First fossil Mecysmaucheniidae (Arachnida, , Araneae), from Lower (uppermost Albian) amber of Charente-Maritime, France

Erin E. SAUPE Paul A. SELDEN Paleontological Institute, University of Kansas, Lindley hall, 1475 Jayhawk boulevard, Lawrence, KS 66045 (USA) [email protected] [email protected]

Saupe E. E. & Selden P. A. 2009. — First fossil Mecysmaucheniidae (Arachnida, Chelicerata, Araneae), from Lower Cretaceous (uppermost Albian) amber of Charente-Maritime, France. Geodiversitas 31 (1) : 49-60.

ABSTRACT Th e fi rst known fossil mecysmaucheniid , Archaemecys arcantiensis n. gen., n. sp., is described, from Lower Cretaceous (Upper Albian) amber of Charente- Maritime, France. Th is is the fi rst fossil spider to be formally described from French Cretaceous amber and extends the geological record of Mecysmauche- niidae back into the Cretaceous, the family having previously been known only from the Recent. Th e fossil diff ers from other Mecysmaucheniidae in having four, rather than two spinnerets, so it can be considered plesiomorphic with KEY WORDS respect to modern members of the family in this character. Th e amber of the Arachnida, Archingeay-Les Nouillers area is uniquely considered to have a largely preserved Araneae, Mecysmaucheniidae, litter fauna and our specimen corroborates this hypothesis. , and now amber, their sister group the Mecysmaucheniidae, have been found as fossils solely in Cretaceous, the northern hemisphere, yet their Recent distributions are entirely southern France, new genus, hemisphere (Gondwanan). Th e fi nd suggests a former pancontinental distribu- new species. tion of Mecysmaucheniidae.

GEODIVERSITAS • 2009 • 31 (1) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 49 Saupe E. E. & Selden P. A.

RÉSUMÉ Premier Mecysmaucheniidae fossile (Arachnida, Chelicerata, Araneae), de l’ambre crétacé inférieur (Albien terminal) de Charente-Maritime, France. Le premier fossile connu d’une araignée mécysmaucheniidée, Archaemecys arcan- tiensis n. gen., n. sp., est décrit de l’ambre crétacé inférieur (Albien terminal) de Charente-Maritime, France. Il s’agit de la première araignée fossile décrite de l’ambre crétacé de France, qui étend le registre géologique des Mecysmauche- niidae jusqu’au Crétacé, la famille étant jusque-là connue seulement dans l’Ac- tuel. La présence de quatre fi lières au lieu de deux le distingue de tous les autres MOTS CLÉS Mecysmaucheniidae et peut être considérée comme plésiomorphique. L’ambre Arachnida, d’Archingeay-Les Nouillers est supposé unique pour avoir largement préservé la Araneae, Mecysmaucheniidae, faune de litière et notre spécimen corrobore cette hypothèse. Les Archaeidae, et ambre, maintenant leur groupe frère, les Mecysmaucheniidae, ont leurs fossiles repré- Crétacé, sentés uniquement dans l’hémisphère nord, alors que leur distribution moderne France, genre nouveau, est entièrement sud-hémisphérique (Gondwana). Cette découverte suggère une espèce nouvelle. ancienne distribution pancontinentale des Mecysmaucheniidae.

