2003. The Journal of Arachnology 31:371±378

THE FIRST CYPHOPHTHALMID (ARACHNIDA, ) FROM BITTERFELD , GERMANY

Jason A. Dunlop: Institut fuÈr Systematische Zoologie, Museum fuÈr Naturkunde der Humboldt-UniversitaÈt zu Berlin, Invalidenstraûe 43, D-10115 Berlin, Germany. E-mail: [email protected] Gonzalo Giribet: Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Avenue, Cambridge, Massachusetts 02138, USA

ABSTRACT. The ®rst fossil cyphophthalmid harvestman, Siro platypedibus new species (Arachnida, Opiliones, Cyphophthalmi), is described from Bitterfeld amber, Sachsen-Anhalt, Germany. The age of this amber is in dispute. Geological studies support a Miocene (20±22 Ma) date for the deposit, but the presence of insect species identical to those in (dated at ca. 35±40 Ma) has led other authors to suggest that the Bitterfeld amber comprises older, redeposited material, contemporary with Baltic in- clusions. Two features in this harvestman fossil are consistent with the Recent genera Siro, Paramiopsalis and Tranteeva: (a) smooth tarsi and metatarsi in legs 1 and 2 and (b) the apparent absence of a dorsal crest on the basal article of the chelicera. Unequivocal autapomorphies of any one of these genera are not clearly preserved in this fossil, but Paramiopsalis is a monotypic Iberian genus, and Tranteeva is a monotypic genus from Bulgaria, while Siro is more diverse and widely distributed, including living rep- resentatives in Central Europe relatively close to the Bitterfeld type locality. For this reason we assign the fossil to Siro. Keywords: Cyphophthalmi, , Siro, taxonomy, paleontology, new species

Fossil harvestmen are rare and their fossil are unconvincing. These Coal Measures fos- record is currently restricted to a few Paleo- sils lack autapomorphies of Cyphophthalmi zoic and Mesozoic examples together with a and are relatively large with long, slender legs more diverse Tertiary record based principally in at least some specimens. on the Florissant Formation and on Baltic and Cyphophthalmi are small to medium-sized, Dominican ; see e.g., Petrunkevitch inconspicuous, almost mite-like, creatures (1955), Cokendolpher & Cokendolpher with short, stubby legs which typically live in (1982) and Selden (1993) for reviews. The soil, leaf litter or caves (Shear 1980). They are majority of the fossil harvestmen have been often regarded as primitive harvestmen and referred to, or strongly resemble members of, recent phylogenetic studies (Shultz 1998; Gi- the Eupnoi and Dyspnoi clades. Among those ribet et al. 1999, 2002) have consistently specimens which have not been formally de- placed them in a basal position, as sister-group scribed there is a very old (c. 340 Ma), but to the remaining opilionids. The fossil record remarkably modern-looking, phalangioid har- provides minimum divergence times for vestman (Wood et al. 1985), which implies for clades, thus the recently published cladograms this group a high degree of morphological predict that the cyphophthalmid lineage conservatism over geological time. Laniatores should go back to at least the mid-Paleozoic, is currently known only from Tertiary ambers, the age of the oldest recorded harvestman (see and all of the Dominican amber harvestmen above). The systematics of the Recent cypho- described thus far are Laniatores (Cokendol- phthalmids have been summarized by the cat- pher & Poinar 1998). The remaining suborder, alogue of Giribet (2000), who recognized 113 Cyphophthalmi, has not previously been re- extant species in 26 genera, and the cladistic corded in the fossil record. Petrunkevitch analysis of generic relationships by Giribet & (1949) claimed that some of the Pennsylva- Boyer (2002). We refer to these publications nian Coal Measures harvestmen distinctly re- for additional background literature on the sembled cyphophthalmids, but his arguments group. In this paper we describe the ®rst fossil

