneae: Mesothelae). Bulletin of the British Arachnological Society 13: 111–116. Google Scholar

    20.

    K.Y. Eskov & J. Wunderlich 1994. On the spiders from Taimyr ambers, Siberia, with the description of a new family and with general notes on the spiders from the Cretaceous resins. Beiträge zur Araneologie 4: 95–107. Google Scholar

    21.

    K.Y. Eskov & S.L. Zonshtein 1990. First Mesozoic mygalomorph spiders from the Lower Cretaceous of Siberia and Mongolia, with notes on the system and evolution of the order Mygalomorphae (Chelicerata: Araneae). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 178: 325–368. Google Scholar

    22.

    R.M. Feldmann , F.J. Vega , S.P. Applegate & G.A. Bishop 1998. Early Cretaceous arthropods from the Tlayúa Formation at Tepexi de Rodríguez, Puebla, México. Journal of Paleontology 72: 79–90. Google Scholar

    23.

    R.R. Forster & N.I. Platnick 1984. A review of the archaeid spiders and their relatives, with notes on the superfamily Palpimanoidea (Arachnida, Araneae). Bulletin of the American Museum of Natural History 178: 1–106. Google Scholar

    24.

    T.M. Gernon , M. Field & R.S.J. Sparks 2009. Depositional processes in a kimberlite crater: the Upper Cretaceous Orapa South Pipe (Botswana). Sedimentology 56: 623–643. Google Scholar

    25.

    D. Grimaldi , A. Shedrinsky & T. Wampler 2000. A remarkable deposit of fossiliferous amber from the Upper Cretaceous (Turonian) of New Jersey, U.S.A. In : D. Grimaldi , ed., Studies in fossil amber, with particular reference to the Cretaceous of New Jersey. Leiden: Backhuys, pp. 1–76. Google Scholar

    26.

    C.E. Griswold 1993. Investigations into the phylogeny of the lycosoid spiders and their kin (Arachnida: Araneae: Lycosoidea). Smithsonian Contributions to Zoology 539: 1–39. Google Scholar

    27.

    C.E. Griswold , J.A. Coddington , G. Hormiga & N. Scharff 1998. Phylogeny of the orb-web building spiders (Araneae, Orbiculariae: Deinopoidea, Araneoidea). Zoological Journal of the Linnean Society 123: 1–99. Google Scholar

    28.

    C.E. Griswold , J.A. Coddington , N.I. Platnick & R.R. Forster 1999. Towards a phylogeny of entelegyne spiders (Araneae, Araneomorphae, Entelegynae). Journal of Arachnology 27: 53–63. Google Scholar

    29.

    Huang D.-Y ., A. Nel , Y.-B. Shen , P.A. Selden & Q.-B. Lin 2006. Discussions on the age of Daohugou fauna—evidence from invertebrates. Progress in Natural Science 16 (Special issue): 308–312. Google Scholar

    30.

    P.A. Jell & P.M. Duncan 1986. Invertebrates, mainly insects, from the freshwater, Lower Cretaceous, Koonwarra Fossil Bed (Korumburra Group), South Gippsland, Victoria. Memoirs of the Association of Australasian Palaeontologists 3: 311–205. Google Scholar

    31.

    H.F. Kaddumi 2007. Amber of Jordan. The oldest prehistoric insects in fossilized resin. 2nd edition. Amman: Eternal River Museum of Natural History. Google Scholar

    32.

    C.C. Labandeira & J.J. Sepkoski Jr. 1993. Insect diversity in the fossil record. Science 261: 310–315. Google Scholar

    33.

    C.C. Labandeira , K.C. Johnson & P. Wilf 2002. Impact of the terminal Cretaceous event on plant insect associations. Proceedings of the National Academy of Sciences of the USA 99: 2061–2066. Google Scholar

    34.

    P.-A. Latreille 1806. Genera crustaceorum et insectorum. Vol. 1, Aranéides. Paris: A . Koenig, pp. 82–127. Google Scholar

    35.