INTRODUCTION (1984) enlarged the (originally it only included the Huttoniidae Simon, 1893, (Araneae) are an extremely diverse order Th orell, 1870 and of , with 40 462 extant species in 3694 Th orell, 1873) to include the Archaeidae (and genera and 109 families (Platnick 2009). Th ey therefore the Mecysmaucheniidae), removing are characterized by numerous synapomorphies, them from the araneoids. Th ey also signifi cantly including the presence of spinnerets on the abdo- enlarged the superfamily by including the Micro- men for producing silk, naked fangs and associated pholcommatidae Hickman, 1944, Mimetidae venom glands (in most species), two body regions, Simon, 1881, Pararchaeidae Forster & Platnick, eight legs, and pedipalps modifi ed in the male for 1984 and Holarchaeidae Forster & Platnick, 1984 sperm transfer. on the basis of two diagnostic characters: cheliceral Th e Mecysmaucheniidae, a relatively small fam- peg teeth and a raised cheliceral gland. Although ily within the Araneae, is composed of seven some accepted Forster & Platnick’s revision (Cod- genera and 25 known species (Platnick 2009). dington & Levi 1991; Coddington et al. 2004), Th ey are small, haplogyne, ecribellate spiders others have contested the arrangement. According found in New Zealand and southern parts of to Schütt (2000), Micropholcommatidae and Mi- South America (primarily Chile and Argentina) metidae should be placed within the Araneoidea, (Jocqué & Dippenaar-Schoeman 2006). Th e type although the placement of the Archaeidae and genus Mecysmauchenius was fi rst described by Mecysmaucheniidae in Palpimanoidea was still Simon (1884) as a member of the Archaeidae dubious at best. Griswold et al. (2005) agreed with Koch & Berendt, 1854. Mecysmaucheniids are Schütt in that the Mimetidae should belong to distinguished from other spiders by the pres- the Araneoidea; however, they claimed that many ence of chelicerae originating from a foramen in of the palpimanoid familial placements are still the carapace and only two spinnerets (Forster & debatable. Others have suggested that peg teeth Platnick 1984). Mecysmaucheniids belong to are homoplasic (Schütt 2000; Coddington et al. the superfamily Palpimanoidea, which has had a 2004) and therefore should not serve as one of the long and controversial history. Forster & Platnick defi ning characters of the Palpimanoidea.

50 GEODIVERSITAS • 2009 • 31 (1) A new Mecysmaucheniidae (Arachnida, Araneae) from Cretaceous French amber

Th e archaeids were fi rst described from Baltic fl owed directly onto the ground (Néraudeau et amber fossils by Koch & Berendt (1854) and were al. 2002). Th e fossil assemblage of Archingeay is considered an extinct group for some 30 years un- mainly composed of insects, but other til extant species were found in Madagascar. Th e are also present (Néraudeau et al. 2002; Perrichot mecysmaucheniids were described soon after from et al. 2007). Nine percent of the total specimens in Chile and were placed as a genus under inclusions in Albian French amber are arachnids the Archaeidae (Simon 1884). Mecysmaucheniids (Perrichot et al. 2007). Hitherto, no spider fos- were considered a subfamily of the Archaeidae by sils have been described from the Cretaceous of Simon (1895), and some people continued to hold France, although they were referred to by Schlüter this view (see Wunderlich 1986, 2004; Eskov 1987, (1978) and Néraudeau et al. (2002), and mem- 1992). Lehtinen (1967) suggested the subfamily bers of the family Zodariidae were mentioned as be raised to family level, which would include being present by Perrichot (2004) and Perrichot Mecysmauchenius Simon, 1884, Pararchaea For- et al. (2007). ster, 1955 and Zearchaea Wilton, 1946. Although Cretaceous spiders are relatively rare but have Lehtinen’s placement of the Pararchaea and Zear- been described from Siberia (Eskov & Zonshtein chaea was contested by Forster & Platnick (1984), 1990; Eskov & Wunderlich 1994), New Jersey these authors retained the Mecysmaucheniidae as (Penney 2002, 2004a), the Isle of Wight (Selden a distinct family (see Forster & Platnick 1984 for 2002), Lebanon (Penney & Selden 2002; Penney a detailed taxonomic history of the archaeids and 2003a; Wunderlich & Milki 2004), Canada (Pen- related taxa). ney 2004c; Penney & Selden 2006), Myanmar Th ere has been no formal description of a fossil (Grimaldi et al. 2002; Penney 2003b, 2004b, mecysmaucheniid to date. Eskov (1987) tentatively 2005), Botswana (Rayner & Dippenaar-Schoe- assigned Archaea conica (see Koch & Berendt 1854: man 1995), Brazil (Mesquita 1996; Selden et al. fi g. 10) to the subfamily Mecysmaucheniinae because 2002, 2006), Australia (Jell & Duncan 1986) of its short chelicerae and only slightly elevated ce- and Spain (Selden 1989, 1990; Selden & Penney phalic region of the carapace, even though the Baltic 2003; Penney 2006; Penney & Ortuño 2006). amber type specimen had been lost. Eskov (1992) Here, we provide the fi rst description of a fossil later created a new genus, Baltarchaea, for A. conica; mecysmaucheniid, from Cretaceous (Late Albian) the species was listed under Mecysmaucheniidae in French amber. Living mecysmaucheniids are litter- Penney (2003b: table 1). A description of a fossil dwellers, like most of the fauna from Archingeay mecysmaucheniid was said to be in preparation amber, but are confi ned to South America and in Eskov & Golovatch (1986), but no such paper New Zealand. Th e fi nd of a mecysmaucheniid in resulted, and the designation was likely changed to Cretaceous Laurasia suggests a more widespread an archaeid (see Eskov 1987). distribution for this family in the Mesozoic than Lacroix (1910) was the fi rst to describe Cretaceous today. amber from France, but it was not until the 1970s that extensive study of the fossiliferous material in French ambers was undertaken (Perrichot et METHODS al. 2007). Th e most fossiliferous French amber deposit from the Cretaceous is the Archingeay- Les Th e specimen was cut, polished and encased in clear Nouillers (herein referred to simply as Archingeay) resin before the authors received it. Drawings were locality (Perrichot et al. 2007). Th e amber from done under both incident and transmitted light Archingeay is Late Albian in age and is unique in with a camera lucida attached to a Leica MZ16 that a large percentage of the inclusions represent stereomicroscope. Drawings were scanned and re- litter fauna (Néraudeau et al. 2002). It is thought traced using Adobe Illustrator. Photographs were that the resin was exuded from a member of the taken with a Canon Digital Rebel XTi attached to plant family Araucariaceae, and the abundant resin the microscope and manipulated in Adobe Pho-

