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The Fossil Arachnida

  • Petar Beron
Chapter
Part of the Monographiae Biologicae book series (MOBI, volume 94)

Abstract

Concerning the fossil orders, Dunlop (1996a, b) presents a revised concept of the formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil orders of Arachnida. The subclasses of Petrunkevitch (1949) have been rejected, Anthracomartida – synonymized with Trigonotarbida. It was concluded that the position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida, according to Dunlop (1996a, b), represents misidentified opilionids.

Keywords

Arachnida Fossils Petrunkevitch Pocock Dunlop Selden Amber Devonian 

Concerning the fossil orders, Dunlop (1996a, b) presents a revised concept of the formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil orders of Arachnida. The subclasses of Petrunkevitch (1949) have been rejected, Anthracomartida – synonymized with Trigonotarbida. It was concluded that the position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida, according to Dunlop (1996a, b), represents misidentified opilionids.

According to the table in the paper of Dunlop et al. (2008), by March 2008, there were 1593 spp. of fossil Arachnida, including 111 Scorpiones, 25 Opiliones, 38 Pseudoscorpiones, 5 Solifugae, 294 “Acari,” 1 Palpigradi, 15 Ricinulei, 979 Araneae, 9 Amblypygi, 7 Uropygi, and 4 Schizomida (actually the Schizomida are 6). Extremly rich fossil fauna has been found in the Burmese amber. “Fossils from the mid-Cretaceous (c.99 Ma) Myanmar (Burma) amber include all extant orders of Arachnida, including the earliest representatives of Schizomida, Parasitiformes, and Palpigradi…The most abundant and diverse arachnid order is the Araneae, with 38 families, 93 genera, and 165 species recorded tom date” (Selden and Dong Ren 2017).

Going through the recent analytical work on the position of different (recent) orders in the system of Arachnida, the following summary appears:

Palpigradi

Palaeokoenenia mordax Rowland et Sissom, 1980, from the late Tertiary (?Pliocene) of Arizona and the Electrokoenenia yaksha Engel et Huang from the Cenomanian Burmese amber remain the only known fossil Palpigrades.

Solifugae

Only five fossil species of these large arachnids, living mostly in dry areas, are known (from Poland, USA, Brazil, and two from the Baltic and Dominican amber, respectively) (Poinar and Santiago-Blay 1989; Dunlop et al. 2004).

Ricinulei

Selden (1992) revised the fossil ricinuleids, describing 15 valid fossil species in 4 genera and 2 families from the Pennsylvanian Coal Measures of Europe and North America.

Amblypygi

Eight valid species of fossil whip spider are accepted by Dunlop et al. (2008): four from Europe and North America; two from Mexican and Dominican amber, respectively; another one from Mexican amber (Poinar and Brown 2004); and one from the Crato Formation in Brazil (Dunlop and Martill 2002). According to Dunlop and Mrugalla (2015), there are already 11 spp. The Late Carboniferous whip spider Graeophonus anglicus Pocock 1911, was redescribed by Dunlop et al. (2007a, b). The species Electrophrynus mirus Petrunkevitch 1971, was considered representing new genus and family, but both have been strongly contested by Armas (2006) and Dunlop and Mrugalla (2015). According to Selden et al. (1991), Amblypygi originated at least by the mid-Devonian (ca. 380 Ma). Presently, there are no Amblypygi living in Europe, and very few are known from North America, north of Rio Grande.

Thelyphonida (Uropygi)

According to Harvey (2003), there are nine species of fossil Uropygi, but after the recalculation of Tetlie and Dunlop (2008) and Dunlop et al. (2008), their number has been reduced to six to seven. Cai and Huang (2017) described from Upper Cretaceous Burmese amber the new genus and species of Thelyphonida Mesothelyphonus parvus- the earliest Thelyphoninae known so far.

Schizomida

Five fossil Schizomida genera have been described so far: Antillostenochrus Armas et Teruel and Rowlandius Reddell et Cokendolpher from the Miocene amber of the Dominican Republic; Calcitro Petrunkevitch from China, Oligocene, and the USA, Pliocene (Calcitronidae); and Onychothelyphonus Pierce and Calcoschizomus Pierce from the USA, Pliocene (Hubbardiidae) (Petrunkevitch 1945; Pierce 1951). From them, only Rowlandius contains extant species.

