Table Of Content2002. The Journal of Arachnology 30:487-493
THE OLDEST LINYPHIID SPIDER, IN LOWER CRETACEOUS
LEBANESE AMBER (ARANEAE, LINYPHIIDAE, LINYPHIINAE)
David Penney and Paul A. Selden: Earth Sciences, University of Manchester,
Manchester, M13 9PL, UK, E-mail: [email protected],uk
ABSTRACT. A new fossil Linyphiidae: Linyphiinae is described from 125-135 Ma old (Upper Neo-
comian-basal Lower Aptian) Cretaceous amber from the Kdeirji/Hammana outcrop, Lebanon. This is the
oldest known linyphiid as well as the oldest described amber spider. The first major radiation of the
linyphiid subfamilies occurred in the early Cretaceous, if not before, and the presence of Linyphiidae in
this period predicts the presence of Pimoidae then too. Current evidence, which suggests the higher
araneoids did not radiate and diversify until after the end=Cretaceous mass extinction event may be an
artefact of sample size.
Keywords: Linyphiidae, Linyphiinae, Lebanese amber fossil
The spider family Linyphiidae contains taxonomic comments of Wunderlich 1986)
4,129 extant species in 550 genera (Platnick and from Upper Cretaceous (Turonian, 90-94
2001), which represents approximately nine Ma) amber from New Jersey (Penney in
percent of total spider species diversity. It press). Wunderlich (1998) described a liny-
ranks second, after Salticidae, in terms of phiid from what was thought to be Dominican
number of described species, but first for Republic amber, but has since been shown to
number of genera (Platnick 2001). However, be Madagascan copal (J. Wunderlich, pers.
many of these genera are monotypic and comm.), which is semi-fossilized resin less
A
would probably not withstand phylogenetic than two million years old. non-amber fos-
scrutiny (Hormiga 2000). The family consists sil linyphiid was described by Berland (1939)
mainly of very small spiders, mainly sheet- from the Oligocene of Alsace, France. The
web builders. It has a global distribution, but specimen in Mexican amber is an exuvium
linyphiids are most diverse in northern tem- from an immature spider, and the French spec-
&
perate regions (Coddington Levi 1991). imen is poorly preserved. Both were described
Their niche may be occupied by Theridiidae and named as linyphiids, but we consider their
in the lower and southern latitudes (e.g. Nen- current placement in the Linyphiidae to be un-
twig 1993: table 8). There are six recognized reliable. Fossil linyphiids have been reported
extant linyphiid subfamilies: Dubiaraneinae as present, but not described, from Eocene
&
Millidge 1993, Erigoninae Emerton 1882, Bitterfeld amber (44 Ma) (Schumann
Linyphiinae Blackwall 1859, Micronetieae Wendt 1989) and Upper Cretaceous Canadian
&
Hull 1920, Mynogleninae Lehtinen 1967, and amber (65-83 Ma) (McAlpiee Martin
Stemonyphantinae Wunderlich 1986 (A. Tan- 1969).
asevitch pers. comm.), although the Linyphi- This paper describes the oldest known lin-
inae are considered by some (e.g. Hormiga yphiid spider from upper Neocomian-basal
2000) to consist of two tribes: Micronetini and Lower Aptian (c. 125—135 Ma) Cretaceous
Linyphiini, reducing the number to five (e.g. Lebanese amber from the Kdeirji/Hammana
Hormiga 1994a, 2000). outcrop, which represents one of the oldest
Fossil linyphiids have been described from insect inclusion-bearing amber deposits (Azar
Tertiary ambers from the Dominican Republic 1998). This specimen is the oldest described
(Miocene, 15-20 Ma) (Wunderlich 1988), amber spider, the previous being a new genus
Mexico (Miocene-Oligocene, 19-27 Ma) (Pe- and species of Nemesiidae described in Bar-
UK
trunkevitch 1971), the Baltic region (Eocene, remian amber from the Isle of Wight,
44 Ma) (Petrunkevitch 1942, 1958; see also (Selden in press).
487
—
488 THE JOURNAL OF ARACHNOLOGY
—
Figures 1-2. Female linyphiine in Lebanese amber, photomicrographs of holotype. 1. Dorsal view
x50; 2. Ventral view x50.