GEODIVERSITAS • 2009 • 31 (1) 51 Saupe E. E. & Selden P. A.

toshop. Figure 1D was taken with a Leica DFC420 C ps prosoma; camera attached to a Leica DM 2500 microscope; pt patella; Leica Application Suite software was used to take RC right chelicera; sr spiracle; a multifocus z-stack so as to achieve sharp focus st sternum; throughout the image. Measurements were made T trichobothrium; using an ocular graticule. ti tibia; Microtomography at the European Synchrotron tr trochanter; Radiation Facility (ESRF) in Grenoble, France ts tarsus; was attempted; however, the competing mediums 2-4 walking legs 2-4. within the amber piece (air, resin, arthropod cuticle, All measurements are in mm. glue, and Canada balsam used for mounting) were problematical, and obtaining an image proved to be impossible. In order to prepare the specimen PRESERVATION AND for microtomography, it was removed from the MORPHOLOGICAL INTERPRETATION clear resin by gentle heating on a hot plate. Excess amber was cut away from the specimen so as to Th e specimen is preserved in a piece of cloudy, increase imaging ease; during this process, a small light orange amber; approximately 4.5 × 3.5 mm. portion of the opisthosoma was unintentionally Prior to receipt by the authors, the piece of am- removed. ber containing the specimen had been cleaved in Fortuitously, removal of the specimen from the half to the left of the abdomen and subsequently resin allowed for improved views of diffi cult to glued back together. Small air bubbles and pieces observe areas (i.e. we were able to observe cheli- of organic material are present. Th e opisthosoma ceral peg teeth). Th e specimen was mounted on of the spider is translucent and the spinneret at- a pin on a microscope stage so that the piece was tachment points can be viewed internally. Many rotatable at all angles. Further imaging was done features of the specimen are diffi cult to study due with a Leica DFC420 C camera attached to a to the cloudiness of the amber (many, tiny air in- Leica M205 C microscope. Figure 3 is a stack of clusions). No other syninclusions co-occur with three images merged using Helicon Focus software the specimen. (www.heliconsoft.com/heliconfocus.html). Penney (2003b) considered the wrinkled opistho- soma in his Cretaceous Burmese amber specimen ABBREVIATIONS of Afrarchaea grimaldii Penney, 2003 (Araneae, Leg formula (e.g., 1423) indicates the length of Archaeidae) to be the consequence of typical pres- each leg relative to the other legs from longest to the ervational processes that occur in amber. Repeat- shortest (in the example, leg 1 is longest, followed ed mention, however, of a wrinkled abdomen in by leg 4). Abbreviations are as follows: archaeids, and the presence of wrinkles in our ALS anterior lateral spinnerets; specimen, leads us to suggest this may be a specifi c AME anterior median eyes; preservational trait of mecysmaucheniids and closely BL book lung; cl claw; related families. Koch & Berendt (1854: fi gs 9, 10) co colulus; fi gured parallel abdominal folds in their drawings cx coxa; of Baltic amber archaeids, and Eskov (1992), Pen- EF epigastric furrow; ney (2003b) and Wunderlich (2004) all mention fe femur; wrinkles and folds in the abdomens of their archaeid LC left chelicera; amber specimens. mt metatarsus; mx maxilla; Th e organic material has shrunk in most of the op opisthosoma; specimen, as is the rule in amber preservation, PLS posterior lateral spinnerets; leaving a ghostly but faithful outline of the original pp pedipalp; surface. Th e cuticle has shrunk from the distal end