Scorpiones

The phylogenetic position of the scorpions remains a key question to resolve in studies of Chelicerate evolution (Weygoldt 1998; Dunlop and Braddy 2001). Scorpions are considered since longtime as the oldest known arachnids (Selden 1993a, b). Pocock (1893) divided the Arachnida into Ctenophora (scorpions) and Lipoptena (the remaining Arachnida). Dubinin (1962) unites Scorpiones and Eurypterida in a class (now clade) Scorpionomorpha. As scorpions are known as (sea ?) forms from the Silurian and the Devonian, very different from the extant forms, the dispute is between paleontologists and neontologists. The familial and suprafamilial classification of the scorpions is very controversial (see Soleglad and Fet 2003; Prendini and Wheeler 2005). The extant scorpions are assigned to the Orthosterni – a group or “parvorder” (Soleglad and Fet 2003) – known from the Carboniferous to the present. The number of superfamilies is not definitely established.

Despite their hard cover, the fossil scorpions are relatively few in numbers. Fet et al. (2000), based on the monograph of Kjellesvig-Waering (1986), quote 78 spp. The further transformations and new descriptions from Baltic amber (Lourenco 2012), Burmese amber (Santiago-Blay et al. 2004), etc. increased this number to 111 by 2008 (Dunlop et al. 2008). The table in the last mentioned paper indicates one curious (and unique among Arachnida) particularity of fossil scorpions: 79 spp. are from Paleozoic age, 16 from Mesozoic, and 16 from Cenozoic.

The older concept that scorpions are very primitive (partly because are very ancient) and are the ancestors of the other arachnids is strongly denied by cladists, which find (Shultz 1990) that they are derived arachnids forming a Dromopoda clade together with Opiliones, Solifugae, and Pseudoscorpiones. Some authors (Uchida 1966) still support the primitive character of the scorpions. The cladistic analysis of Dunlop and Braddy (2001) did not provide an undisputable end of the contest between the three main hypotheses: (1) Scorpions are the sister group of all other arachnids; (2) scorpions are derived group of arachnids as part of Dromopoda clade (and perhaps sister group of opilionids); and (3) scorpions are sister group of eurypterids. Support exists for any of the alternative hypotheses, based on morphological and molecular data and their interpretation. We can only admire the perception of the older authors (Pocock, Dubinin), which had in hand much less instruments for comparative analysis.

Gess (2013) published “the earliest record of terrestrial animals in Gondwana” from South Africa. This was the new genus and species Gondwanascorpio emzantsiensis, a scorpion from a Famennian (Late Devonian) formation.

Pseudoscorpiones

In the catalogue of Harvey (1990) were listed 32 valid fossil pseudoscorpions from amber (Burmese, Chinese, Baltic, and Dominican). Schwaller et al. (1991) published the oldest pseudoscorpion from the mid-Devonian of Gilboa, New York State (the first Paleozoic pseudoscorpion). More details could be found in Harvey (1990) and Spahr (1993). Dunlop et al. (2008). After the new data of Hendericks (2005) and Judson (2007), respectively, from Baltic and Dominican amber, the total number of fossil pseudoscorpions amounts on 38 species (Dunlop et al. 2008). According to Harms and Dunlop (2017), overall 16 of the 26 [27?] families of living pseudoscorpions have been documented from fossils and 49 currently valid species are recognized. “The present-day distributions of some group (Faellidae and Pseudogarypidae) is relictual and highlights past extinction events” (Harms and Dunlop 2017). Harvey et al. (2017) described from the Burmese amber (mid-Cretaceous, ca. 90 Mya) the oldest member of Chthoniidae Weygoldtiella plaususHarvey et al.