    W.R. Lourenço 2000. Premier cas d'un sub-fossile d'araignée appartenant au genre Archaea Koch & Berendt (Archaeidae) dans le copal de Madagascar. Comptes Rendus de l'Académie des Sciences, Series IIA - Earth and Planetary Sciences 330: 509–512. Google Scholar

    36.

    J.F. Mcalpine & J.E. Martin 1969. Canadian amber - a paleontological treasure-chest. Canadian Entomologist 101: 819–838. Google Scholar

    37.

    M.V. Mesquita 1996. Cretaraneus martinsnetoi n. sp. (Araneoidea) da Formação Santana, Cretáceo Inferior da Bacia do Araripe. Revista Universidade Guarulhos, Série Geociências 1: 24–31. Google Scholar

    38.

    J. Morris 1845. Fossil flora. In : P.E. De Strzelecki , ed., Physical description of New South Wales and Van Diemans Land. London: Longman, Brown & Green, pp. 245–254. Google Scholar

    39.

    D. Penney 2002. Spiders in Upper Cretaceous amber from New Jersey (Arthropoda, Araneae). Palaeontology 45: 709–724. Google Scholar

    40.

    J. Morris 2003a. A new deinopoid spider from Cretaceous Lebanese amber. Acta Palaeontologica Polonica 48: 569–574. Google Scholar

    41.

    J. Morris 2003b. Afrarchaea grimaldii, a new species of Archaeidae (Araneae) in Cretaceous Burmese amber. Journal of Arachnology 31: 122–130. Google Scholar

    42.

    J. Morris 2006. The oldest lagonomegopid spider, a new species in Lower Cretaceous amber from Álava, Spain. Geologica Acta 4: 377–382. Google Scholar

    43.

    J. Morris 2008. Dominican Amber Spiders: a comparative palaeontological-neontological approach to identification, faunistics, ecology and biogeography. Manchester: Siri Scientific Press. Google Scholar

    44.

    D. Penney & P.A. Selden 2002. The oldest linyphiid spider, in Lower Cretaceous Lebanese amber (Araneae, Linyphiidae, Linyphiinae). Journal of Arachnology 30: 487–493. Google Scholar

    45.

    D. Penney & P.A. Selden 2006. Assembling the Tree of Life — Phylogeny of Spiders: a review of the strictly fossil spider families. In : C. Deltshev & P. Stoev , eds, European Arachnology 2005. Acta Zoologica Bulgarica Supplement 1: 25–39. Google Scholar

    46.

    D. Penney , C.P. Wheater & P.A. Selden 2003. Resistance of spiders to Cretaceous-Tertiary extinction events. Evolution 57: 2599–2607. Google Scholar

    47.

    A. Petrunkevitch 1958. Amber spiders in European collections. Transactions of the Connecticut Academy of Arts and Sciences 41: 97–400. Google Scholar

    48.

    N.I. Platnick 2009. The world spider catalog, version 9.5. New York: American Museum of Natural History. ( http://research.amnh.org/entomology/spiders/catalog/index.html; accessed April 3 , 2009) Google Scholar

    49.

    M.J. Ramírez 2000. Respiratory system morphology and the phylogeny of haplogyne spiders (Araneae, Araneomorphae). Journal of Arachnology 28: 149–157. Google Scholar

    50.

    A.P. Rasnitsyn & A.J. Ross 2000. A preliminary list of arthropod families present in the Burmese amber collection at The Natural History Museum, London. Bulletin of the Natural History Museum, London (Geology) 56: 21–24. Google Scholar

    51.

    A.P. Rasnitsyn & V.V. Zherikhin 2002. Appendix: Alphabetic list of selected insect fossil sites. In : A.P. Rasnitsyn & D.L.J. Quicke , eds, History of Insects. Dordrecht: Kluwer, pp. 437–444. Google Scholar

    52.