52 GEODIVERSITAS • 2009 • 31 (1) A new Mecysmaucheniidae (Arachnida, Araneae) from Cretaceous French amber

of the leg segments, leaving only the outline of the Forster & Platnick (1984) mentioned that the original surface (Fig. 1E). In the tarsi, however, the spiracle is often sclerotized in mecysmaucheniids, a cuticle is absent from the proximal end of the podo- character observed in our fossil. Archaemecys n. gen., mere, and a slight constriction occurs, indicating like other mecysmaucheniids, does not have tuber- the area was unsclerotized in the living organism cles on the carapace, and the chelicerae in the fossil (a character that is present in both archaeids and are shorter and stouter than those usually found mecysmaucheniids). in archaeids. Archaeids have spatulate hairs on the tibia and metatarsus of leg 1 (Forster & Platnick 1984), but the Mecysmaucheniidae, including our SYSTEMATICS specimen, do not. Additionally, our specimen does not have a femoral hump, as is present in archaeids Order ARANEAE Clerck, 1757 (see Wunderlich 2004). A Pararchaeidae affi nity Suborder Pocock, 1892 can be ruled out because, as mentioned above, Superfamily PALPIMANOIDEA the pararchaeids do not possess the unsclerotized, sensu Forster & Platnick, 1984 constricted area at the bases of the tarsi. Th is aside, our specimen has plumose leg setae, not the serrate Family MECYSMAUCHENIIDAE Simon, 1895 or smooth setae found in pararchaeids, and the legs are stout, unlike the slender legs characteristic of REMARKS pararchaeids. Mecysmaucheniidae are diagnosed as having two spinnerets and chelicerae originating from a foramen in the carapace. Although our specimen has four Genus Archaemecys n. gen. spinnerets and we cannot be certain the chelicerae originate from a foramen in the carapace, we place TYPE SPECIES. — Archaemecys arcantiensis n. gen., n. sp., the specimen in Mecysmaucheniidae on account by present designation and monotypy. of the following characters. Th e unsclerotized, MATERIAL EXAMINED. — Holotype and only known constricted area on the base of the tarsal segments specimen, subadult male, MNHN ARC11R deposited in our specimen appears to be a synapomorphy in the Muséum national d’Histoire naturelle, Paris. uniting the archaeids and mecysmaucheniids (For- ETYMOLOGY. — Th e genus name is derived from the ster & Platnick 1984; H. Wood pers. comm.) and Greek archae, meaning primitive or ancient, and the places our specimen fi rmly among these families. modern genus Mecysmauchenius, which the fossil some- Th e lack of a constricted neck or greatly heightened what resembles. cephalic area (characteristic of Archaeidae), and the STRATIGRAPHIC HORIZON. — Lower Cretaceous, upper- presence of four spinnerets (vs six in archaeids), most Albian, subunit A1s12 (Néraudeau et al. 2002). suggest our specimen should be included within Mecysmaucheniidae. Our reasoning is that, while DIAGNOSIS. — Archaemecys n. gen. can be distinguished from all other genera of Mecysmaucheniidae by the pres- mecysmauche niids have only two spinnerets, this ence of four spinnerets, a strongly sclerotized ring around is a reduction from the plesiomorphic six (i.e. an the spinnerets, and a sclerotized tracheal spiracle. apomorphy). Loss of spinnerets (or their reduction to patches of spigots) has occurred several times REMARKS within the Palpimanoidea (Forster & Platnick 1984) Th e genus diff ers from other mecysmaucheniids and is a character of known polarity. It is likely that by the presence of four spinnerets. All extant reduction is a trend within the Mecysmaucheniidae Mecysmaucheniidae have only two spinnerets, a and the condition in Archaemecys n. gen. represents derived condition. Additionally, Archaemecys n. gen. a stage in the reduction. Archaeids have two spira- has a particularly robust sclerotized ring around the cle openings, unlike the single opening seen in the spinnerets, and its legs are much shorter and stouter mecysmaucheniids and our specimen. Additionally, than in extant mecysmaucheniids.