371 372 THE JOURNAL OF ARACHNOLOGY cyphophthalmid, a specimen discovered in the taxa, including some spiders and a well-pre- Berlin collection of inclusions from the Bit- served phalangiid harvestman, and provided terfeld amber deposit of eastern Germany. an overview of the geological setting. Further geological details can be found in FuÈhrmann METHODS & Borsdorf (1986) and Kosmowska-Ceranow- The holotype, and only specimen, was in- icz & Krumbiegel (1989). formally recognized as a cyphophthalmid in Geological setting.ÐHistorically, amber 1989 by Manfred Moritz, the then Curator of has been recovered from a number of sites in in the Zoology Department of the the vicinity of Bitterfeld and the adjacent Mul- Museum fuÈr Naturkunde, Berlin. It was thus de river in the Sachsen-Anhalt region of east- discovered too late to be listed in the sum- ern Germany (see e.g. Kosmowska-Ceranow- mary paper of Schumann & Wendt (1989) on icz & Krumbiegel 1989, ®g. 1). Since 1955 Bitterfeld inclusions. The fossil was not sub- the principal source of amber, almost certainly sequently formally described. The specimen is including the specimen described here, has held in the Section of the Paleon- been the now disused, open-cast Braunkohl tology Department of the Museum fuÈr Natur- mine of Goitsche near Bitterfeld. The amber- kunde Berlin (MB.A.) under the repository producing horizon consists of a series of mas- number 1086. Drawings were prepared with sive, sandy-clay lenses which, according to the aid of a camera lucida attachment and the the local geological terminology, lie between fossil was compared to all extant genera of the Bitterfeld main coal seam and the Breiten- Cyphophthalmi and to most species of Siron- feld seam (Kosmowska-Ceranowicz & Krum- idae (see e.g. Giribet & Boyer 2002, Appen- biegel 1989, ®g. 3). The sand and clay have dix 2). Digital photographs (Figs. 1±2) were been interpreted as representing a period of taken using a JVC Digital Camera KY-F70B marine ingression which was dated on spore mounted on a Leica MZ 12.5 stereomicro- evidence to sporomorph zone IIIA according scope. Series of ca. 20 images were taken at to the German Democratic Republic strati- different focal planes and assembled with the graphic scale for the Tertiary (Krutsch in Bar- dedicated software package Auto-Montage thel & Hetzer 1982). This spore zone corre- 4.01.0085 by Synoptics Ltd. All measure- lates to a lower Miocene (lower Aquitanian) ments are in mm. age. Age of the amber.ÐAmber is notoriously BITTERFELD AMBER dif®cult to date precisely and the Bitterfeld Amber from the Bitterfeld region of Ger- inclusions have been assigned to anything be- many has been known since at least the mid tween an Eocene and a Miocene age. Barthel 17th century; see Kosmowska-Ceranowicz & & Hetzer (1982) interpreted Bitterfeld amber Krumbiegel (1989) for a review. This Bitter- as younger than Baltic amber, dating the for- feld, or Saxon (ϭSaxonian), amber is not as mer at lower Miocene (ca. 22 Ma) based on well known as Baltic or Dominican amber. a combination of the regional geology and mi- The locality was originally kept secret by the crobotany (see above). The inclusion-bearing German Democratic Republic on the suspi- pieces unequivocally lie in situ in strata as- cion that it was associated with uranium de- sociated with the Miocene coal seams (M. posits (M. Barthel, pers. comm., 2003), but in Barthel, pers. comm., 2003). Wunderlich recent years it has seen increased activity from (1983) suggested that the Bitterfeld amber amateur collectors. It nevertheless contains a was merely part of the Baltic amber complex diverse range of inclusions, provisionally list- and thus implicitly late Oligocene/Eocene (ca. ed by Schumann & Wendt (1989). Although 35±40 Ma) in age. He derived these conclu- there have been numerous papers on the sions from a (very provisional) survey of sim- plants, fungi and, especially, the insect fauna ilar faunal (spider) and ¯oral elements in both (e.g. RoÈschmann 1997; Wagner et al. 2000), ambers. Fuhrmann & Borsdorf (1986) sup- the arachnids remain quite poorly studied with ported the Miocene age and rejected this re- only a few formal species descriptions (e.g. deposition hypothesis. These authors present- Wunderlich 1993). The ®rst concerted study ed a detailed physico-chemical analysis in of the inclusions was by Barthel & Hetzer which they argued that the composition of (1982). These authors ®gured a number of mineralogical species in Bitterfeld amber dif- DUNLOP & GIRIBETÐFOSSIL CYPHOPHTHALMID IN AMBER 373