    R.J. Rayner , M.K. Bamford , D.J. Brothers , A.S. Dippenaar-Schoeman , I.J. Mckay , R.G. Oberprieler & S.B. Waters 1997. Cretaceous fossils from the Orapa diamond mine. Palaeontologia Africana 33: 55–65. Google Scholar

    53.

    R.J. Rayner & A.S. Dippenaar-Schoeman 1995. A fossil spider (superfamily Lycosoidea) from the Cretaceous of Botswana. South African Journal of Science 91: 98–100. Google Scholar

    54.

    G.J. Retallack 1981. Middle Triassic megafossil plants from Long Gully, near Otematata, North Otago, New Zealand. Journal of the Royal Society of New Zealand 11: 167–200. Google Scholar

    55.

    E.E. Saupe & P.A. Selden 2009. First fossil Mecysmaucheniidae (Arthropoda: Chelicerata: Araneae), from Lower Cretaceous (Upper Albian) amber of Charente-Maritime, France. In : V. Perrichot & D. Néraudeau , eds, Cretaceous ambers from southwestern France: geology, taphonomy, and paleontology. Geodiversitas 31: 49–60. Google Scholar

    56.

    N Scharff , & J.A. Coddinoton 1997. A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae), Zoological Journal of the Linnean Society 120: 355–434. Google Scholar

    57.

    D.E. Shcherbakov 2008. Madygen, Triassic Lagerstätte number one, before and after Sharov. Alavesia 2: 113–124. Google Scholar

    58.

    T. Schlüter 1978. Zur Systematik und Palökologie harzkonservierter Arthropoda einer Taphozönose aus dem Cenomanium von NW-Frankreich. Berliner Geowissenschaftliche Abhandlungen (Serie A) 9:1–150. Google Scholar

    59.

    K. Schütt 2002. The limits of the Araneoidea (Arachnida: Araneae), Australian Journal of Zoology 48: 135–153. Google Scholar

    60.

    P.A. Selden 1989. Orb-web weaving spiders in the early Cretaceous, Nature 340: 711–713. Google Scholar

    61.

    P.A. Selden 1990. Lower Cretaceous spiders from the Sierra de Montsech, northeast Spain. Palaeontology 33: 257–285. Google Scholar

    62.

    P.A. Selden 1991. Aranyes del Cretaci inferior de la Serra del Montsec (Espanya). In : X. Martínez-Delclòs , ed., Les calcàries litogràfiques del Cretaci Inferior del Montsec. Deu anys de campanyes paleontològiques. Lleida: Institut d'Estudis Ilerdencs. pp. 77–85. Google Scholar

    63.

    P.A. Selden 1996a. Fossil mesothele spiders, Nature 379: 498–499. Google Scholar

    64.

    P.A. Selden 1996b. First fossil mesothele spider, from the Carboniferous of France. Revue Suisse de Zoologie Volume hors série 2: 585–596. Google Scholar

    65.

    P.A. Selden 2000. Palaeothele, a replacement name for the fossil mesothele spider Eothele Selden non Rowell. Bulletin of the British Arachnological Society 11: 292. Google Scholar

    66.

    P.A. Selden 2002. First British Mesozoic spider, from Cretaceous amber of the Isle of Wight, southern England. Palaeontology 45: 973–983. Google Scholar

    67.

    Selden. P.A ., J.M. Anderson , H.M Anderson , & N.C. Fraser 1999. Fossil araneomorph spiders from the Triassic of South Africa and Virginia, Journal of Arachnology 27: 401–414. Google Scholar

    68.

    P.A. Selden , F. Da C. Casado & M.V. Mesquita 2002. Funnel-web spiders (Araneae: Dipluridae) from the Lower Cretaceous of Brazil. Boletim do 6° Simpósio sobre o Cretácio do Brasil/2 do Simposio sobre el Cretácico de América del Sur (2002), pp. 89–91. Google Scholar

    69.