GEODIVERSITAS • 2009 • 31 (1) 53 Saupe E. E. & Selden P. A.

A C

D

B E

FIG. 1. — Archaemecys arcantiensis n. gen., n. sp. (MNHN ARC11R): A, dorsal view; B, ventral view; C, opisthosoma, notice the pro- nounced wrinkling of the cuticle and the sclerotized rings around the tracheal spiracle and the spinnerets; D, lateral view of the spin- nerets, anterior to the left; the anterior two spinnerets (left) are relatively large, with two segments, while the posterior two spinnerets (right) are smaller; E, close-up of metatarsus and tarsus third leg segment showing the unsclerotized portion at the base of the tarsus (arrowed). Scale bars: A, B, D, E, 0.5 mm; C, 0.2 mm.

54 GEODIVERSITAS • 2009 • 31 (1) A new Mecysmaucheniidae (Arachnida, Araneae) from Cretaceous French amber

A

mt 2

ts pt 1 pt ti ti ti pp mt fe fe ts LC 3 ti pt mt tr tr RC fe pt cx AME mt tr ti fe cx mt pt tr ps cx ti ti fe tr cx pt fe fe pt 4 BL 1 tr ti co? fe sr pt sclerotized plate ALS 4 PLS op ALS anal tubercle? PLS

3 B cl ts T mt T T ti 4 pt pt ti PLS pt T anal tubercle? T mt T? ti ALS ts fe 2 PLS cl co? ALS mt fe fe ts tr sr cl EF tr tr mt pt ti ts cx fe cx op 1 tr cx cx ts BL fe ts pt tr mt cx T st tr ti mt mx LC pp fang ti fe ts ti RC mx fe 4 3 cl pt pt T mt pp ts fe ti fe pt 2 T mt pt ti

FIG. 2. — Interpretive drawings of Archaemecys arcantiensis n. gen., n. sp.: A, dorsal view; B, ventral view. See text for explanation of abbreviations. Scale bar: 0.5 mm.

GEODIVERSITAS • 2009 • 31 (1) 55 Saupe E. E. & Selden P. A.