Figures 1±2.ÐSiro platypedibus new species; the ®rst fossil cyphophthalmid. MB.A. 1086 from Bit- terfeld amber, Sachsen-Anhalt, Germany. 1. Dorso-lateral aspect; 2. Ventro-lateral aspect. Scale bars ϭ 1 mm. fered signi®cantly from that of Baltic amber. region. Their main conclusion was that an Eo- Kosmowska-Ceranowicz & Krumbiegel cene age for the amber-bearing sediments (1989) analyzed the heavy mineral composi- could be ruled out, but that the sediments tion in the amber-bearing sediments and com- could represent redeposited Oligocene mate- pared them to strata of known age in the same rial. Note that these results apply to the sedi- 374 THE JOURNAL OF ARACHNOLOGY ments and not the amber itself. These authors fortunately there is a series of internal frac- also noted a degree of color variation in the tures in the matrix around the body, which, Bitterfeld amber which is not seen in Baltic along with numerous bubbles, obscures some amber, again implying that these ambers come details. There also appears to be some sort of from different sources. foreign body (a spore?) directly underneath However, RoÈschmann (1997, and references the which partly covers the coxo-ster- therein) has challenged the assumption that nal region and ventral surface of the opistho- the Baltic and Bitterfeld ambers are funda- soma. Grinding the amber would not clearly mentally different, based on a detailed com- reveal the entire ventral surface without dam- parison of insect faunas; particularly Diptera. aging the legs. Some parts of the legs are en- In this study, 14 species of fossil Sciaridae crusted with a refractant substance, but the un- were found to be common to both ambers and, derlying morphology is still visible. The fossil in general, the Baltic and Bitterfeld ¯y faunas is undoubtedly a cyphophthalmid and, like score similarly on various ecofaunistic indi- modern examples, it has a small, compact and ces. The longevity of arthropod species has densely tuberculate body (Murphree 1988), been estimated at 2.5±7 million years, thus if short, stubby legs with somewhat swollen and there really is an age difference of at least 13 rounded podomeres, and a tarsus which ends million years between the Baltic and Bitter- in a single claw. The animal is complete and feld ambers then, assuming these longevity es- the body is c. 2 mm long, which is within the timates are accurate and widely applicable, we range of modern species. There is no project- would not expect to ®nd identical species in ing adenostyle on the fourth tarsus; therefore, them. RoÈschmann thus implied an older age since the specimen looks mature due to the and redeposition of the Bitterfeld material and degree of sclerotization, we interpret it as a suggested that the Baltic and Bitterfeld amber- female. producing forests were of a rather similar age; The carapace is mostly hidden in the ma- unlike the host sediments of the amber pieces. trix. There is no evidence for the presence of It should be added that Wagner et al. (2000) eyes. Ozopores on a pair of raised tubercles gave an Eocene (50 Ma) date for both Bitter- are characteristic for cyphophthalmids (Giri- feld and Baltic amber, but the two papers they bet et al. 2002) and one of these paired, cite in support of this (Noonan 1986, 1988) raised, horn-like structures, sometimes are essentially biogeographical studies, one of termed ozophores, can be seen on one side which mentions a date for Baltic amber of 30 of the fossil (Figs. 2, 4). It is deep within the Ma and neither of which mention the Bitter- matrix and details of morphology are lacking. feld deposit! This illustrates the danger of Its apparently dorso-lateral position is con- poorly-justi®ed dates becoming perpetuated in sistent with the `type 2' orientation (cf. Jub- the literature. It is beyond the scope of the erthie 1970; Giribet & Boyer 2002). On the present paper to resolve the problems of dat- other side (Figs. 1, 3) the ozophore region is ing the Bitterfeld . Rikkinen & Poinar obscured, almost as if a secretion from it has (2000) used the younger, Miocene age of formed an ill-de®ned bubble in the matrix around 20±22 Ma, but Poinar (pers. comm., over the antero-lateral corner of the carapace. 