Opiliones

The fossil record of this important group has been summarized by Dunlop (2007a, b). As we can see from the summary table in Dunlop’s paper, listing all known by this time fossil Opiliones, the fossil record of this group, containing now more than 6500 described species, consists of only 31 entries, part of them unnamed. Eight of them are from the Paleozoic (from Lower Devonian in Scotland to Upper Carboniferous in France). Only two taxa are recognized as genuine Mesozoic harvestmen (from the Lower Cretaceous in Australia and from Upper Cretaceous Myanmar amber). None of them belongs to the modern families. The remaining 21 entries are Cenozoic and belong to the present-day families Trogulidae, Phalangiidae, Sclerosomatidae, Nemastomatidae, Caddidae, Kimulidae, Samoidae, and Cladonychiidae and are even assigned to the modern genera Trogulus, Kimula, Caddo, Dicranopalpus, Leiobunum, Nemastoma, Sabacon, etc. Sometimes these findings cast new light over the present-day distribution of families like Caddidae or Kimulidae and help in restoring the paleodistribution of the Opiliones. Actually, only 25 species are recognized as valid (Dunlop et al. 2008), with only one sp. from the Mesozoic. Ten of the findings come from the Baltic amber, considered to be of Eocene age (some 38–54 million years ago), mostly old information, summarized by Koch and Berendt (1854) and Menge (1854). The modern research in the Dominican amber by Cokendolpher (1986) and Cokendolpher and Poinar (1982, 1998) brought another four species. They belong to the family of Phalangodidae and Kimulidae, still living in the Antillean area. The age of this fauna is not very different from the age of Baltic amber fauna, but the research in the Dominican amber has much shorter history (for the harvestmen since 1978), and there is a lot more to expect.

Only 14 fossil Opiliones species have been recorded before 1955, so the task of Petrunkevitch (1955), who summarized the existing scant information concerning fossil Opiliones, seems to be relatively simple. But it is amazing that these widespread animals are so rare in the fossil record (the reasons are explained by Dunlop, op. cit.). The oldest known Devonian harvestmen look very much like the modern forms, and it is a well-founded guess that there are also Silurian or other pre-Devonian Opiliones-like creatures, which are to be discovered. By Devonian (the Opiliones are considered to be at least 400 million years old), most of the known arachnid orders already existed as terrestrial, air-breathing animals. However, Garwood et al. (2014) described a new fossil suborder and concluded that “The discovery of Tetrophthalmi alters molecular divergence time estimates, supporting Carboniferous rather then Devonian diversification for extant suborders and directly impacting inference of terrestrialization history and biogeography.”

The peculiar suborder Cyphophthalmi is represented by Siro platypedibus Dunlop et Giribet, 2003, a single specimen, described from Tertiary Bitterfeld amber, and by a new genus and species Palaeosiro burmanicum Poinar, 2008, from the Early Cretaceous Burmese amber (Upper Albian (100 to 105 m.y.B.P.)

Araneae

The bulk of Arachnida (the Acari excluded), both living over 40 000 spp. and fossil (979 spp.), belong to the order of spiders. A general review of the spider fossil record is provided by Penney and Selden (2006a, b). The oldest record is from Gilboa, New York (mid-Devonian, 380 Ma) (Selden et al. 1991). According to Dunlop et al. (2008), the vast majority of fossil spiders (c. 820 spp.) originate from amber: 540 spp. from the Baltic amber and 170 spp. from the Dominican amber. There are also findings in the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc. Spiders are described recently also from Cretaceous ambers in Siberia, Canada, Burma, the USA, and Spain (see Dunlop et al. 2008). Significant findings from the Cretaceous occurred also in Siberia and Mongolia (Eskov and Zonshtein 1990) and Spain (Selden and Penney 2003) and from the Crato Formation in Brazil (Selden et al. 2006).

Acari

As tiny creatures, mites are not many in the fossil record. After the publication of Krivolutsky (1973) about finding fossil Oribatida (including some recent genera (Cultroribula, Achipteria) from the Jurassic and Cretaceous deposits in Russia, it was accepted (Hammer and Wallwork 1979) that “families of oribatids that we can recognize today were present during the Jurassic and undoubtedly earlier.” It was proved some time later by Norton et al. (1988) with the finding of oribatids in the terrestrial Devonian deposit near Gilboa, New York.