Archaemecys arcantiensis n. sp. spines; each metatarsus with single trichobothrium, (Figs 1-3) most likely on dorsal surface (trichobothrium located seven-tenths of way along metatarsus from proximal Arachnida Salticidae – Néraudeau et al. 2002: fi g. 6.8. end in legs 2-4); tibiae 2-4 with at least one (up to TYPE LOCALITY. — Archingeay-Les Nouillers, Charente- three) trichobothria (Fig. 2). Legs relatively short Maritime, south-west France. and stout. Tarsi with unsclerotized constriction at the base of the tarsus (Fig. 1E). Th ree tarsal claws ETYMOLOGY. — Th e specifi c epithet is based on Arcan- tiatum, former Latin name of the locality Archingeay on unsclerotized onychium; paired claws with four, from which the fossil originates. possibly fi ve teeth, unpaired claw simple. Pedipalp rounded and bulbous, 0.37 long, 0.16 wide. DIAGNOSIS. — As for the genus. REMARKS DESCRIPTION While we are only able to view four spinnerets, we Body length 3.10. Carapace 1.31 long, 0.94 wide, cannot completely rule out the possibility of six, as ≥ 0.66 high; pars cephalica highly elevated so that there may be a very small median pair (this is unlikely carapace, when viewed from side, appears sub- since we are able to view inside the opisthosoma rectangular in outline; without tubercles. Only AME to the point of spinneret attachment). Regard- visible on anterior face of carapace. Chelicerae with less, even if the specimen possesses six spinnerets, ≥ 11 peg teeth on promargin (Fig. 3); peg teeth this would not change its placement within the ≈ 0.17 long, chelicerae 0.71 long; fang ≥ 0.35 long; Mecysmaucheniidae. Although the archaeids and chelicerae appear to originate from a foramen, al- pararchaeids have six spinnerets, these families do though this is not certain; sclerotized lip/projection not share the other characters found in our specimen above where chelicerae originate (Fig. 3). Sternum (see family remarks). Additionally, losing spinnerets 0.34 wide, 0.57 long, lateral margins project slightly is a fairly common phenomenon within the Araneae. between coxae. Maxillae converge slightly, almost Extant mecysmaucheniids have lost four spinnerets circular in shape. Petiole encircled by sclerotized in the reduction to the two the family currently plate (Fig. 2A); rugose epigastric plate, fl anked by possesses; given this, it seems quite possible that book lung covers (Fig. 2B). Opisthosoma 1.42 long, a primitive mecysmaucheniid would have four, or approximately 1.10 wide, height uncertain; likely even six, spinnerets. sub-globular in life; dorsal surface not preserved so presence of scutum not known. Opisthosomal cuticle wrinkled in subparallel lines, bearing short, DISCUSSION plumose setae, each originating from a prominent follicle. Four spinnerets and anal tubercle surrounded Th is is the fi rst description of a fossil mecysmaucheniid by well-developed chitinous ring (Fig. 2B); ALS and it extends the range of the family back to the well defi ned in two segments (Fig. 1D), posterior Cretaceous Late Albian). Archaeids have been de- spinnerets smaller; patch of cuticle with short setae scribed from Late Cretaceous amber (Penney 2003b) in front of ALS, possibly a colulus. Spiracle situ- and are known from the (Eskov1987; Selden ated well anterior to spinnerets; well fortifi ed with et al. 2008); it follows that mecysmaucheniids are cuticle ring (Fig. 1A-C). also present in the Mesozoic, since they are the Leg formula 1423; leg 1 cx 0.23, tr 0.16, fe 0.57, sister to the archaeids. pt 0.25, ti 0.39, mt 0.39, ts 0.39, total 2.38; leg 2 A cladistic analysis was attempted on the Palpima- cx 0.18, tr 0.19, fe 0.52, pt 0.20, ti 0.38, mt 0.33, noidea sensu Forster & Platnick (1984) in an eff ort ts 0.33, total 2.13; leg 3 cx 0.17, tr 0.18, fe 0.45, pt to determine where Archaemecys n. gen. grouped in 0.21, ti 0.39, mt 0.33, ts 0.28, total 2.01; leg 4 cx relation to the other palpimanoid families. Th e re- 0.25, tr 0.16, fe 0.54, pt 0.20, ti 0.44, mt 0.41, ts sult was a polytomy which failed to provide us with 0.33, total 2.33. Plumose setae on all leg segments; no useful information. Th e polytomy likely resulted

56 GEODIVERSITAS • 2009 • 31 (1) A new Mecysmaucheniidae (Arachnida, Araneae) from Cretaceous French amber

FIG. 3. — Archaemecys arcantiensis n. gen., n. sp., anterior view of the chelicerae and pedipalps. Notice the peg teeth on the cheli- cerae and the heightened profi le of the carapace. Scale bar: 0.1 mm. from a lack of informative characters used in the the use of combined characters to diagnose families. analysis and because we were dealing with family- In order to accurately perform an analysis on this level data that is possibly unnatural. For example, superfamily, one should include many possible the tendency to lose spinnerets is prevalent within outgroups, such as those within the Araneoidea, the Palpimanoidea: Palpimanidae, Stenochilidae, and re-evaluate the characters used to diagnose each Huttoniidae and Mecysmaucheniidae all have a family. Additionally, cladistic analyses should be reduced number (Forster & Platnick 1984), but performed at the species level. Th e analysis suggests the Archaeidae (sister to the Mecysmaucheniidae more work needs to be done on the systematics of and phylogenetically removed from many of the the Palpimanoidea. previously cited taxa) still retain the primitive six. Mecysmaucheniids are commonly found in the It is important to note that the mecysmaucheniids litter layer of forests (Forster & Platnick 1984; are diagnosed by the combined characters of having Jocqué & Dippenaar-Schoeman 2006). Since the only two spinnerets (a character present in other Archingeay amber fauna is considered a unique palpimanoid families) and chelicerae originating representative of a litter fauna (Néraudeau et al. from a foramen in the carapace. Th e mosaic of 2002; Perrichot et al. 2007), our spider specimen characters found in palpimanoid taxa is refl ected in corroborates this observation and supports the

GEODIVERSITAS • 2009 • 31 (1) 57 Saupe E. E. & Selden P. A.