2002) now accepts the idea that Bitterfeld am- The opisthosoma expresses at least seven ter- ber could be equivalent to Baltic amber, but gites dorsally, each densely tuberculate with perhaps originating from a different geograph- circular tubercles and separated from adja- ical source to the Baltic amber forest. Our fos- cent tergites by a narrow band of non-tuber- sil can thus be constrained to, at best, an age culate cuticle. of between 20±40 Ma. The chelicerae are mostly obscured behind other limbs, but the dorsal surface of the basal MORPHOLOGICAL INTERPRETATION article can be seen. Proximally, there is no ev- The fossil is preserved in an oval piece of idence for a dorsal crest (ϭ dorsal ridge) relatively clear and translucent amber. Its ori- which is characteristic of many living cypho- entation makes it primarily visible from both phthalmids (e.g. Giribet & Boyer 2002, Figs. sides in a dorso-lateral and ventro-lateral view 1±3). Characters relating to the ventral surface respectively, thus details of the legs are easier of the second article or the movable ®nger of to see than those of the body (Figs. 1±4). Un- the chela are equivocal. The pedipalps are DUNLOP & GIRIBETÐFOSSIL CYPHOPHTHALMID IN AMBER 375 slender with elongate podomeres which are tibia 0.29, tarsus 0.27. Leg article lengths as distally densely setose. The legs are robust follows. Leg 1: femur 0.75, patella 0.32, tibia and tend to converge distally beneath the an- 0.56, metatarsus 0.30, tarsus 0.55; total length imal. Like the body, the legs are densely tu- c. 2.5. Leg 2: femur 0.65, patella 0.32, tibia berculate. The leg tuberculation tends to be 0.41, metatarsus 0.29, tarsus 0.50; total length formed from more elongate, oval tubercles, c. 2.2. Leg 3: femur 0.45, patella 0.33, tibia especially on the dorsal surface of the articles 0.33, metatarsus 0.30, tarsus 0.47; total length in legs 3 and 4. Signi®cantly, this tubercula- c. 1.9. Leg 4: femur 0.58, patella 0.30, tibia tion is not apparent on the metatarsus and tar- 0.38, metatarsus 0.26, tarsus 0.53; total length sus of legs 1 and 2 (Figs. 1±4). The leg for- c. 2.1. Legs tuberculate like body, but tuber- mula is, from longest to shortest:1243.The culation absent on distal articles of legs 1 and patellae and tibiae of leg 4 are notably swollen 2. All legs end in single, smooth, hook-shaped and ovate in appearance. The legs preserve a claw, c. one-third the length of the tarsus. number of setae, which become more numer- Remarks.ÐThis specimen represents the ous on the more distal articles. There is a par- ®rst, and so far only, fossil record of Cypho- ticular concentration of setae on the ventral phthalmi. Their rarity as fossils is undoubtedly surface of the distal tarsus near the origin of due to a combination of their unmineralized cu- the claw, but not forming a distinct solea. All ticle, small size and cryptic, soil-living habits; legs end in a single, smooth, hook-shaped tar- all of which reduce their chances of preserva- sal claw. tion. Amber nevertheless has the potential to trap soil or leaf-litter organisms. Schumann & SYSTEMATIC PALEONTOLOGY Wendt's (1989) faunal list for Bitterfeld amber includes various elements of the soil meso- and Suborder Cyphophthalmi Simon 1879 macrofauna such as nematodes, isopods, oriba- Family Sironidae Simon 1879 tid mites, collembollans and myriapods. Genus Siro Latreille 1796 Although there has been some northward Siro platypedibus new species drift of continental Europe by a few degrees Figs. 1±4 over the last 40 million years, our fossil still Type and only material.ÐMB.A. 1086. probably represents the most northerly record Holotype and only known specimen. Bitter- of European Sironidae, c. 51±52Њ N. The most feld amber. From the site of the Goitsche (or northerly distributed extant species in Europe Goitzsche) Mine, near Bitterfeld, Sachsen-An- is Siro carpaticus Rafalski 1956 which occurs halt, Germany (c. 51Њ36ЈN, 12Њ22ЈE). Tertiary in the Carpathian mountains of south-eastern (?Oligocene±Miocene) in age. Specimen also Poland (c. 40Њ08ЈN). The amber fossil sug- bears the identi®cation number ``Ser. 15/4''. gests that Siro was at one stage more wide- Etymology.