Describing the second fossil Opilioacarid, Dunlop et al. (2010a) analyzed the status of the fossil Acari by this time. According to them and to Dunlop et al. (2010b), 292 fossil actinotrichid species (including recent forms found as subfossils) have been recorded. Actinotrichids were certainly present by the Early Devonian (ca.410 Ma) and potentially much earlier. There are only 14 fossil species of anactinotrichids, and their record goes back to Cretaceous (Poinar and Buckley 2008) – two opilioacarids, four gamasids, no holothyrids, and eight ticks. The small number of fossil gamasids is really “puzzling.” Most genera of the fossil Gamasida and Ixodida are the same as in the extant fauna. Only in the Burmese amber have been found two new genera of Ixodidae (Cornupalpatum Poinar et Brown, 2003, and Compluriscutula Poinar et Buckley, 2008).

Many years since this time, the fossil record of Oribatida is still poor (Dunlop et al. 2008). According to the table of Dunlop et al. (2008), by March 2008, there were known 294 “Acari.” Obviously, Oribatida are privileged as being preserved in the older rock layers (not amber), because of their hard armor. Such armor, even harder, has also the caeculids, but they are much smaller group, and only two species have been described recently from Baltic and Dominican amber (Eocene). From the Canadian amber has been described as new genus and species Proterythraeus southcotti Vercammen-Grandjean, 1972. From the Baltic amber have been recorded another new genus (Arythaena Menge, 1854) and eight other Erythraeidae species, assumed to belong to the present-day widespread genera Erythraeus, Balaustium, and Leptus. Provided that the material still exists, these identifications should be revised (Zacharda and Krivolutsky 1985).

The fossil Mesostigmata are rare. From the Baltic amber (Eocene, ca. 44–49 Ma), Dunlop, Kontschán, and Zwanzig (2013) recorded Uropodina and Dunlop et al. (2014) Laelapidae of the extant genus Myrmozercon.

Opilioacarida

The first fossil opilioacarid to be described is Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004, from Eocene Baltic amber. It has been attached to the genus Paracarus Chamberlin et Mulaik, 1942, represented by a modern species from Central Asia, and represents the northernmost species of Opilioacarida. The second fossil opilioacarid (? Opilioacarus aenigmus Dunlop, Sempf et Wunderlich, 2010a) was described again from Baltic amber (Eocene, 45–50 Ma). However, it has been assigned tentatively to the now Mediterranean genus Opilioacarus With, 1903.

Paleozoogeography of Arachnida

Important information concerning the past distribution of the Arachnida comes from the amber – Baltic, Dominican, Burmese, Lebanese, Chinese, etc. The amber is of different geological age. Other findings of fossil Arachnida are from the Crato Member of the Santana Formation in Brazil (Martill et al., Eds, 2007), Kazakhstan, etc.

Here are some data showing the differences between the past and present distribution of Arachnida:

Palpigradi

Remaining are the findings of Palaeokoenenia mordax Rowland et Sissom from the late Tertiary (?Pliocene) of the USA (Arizona) and of the Electrokoenenia yaksha Engel et Huang from the Cenomanian Burmese amber (Myanmar).

Ricinulei

The findings of the ancient group in the Pennsylvanian Coal Measures of Europe and North America (Nam.- Steph., 327–290 Ma) (Selden 1992) show a much broader distribution in the distant past. Actually, no ricinuleid lives in the Palearctic and (practically) in the Nearctic (only one sp. Pseudocellus dorotheae from Texas, on the border of Mexico). There are no extant Ricinulei also in Asia, but Wunderlich (2012) described? Poliocera cretacea from Burmese Cretaceous amber.

Scorpiones

There are at least 45 fossil families of scorpions (almost entirely from the suborder Branchioscorpionina).

Solifugae

Fossil species have been described not only from countries, where there are Solifugae even now (USA, Brazil), but also from Poland and the Baltic amber.

Amblypygi

Most of the known fossil Amblypygi come from countries where the order is represented now (Brazil, Mexico, Dominican Republic). However, four species are recorded from Europe and North America (Nam.-Steph., 327–290 Ma).