hypothesis that resin fl owed directly onto the forest to Pangaea that has experienced extinction in the fl oor, thereby engulfi ng our spider and the many northern hemisphere. Th us, it appears as if the other organisms found in this amber. extant distribution of mecysmaucheniids consists Recent mecysmaucheniids occur only in New of relicts of a once more widespread distribution, Zealand and South America (specifi cally Argentina as is mimicked in their archaeid sisters. and Chile) (Jocqué & Dippenaar-Schoeman 2006; Platnick 2009), while archaeids are found only in Australia, South Africa and Madagascar. Th e pres- Acknowledgements ence of fossil archaeids and mecysmaucheniids in We thank Vincent Perrichot for providing the speci- Eurasia poses interesting biogeographical questions. men for study and help with preparation, Hannah Although the present distribution of mecysmau- Wood (University of California, Berkeley) and Dave cheniids and archaeids may be an artifact of sam- Penney (Th e Gambia) for useful discussions, and pling bias, this seems unlikely, and it is especially two anonymous reviewers for providing comments curious that mecysmaucheniids occur in precisely on the manuscript. Th is work is a contribution to those parts of Gondwana from which archaeids the project AMBRACE no. BLAN07-1-184190 to are absent. Th e particular distributions of these Didier Néraudeau from the Agence nationale de la two families may represent a case of competitive Recherche (ANR). exclusion or, more likely, since they are sister taxa, may result from a familial divergence in the distant NOTE ADDED TO PROOFS geological past. Th ere are two broad, opposing hypo- A work published recently by Wunderlich (2008) theses to explain the Gondwanan distribution of referred to Archaemecys arcantiensis n. gen., n. sp. extant taxa: mobilistic or vicariance biogeography, (this paper) as Palaeomecysmauchenius (this was a whereby Gondwana fragmented and continental manuscript name – we provided Wunderlich with drift explains current taxa distributions, and the a preliminary draft of the present paper in 2007) theory of a relict population or ousted relicts, which and placed our specimen in his new subfamily proposes a pancontinental distribution of taxa fol- Lacunaucheniinae. We refute this placement on lowed by extinction in the northern hemisphere (see the following grounds: Archaemecys n. gen. pos- Platnick & Nelson 1978; Nelson & Platnick 1981; sesses a ring around the spinnerets (contra mem- Eskov & Golovatch 1986; Briggs 1987, 1995; Eskov bers of Lacunaucheniinae) and does not, to our 1987, 1992; Penney 2003b; and Wunderlich 2004 knowledge, have three pairs of spinnerets (a trait for a review of the above scenarios). Mobilistic and/ of Lacunaucheniinae). Furthermore, Wunderlich or vicariance biogeography is not supported by the (2008) indicated we support the theory of ousted fossil evidence for archaeid and mecysmaucheniid relicts (the hypo thesis that northern lineages were spider families (see Penney 2003b and Eskov 1987 for ousted to the southern hemisphere by more com- further discussion on archaeids). Mecysmaucheniids petitive taxa). Th is is false: we simply stated that are known from the Cretaceous of France (this Archaeidae and Mecysmaucheniidae likely had a paper), whereas archaeids are known from the more widespread distribution in the Mesozoic, so Jurassic of Kazakhstan (Eskov 1987) and China their extant range would therefore be a relict of a (Selden et al. 2008), French Eocene amber (Penney previous pancontinental distribution. 2007), and they are commonly found in Cenozoic Baltic amber. Th is non-Gondwanan distribution of the fossils suggests either an overall migration REFERENCES from the northern to the southern hemisphere from Mesozoic to Recent times (this is unlikely) BRIGGS J. C. 1987. — Antitropical distribution and evolution in the Indo-West Pacifi c ocean. Systematic or, more likely, because few fossil spider localities Zoology 36: 237-247. are yet known from Gondwanaland (Wunderlich BRIGGS J. C. 1995. — Global Biogeography: Developments 2004), a cosmopolitan distribution harking back in Palaeontology and Stratigraphy, Vol. 14. Elsevier,

58 GEODIVERSITAS • 2009 • 31 (1) A new Mecysmaucheniidae (Arachnida, Araneae) from Cretaceous French amber

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Submitted on 12 October 2007; accepted on 18 December 2008.

60 GEODIVERSITAS • 2009 • 31 (1)