ÐFrom the tall, ¯attened patel- spread in Europe and previously occurred fur- la and tibia in leg 4. ther north than its present geographical range. Diagnosis.ÐOrnamentation of legs as in the Two other sironid species occur at high lati- genera Siro, Paramiopsalis and Tranteeva with tudes in North America, S. acaroides (Ewing all tarsi and metatarsi 1 and 2 lacking the nor- 1923) in Washington, up to 47Њ50ЈN, and S. mal pattern of granulation seen in the other po- kamiakensis (Newell 1943) in Washington and domeres. Articles of legs compressed laterally Idaho, c. 47Њ50ЈN. with patella and tibia 4 becoming enlarged and ¯attened. Claws large. The appendages of this AFFINITIES specimen are of particular interest and may in- The specimen described here lacks ventral dicate some special type of habitat. teeth on the claw of leg 2, a character which Description.ÐComplete female cypho- excludes placement in the following genera: phthalmid. Total body length c. 2, but anterior Neogovea Hinton 1938, Huitaca Shear 1979, obscured in matrix. Body with tuberculate or- Metagovea Rosas Costa 1950, Paragovia nament, darker than legs with at least seven Hansen 1921, Troglosiro Juberthie 1979 and clearly de®ned opisthosomal tergites. Prosoma Metasiro Juberthie 1961 (see Giribet & Boyer with anterolateral ozophore. Chelicerae most- 2002). Two preserved features (Figs. 1±4) are ly obscured, but basal article 0.65 long. Ped- of particular interest: (a) the absence of tu- ipalps slender, podomere lengths: patella 0.33, bercular ornament on the metatarsus and tar- 376 THE JOURNAL OF ARACHNOLOGY

Figures 3±4.ÐInterpretive drawings of the specimens shown in Figs. 1, 2. Abbreviations: ch ϭ chelic- era, cx ϭ coxae, in ϭ unidenti®ed inclusion beneath the animal, op ϭ opisthosoma, oz ϭ ozophore, pa ϭ pedipalp, pt ϭ patella, ti ϭ tibia. Legs numbered from 1 to 4. Note the lack of tuberculation at the distal ends of legs 1 and 2 and the shape of the patella and tibia on leg 4. Scale bar ϭ 0.5 mm.

sus of legs 1 and 2 and (b) the apparent ab- tween the genera Siro, Paramiopsalis and sence of a dorsal crest on the basal article of Tranteeva is dif®cult based on the preserved the chelicera. Both of these characters are con- morphology. They are differentiated from sistent with the extant genera Siro, Paramiop- each other on characters relating to the corona salis Juberthie 1962 and Tranteeva KratchovõÂl analis at the posterior end of the opisthosoma, 1958 (see e.g. Juberthie 1970, 1991; Giribet fusion of the coxae of legs 2 and 3 (in Par- & Boyer 2002) all of which belong to the amiopsalis), the shape of the palpal trochanter, Laurasian family Sironidae. The distinct lack the length of the appendages, and the type of of ornamentation on metatarsi 1 and 2 is also adenostyle. All these characters are either ab- found in the Japanese genus Suzukielus Jub- sent or equivocally preserved in the fossil and erthie 1970, but it has a dorsal crest on the further preparation by grinding the amber is basal article of the chelicera. The type 2 po- unlikely to reveal them. The appendages of S. sition of the ozophore is also consistent with platypedibus are of particular interest because these taxa, although this character is seen in they have long claws as in some of the trog- other genera too. However, it indicates that the lobiontic species of Siro and Tranteeva, but fossil does not belong to the sironid genera the legs in the fossil, instead of being elon- Parasiro Hansen & Sùrensen 1904 or Odon- gated, are compressed laterally. This is remi- tosiro Juberthie 1961, which have type 1 niscent in a certain way of the appendages of ozophores inserted in the margin of the cara- the equatorial African genus Ogovea Roewer pace. Resolving the position of the fossil be- 1923 which may show fossorial behavior. DUNLOP & GIRIBETÐFOSSIL CYPHOPHTHALMID IN AMBER 377

Figure 4.ÐSame as Fig. 3.