METHODS
leg formula (e.g. 4123), the legs are ranked in
—
Preservation. The spider is preserved in order of length (longest first). Tml and Tm4
a very small piece (3x3x1 mm) of clear, are measurements of the distance that a tri-
yelloW“Colored amber. There are a few air chobothrium is located along the lengths of
bubbles of varying size and a small number metatarsi 1 and 4 respectively, relative to the
Tml —
of organic and inorganic syninclusions. length of the leg segment, e.g. 0.3
—
Methods. The amber piece had been pre= indicates that the trichobothrium is located
pared by being set in a clear synthetic resin three tenths of the way along metatarsus 1,
disc (22 mm diameter x 2 mm thick), and pol- from the proximal end of the segment.
ished prior to receipt by the authors. The spec- SYSTEMATIC PALAEONTOLOGY
imen was studied, drawn and photographed,
using both transmitted and incident light, us- Family Linyphiidae Blackwall 1859
ing a Nikon Optiphot stereomicroscope, with
Subfamily Linyphiinae Blackwall 1859
a camera lucida drawing tube and a Nikon
gen. et sp. indet.
FX-35DX camera attached by means of a
Figs. 1-3
phototube. An Olympus SZH stereozoom mi-
croscope with incident light revealed addition- Distribution.—Upper Neocomian-basal
al detail of the specimen. All measurements Lower Aptian (c. 125-135 Ma) Cretaceous
are in mm. Lebanese amber from the Kdeirji/Hammana
Abbreviations used in the text and fig- outcrop, Lebanon. —
ures. ab = air bubble; at = anal tubercle; ch Only known specimen, Female, speci-
= chelicera; co = colulus; cx = coxa; ep = men No. 491 preserved in Cretaceous Leba-
epigyne; fe = femur; la = labium; mx = max- nese amber, held in the Laboratoire
illa; mt — metatarsus; op = opisthosoma; pa d’Entomologie, Museum National d'histoire
= patella; pp = pedipalp; sp = spinneret; st Naturelle, Paris (MHNP), examined.
= sternum; ta = tarsus; ti = tibia; tr = tri- I^escTiptmn.-—Measurements: body length
chobothria; 1-4 = walking legs 1-4. In the 1.86; carapace ground away (Fig. 1), but the
PENNEY & SELDEN—LINYPHIIDAE IN LEBANESE AMBER 489
—
Figure 3. Female linyphiine in Lebanese amber, camera lucida, ventral view. Scale bar = 1 mm.
chelicerae are in place so it is possible to de- long with distinct serrula and a transverse row
termine the body length. Detailed structure of dark, chitinized denticles; labium slightly
and dentition of chelicera are not visible, but rebordered, much wider than long. Sternum
it appears unmodified and lacks stridulatory 0.43 long, 0.40 wide, smooth, with sparse
striae on the ectal surface. Maxilla wider than covering of setae; margin slightly incised to
490 THE JOURNAL OF ARACHNOLOGY
accommodate coxae, sharply truncated poste- genus. This female linyphiine, it is the oldest
riorly where it extends between fourth coxae; representative of the Linyphiidae recorded,
five erect setae along the truncated edge. Op- and is also the oldest described amber spider.
isthosoma 1.23 long, 1.00 high, sub-spherical,
DISCUSSION
without tubercles or scuta, covered with short
setae; the right side has collapsed inwards. The superfamily Araneoidea comprises
Anterior lateral and posterior lateral spinnerets twelve extant families (Griswold et al. 1998)
with numerous spigots, posterior median spin- and the extinct families Ephalmatoridae Pe-
nerets not visible. Colulus relatively large, trunkevitch 1950, redefined by Wunderlich
with at least three bristles. Anal tubercle dis- (1986) from Baltic amber, and Juraraneidae
tinct (Figs. 2, 3). Epigyne projects ventrally, Eskov 1984 from the Jurassic of Kazakhstan.