    P.A. Selden , F. Da C. Casado & M.V. Mesquita 2006. Mygalomorph spiders (Araneae: Dipluridae) from the Lower Cretaceous Crato Lagerstätte, Araripe Basin, north-east Brazil. Palaeontology 49: 817–826. Google Scholar

    70.

    Selden. P.A ., & J.-C. Gall 1992. A Triassic mygalomorph spider from the northern Vosges, France. Palaeontology 35: 211–235. Google Scholar

    71.

    P.A. Selden , D.-Y. Huang & D. Ren 2008. Palpimanoid spiders from the Jurassic of China. Journal of Arachnology 36: 306–321. Google Scholar

    72.

    P.A. Selden , & D. Penney 2003. Lower Cretaceous spiders (Arthropoda: Arachnida: Araneae) from Spain. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2003: 175–192. Google Scholar

    73.

    P.A. Selden , W.A. Shear & M.A. Sutton 2008. Fossil evidence for the origin of spider spinnerets, and a proposed arachnid order, Proceedings of the National Academy of Sciences of the USA 105: 20781–20785. Google Scholar

    74.

    W.A. Shear , P.A. Selden , W.D.I. Rolfe , P.M. Bonamo & J.D. Grierson 1987. New terrestrial arachnids from the Devonian of Gilboa, New York (Arachnida: Trigonotarbida), American Museum Novitates 2901: 1–74. Google Scholar

    75.

    F.P. Smith 1902. The spiders of Epping Forest. Essex Naturalist 12: 181–201. Google Scholar

    76.

    F. Vollrath & P.A. Selden 2007. The role of behavior in the evolution of spiders, silks, and webs. Annual Review of Ecology, Evolution, and Systematics 38: 819–846. Google Scholar

    77.

    J. Wunderlich 1998. Beschreibung der ersten fossilen Spinnen der Unterfamilien Mysmeninae (Anapidae) und Erigoninae (Linyphiidae) im Dominikanischen Bernstein (Arachnida, Araneae). Entomologische Zeitschrift 108: 363–367. Google Scholar

    78.

    J. Wunderlich 1999. Two subfamilies of spiders (Araneae, Linyphiidae: Erigoninae and Anapidae: Mysmeninae) new to Dominican amber—or falsicated amber? In : J. Alonso , J. C. Corral & R. López , eds. Proceedings of the World congress on amber inclusions, Álava, Spain. Estudios de Museo de Ciencias Naturales de Álava 14 (Núm, Espec. 2 ): 167–172. Google Scholar

    79.

    J. Wunderlich , ed . 2004. Fossil spiders in amber and copal, Beiträge zur Araneologie 3: 1–1908. Google Scholar

    80.

    J. Wunderlich , ed . 2008. Fossil and extant spiders, Beiträge zur Araneologie 5: 1–870. Google Scholar

    81.

    V.V. Zerikhin & K.Y. Eskov 1999. Mesozoic and Lower Tertiary resins in former USSR. In : J. Alonso , J.C. Corral & R. López , eds, Proceedings of the World congress on amber inclusions, Álava, Spain. Estudios de Museo de Ciencias Naturales de Álava 14 (Núm, Espec. 2 ): 119–131. Google Scholar

    82.

    V.V. Zherikhin & A.J. Ross 2000. A review of the history, geology and age of Burmese amber (Burmite). Bulletin of the Natural History Museum, London (Geology) 56: 3–10. Google Scholar
    Paul A. Selden, Heidi M. Anderson, and John M. Anderson "A Review of the Fossil Record of Spiders (Araneae) with Special Reference to Africa, and Description of a New Specimen from the Triassic Molteno Formation of South Africa," African Invertebrates 50(1), 105-116, (1 June 2009). https://doi.org/10.5733/afin.050.0103
    Published: 1 June 2009
    KEYWORDS
    Arachnida
    Carnian
    Chelicerata
    fossils
    Mesozoic
    palaeontology
    Palaeozoic
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