Paramiopsalis is a monotypic genus, cur- reported to measure between 2.0 and 2.5 mm. rently restricted to the north-west Iberian Pen- Among the European species, Siro gjorgjevici insula (Juberthie 1962; Giribet 2000). Tran- HadzÏi 1933, S. teyrovskyi KratochvõÂl 1938 and teeva is also monotypic and restricted to some Tranteeva paradoxa KratochvõÂl 1958 measure caves in Bulgaria (KratochvõÂl 1958; Juberthie between 2.0 and 2.5 mm in length. 1991), although there are doubts about the va- lidity of the genus, which seems to be an apo- ACKNOWLEDGMENTS morphic form of the Balkan sironid clade We are very grateful to Manfred Moritz for (Juberthie 1991). By contrast, Siro is more di- information about this specimen, to Christian verse and widely distributed, containing 23 Neumann (MB.A.) for access to material in extant species spread across Europe, Turkey his care, to Jen Fogarty (MCZ) for assistance and the USA. These records include Central with the digital imaging and Manfred Barthel European taxa (Austria, Slovakia, Poland) (Berlin) for information about the locality. We whose, albeit often localized, distribution is also thank Bill Shear, Dave Penney, James relatively consistent with the type locality of Cokendolpher and George Poinar Jr. for valu- the Bitterfeld amber in eastern Germany. We able comments. see no characters in the fossil which would justify the creation of a new genus and for LITERATURE CITED biogeographical reasons we tentatively assign Barthel, M. & H. Hetzer. 1982. Bernstein-Inklusen our fossil to the more widespread Siro and aus dem MiozaÈn des Bitterfelder Raumes. Zeit- suggest that this genus may have inhabited schrift fuÈr angewandte Geologie 28:314±336. Central Europe since at least the mid-Tertiary. Cokendolpher, J.C. & J.E. Cokendolpher. 1982. Re- While most members of the Laurasian Siron- examination of the Tertiary harvestmen from the idae are smaller than 1.8 mm, a few species are Florissant Formation, Colorado (Arachnida: Op- 378 THE JOURNAL OF ARACHNOLOGY

iliones: Palpatores). Journal of Paleontology 56: eastern North America and Europe as shown by 1213±1217. insects. Memoirs of the Entomological Society Cokendolpher, J.C. & G.O. PoinarJr. 1998. A new of Canada 144:39±53. fossil harvestman from am- Petrunkevitch, A.I. 1949. A study of Palaeozoic ber (Opiliones, Samoidae, Hummelinckiolus). Arachnida. Transactions of the Connecticut Journal of Arachnology 26:9±13. Academy of Arts and Science 37:69±315. FuÈhrman, R. & R. Barsdorf. 1986. Die Bernstei- Petrunkevitch, A.I. 1955. Arachnida. Pp. 42±162. narten des UntermiozaÈns von Bitterfeld. Zeit- In Treatise on Invertebrate Paleontology, Part P, schrift fuÈr angewandte Geologie 32:309±316. Arthropoda 2 (R. C. Moore, ed.). Geological So- Giribet, G. 2000. Catalogue of the Cyphophthalmi ciety of America and University of Kansas Press, of the world (Arachnida, Opiliones). Revista Ib- Lawrence. eÂrica de AracnologõÂa 2:49±76. Rafalski, J. 1956. Opis Siro carpaticus sp. n. wraz Giribet, G. & S.L. Boyer. 2002. A cladistic analysis zuwagami o morfologii i systematyce Cypho- of the cyphophthalmid genera (Opiliones, Cypho- phthalmi (Opiliones). Sprawozdania PozanÂskiego phthalmi). Journal of Arachnology 30:110±128. Towarzystwa PrzjacioÂø Nauk 1:49±52. Giribet, G., G.D. Edgecombe, W.C. Wheeler & C. Rikkinen, J. & G.O. PoinarJr. 2000. A new species of Babbitt. 