Thelyphonida (Uropygi)

Different genera from the now existing have been described from the Carboniferous of North America and Europe. Presently, Uropygi are not known from Europe (even closely) and from most of North America.

Schizomida

One representative of the order was found from the Oligocene of China. Presently, the only Schizomida known from this country is Bamazomus siamensis from Hong Kong, within the present range of the order.

Opiliones

There are fossil families of Opiliones, belonging to four recent suborders: Cyphophthalmi (Sironidae), Eupnoi (Phalangiidae), Dyspnoi (Caddidae, Nemastomatidae), and Laniatores. The material from Baltic amber (Eocene, ca. 44–49 Ma) has been presented by Koch and Berendt (1854) and supplemented or corrected by Roewer (1939), Dunlop and Barov (2005), and Dunlop and Mitov (2009). From the Bitterfeld amber (Germany, Oligocene–Chattian, 25.3–25.8 Ma) have been described the first fossil cyphophthalmid Siro platypodibus Dunlop et Giribet, 2003, and many other Opiliones, belonging to extant genera (Lacinius Thorell, Amilenus Martens, Mitostoma Roewer, ?Histricostoma Kratochvil, Caddo Banks, Dicranopalpus Doleschall, and even species like Lacinius erinaceus Staręga, known from Caucasus) (Dunlop and Mitov 2009). It is supposed that Opiliones of the extant genera and even species existed some tens of millions year ago (in the Paleogene). However, the Opiliones are much older and, as specialists say, can be traced back to at least the mid-Paleozoic. With the discovery of Caddo in North Europe, it was supposed even that the now discontinuous species of this genus (Japan and North America) have been spread in the Tertiary much broader. The fact that extant species have been recorded in the Oligocene, more than 20 Ma ago, raises the problem of the pace of evolution. Cokendolpher and Cokendolpher (1982) reexamined the tertiary harvestmen from Colorado.

Araneae

As it was said, the age of the fossil spiders is mostly from the Mesozoic, the oldest known being at 380 Ma (mid-Devonian of Gilboa). The Russian arachnologist Eskov (1984) used the finding of a member of “Gondwanan” family Archaeidae (Jurarchaea zherikhini) in the Upper Jurassic of Kazakhstan, as an argument against the attempts to explain many distributions with Gondwana. The actual distribution of the family (South Africa, Madagascar, Australia, Tasmania, New Zealand, and Patagonia) was considered by Legendre (1977) for “typically Gondwanan.” These spiders are known also from Baltic amber (seven species) and probably their present kind Gondwanan distribution is a relict one from a much more cosmopolitic distribution in the past. About the same time was published also a revision by Forster and Platnick (1984), leaving within Archaeidae only four genera from the Baltic amber, Australia, South Africa, and Madagascar. This does not change very much the conclusions of Eskov. Also, as Wunderlich (1995) pointed, Cyatholipidae are not “representing former parts of the Gondwanaland” (Griswold 1987) but rather a relict distribution of a very wide distribution in the Lower Tertiary period.

Opilioacarida

Paracarus pristinus from Baltic amber has been ascribed by Dunlop, Wunderlich, and Poinar (2004) to the genus Paracarus, represented today only by one species in Central Asia. The other fossil Opilioacarid (Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich, 2010a) is conditionally included in the genus Opilioacarus, living today in the south of Europe (Italy, Greece) and in North Africa.

Trombidiformes

From Canadian amber has been described as new genus and species Proterythraeus southcotti Vercammen-Grandjean, 1972. From the Baltic amber have been recorded another new genus (Arythaena Menge, 1854) and eight other Erythraeidae species. From the Early Cretaceous (Aptian) Crato Formation, Brazil, has been described a new large, fossil mite (Arachnida: Acari, Erythraeoidea), Pararainbowia martilli Dunlop 2007a. Bernini (1991) made general review of the fossil Acarida, known by this time.

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Authors and Affiliations

  • Petar Beron
    • 1
  1. 1.National Museum of Natural History, Bulgarian Academy of SciencesSofiaBulgaria

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