2002. Phylogeny and systematic posi- resinicolous Chaenothecopsis (Mycocaliciaceae, tion of Opiliones: a combined analysis of chelic- Ascomycota) from 20 million year old Bitterfeld erate relationships using morphological and mo- amber, with remarks on the biology of resinicolous fungi. Mycological Research 104:7±15. lecular data. Cladistics 18:5±70. È Giribet, G., M. Rambla, S. Carranza, J. BagunÄaÁ, M. RoÈschmann, F. 1997. Okofanistischer Vergleich von Nematoceren-Faunaen (Insecta; Diptera: Sciari- Riutort & C. Ribera. 1999. Phylogeny of the dae und Ceratopogonidae) des Baltischen und order Opiliones (Arthropoda) inferred SaÈchsischen Bernsteins (TertiaÈr, OligozaÈn-Mioz- from a combined approach of complete 18S and aÈn). PalaÈontologische Zeitschrift 71:79±87. partial 28S ribosomal DNA sequences and mor- Schumann, H. & H. Wendt. 1989. Zur Kenntnis der phology. Molecular Phylogenetics and Evolution tierischen Inklusen des SaÈchsischen Bernsteins. 11:296±307. Deutsche entomologische Zeitschrift, N.F. 36: Â Juberthie, C. 1962. Etude des opilions cyphophthal- 33±44. mes Stylocellinae du Portugal. Description de Selden, P.A. 1993. Fossil arachnidsÐrecent ad- Paramiopsalis ramulosus gen. n., sp. n. Bulletin vances and future prospects. Memoirs of the du MuseÂum National d'Histoire Naturelle 34: Queensland Museum 33:389±400. 267±275. Shear, W.A. 1980. A review of the Cyphophthalmi Juberthie, C. 1970. Les genres d'opilions Sironinae of the United States and Mexico, with a proposed (Cyphophthalmes). Bulletin du MuseÂum Nation- reclassi®cation of the suborder (Arachnida, Opil- al d'Histoire Naturelle 41:1371±1390. iones). American Musuem Novitates 2705:1±34. Juberthie, C. 1991. Sur Trenteeva paradoxa (Kra- Shultz, J.W. 1998. Phylogeny of Opiliones (Arach- tochvil, 1958), opilion troglobie et les opilions nida): an assessment of the ``Cyphopalpatores'' cyphophthalmes de Bulgarie. MeÂmoires de Bios- concept. Journal of Arachnology 26:257±272. peÂologie 18:263±267. Simon, E. 1879. Les Arachnides de France VII. Kosmowska-Ceranowicz, B. & G. Krumbiegel. Contenant les ordes des Chernetes, Scorpiones et 1989. Geologie und Geschichte des Bitterfelder Opiliones. Paris. Bernsteins und anderer fossiler Harze. Hallesch- Wagner, R., C. Hoffeins, & H.W. Hoffeins. 2000. es Jahrbuch fuÈr Geowissenschaften 14:1±25. A fossil nymphomyiid (Diptera) from the Baltic KratochvõÂl, J. 1958. JeskynnõÂ sekaÂeÁi Bulharska and Bitterfeld amber. Systematic Entomology 25: (Cyphophthalmi a Laniatores) [Die HoÈhlenwe- 115±120. berknechte Bulgariens (Cyphophthalmi und Lan- Wood, S.P., A.L. Panchen & T.R. Smithson. 1985. iatores)]. Acta Academiae scientiarum cechos- A terrestrial fauna from the Scottish Lower Car- lovenicae basis brunensis 30:372±396. boniferous. Nature 314:355±356. Latreille, P.A. 1796. PreÂcis des characteÁres geÂneÂ- Wunderlich, J. 1983. Zur Konservierung von Bern- riques des insectes, disposeÂs dans un ordre na- stein-EinschluÈssen und uÈber den ``Bitterfelder Bernstein''. Neue Entomologische Nachrichten turel. PreÂvot, Paris. 4:11±13. Murphree, C.S. 1988. Morphology of the dorsal in- Wunderlich, J. 1993. Die ersten fossilien Becherspin- tegument of ten opilionid species (Arachnida, nen (Fam: Cyatholipidae) in Baltischen und Bitter- Opiliones). Journal of Arachnology 16:237±252. felder Bernstein (Arachnida: Araneae). Mitteilun- Noonan, G.R. 1986. Distribution of insects in the gen aus dem Geologisch-PalaÈontologisch Institut northern hemisphere: continental drift and epi- der UniversitaÈt Hamburg 75:231±241. continental seas. Bulletin of the Entomological Society of America 32:80±84. Manuscript received 24 January 2003, revised 7 Noonan, G.R. 1988. Faunal relationships between April 2003.