appears domed in lateral view (a clear ventral The phylogenetic scheme for the Araneoidea
view is not possible because of the position (Fig. 4) follows Griswold et al. (1998), except
of the spider in the amber matrix so the de- for the placement of the fossil taxa. According
tailed structure is not clear). Epigyne heavily to Griswold et al. (1998) the unambiguous
sclerotized, with a single opening and a flat synapomorphies for the linyphioid families,
dorsal margin; lateral margins appear rounded which includes Linyphiidae and its sister tax-
and project slightly posteriorly. on Pimoidae (see Hormiga 1994b), are strid-
Leg formula 1423. Leg 1 fe 0.71, pa 0.19, ulating striae ectally on the male chelicerae,
ti 0.43, mt 0.41, ta 0.33, total 2.07; leg 2 fe patella-tibia autospasy, and an enlarged base
0.57, pa 0.16, ti 0.29, mt 0.26, ta 0.24, total on the basal posterior lateral spinneret cylin-
1.52; leg 3 fe 0.41, pa 0.14, ti 0.21, mt 0.19, drical gland spigot. The fossil specimen is fe-
ta 0.17, total 1.12; leg 4 fe 0.63, pa 0.16, ti male, all legs are intact, and the fine detail of
0.33, mt 0.31, ta 0.21, total 1.64; all segments the spigots is not clear. The fossil is excluded
with distinct setae and annulate distally. Cox- from the Pimoidae on account of its size, con-
ae and trochantera without modifications; fe 1 siderably smaller than 5 mm, the lower end of
and possibly fe 2 with short median dorsal the range given by Hormiga (1994b), the epi-
spine, apparently lacking on fe 3 and 4; fe 1 gyne does not protrude posteriorly beyond the
with long prolateral spine located just distal epigastric furrow as it appears to in many pi-
to midpoint; all patellae with proximal and moid species, femur 4 lacks dorsal spines, and
distal dorsal spines; tibial spination 2, 2, 2, 2; the legs lack long setae which are curved at
ti 1 also with median, long prolateral spine; their distal end (e.g. Hormiga 1994b). The
metatarsi and tarsi without spines (Eigs. 1-3). systematic placement of many genera within
All tibiae with trichobothria equal to or longer linyphiid subfamilies is based solely on auta-
than the tibial diameter (Fig. 3); Tml = 0.3, pomorphies derived from male secondary
Tm4 lacking. Tarsi with three simple, untooth- genital organs and in some cases no unambig-
ed claws and accessory setae, unpaired claw uous diagnostic character states have been es-
long, pedipalp with a single simple claw. tablished for female specimens. Therefore, we
—
Remarks. It is well appreciated that am- tentatively place this specimen in the Liny-
ber spiders are taxonomically subequal to Re- phiidae: Linyphiinae, based on the remaining
cent spiders (e.g. Eskov 1990). In many fos- somatic and genitalic morphology, for exam-
sils it is difficult to identify and study those ple the legs spination and trichobothrial pat-
characters considered important as diagnostic terns, but to which tribe (Micronetini or Lin-
for extant taxa. The specimen described here yphiini) it belongs is uncertain. We accept that
cannot be diagnosed by any putative autapo- this placement is not based on any putative
morphies, so a specific epithet is not assigned, autapomorphies and are unaware of any stud-
nor can it be placed with certainty in an extant ies, which provide reliable diagnostic or phy-
->
—
Figure 4. Evolutionary tree of the Araneoidea. Dotted fill = described fossil, striped fill = known
geological range, vertical solid lines = range extension, horizontal solid lines = phylogenetic relationships,
tight dashed lines = ancestral lineage, loose dashed lines = ghost lineage, Lagerstatten on right side.
Terminology follows Smith (1994).
PENNEY & SELDEN—LINYPHIIDAE IN LEBANESE AMBER 491
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492 THE JOURNAL OF ARACHNOLOGY
logenetic chaetotaxy or trichobothrial patterns. Cretaceous spiders, many of which are poorly
However, paleoarachnologists are often con- preserved and not identifiable to family, ac-
fronted with fossils which are unique and do count for the only three publications to date
&
not possess, or clearly exhibit, the characters (Eskov Wunderlich 1994; Penney in press;
used in Recent spider taxonomy and system- Selden in press) that describe Cretaceous am-
atics, but important specimens of great antiq- ber spiders. The relatively large number of
uity such as this warrant description and Recent spider families being discovered in
placement as far as is possible. rocks and amber of Mesozoic age suggests a
The presence of fossil Linyphiidae in the great antiquity of modern spider families (Sel-
upper Neocomian-basal Lower Aptian (c, den & Penney 2001), we would expect to find
125-135 Ma) predicts the presence of Pimo- some of these families given enough Creta-
idae, otherwise unknown in the fossil record, ceous specimens and are currently working
in the Cretaceous. It also predicts the occur- through this material.
rence of the spineless femur clade and the
ACKNOWLEDGMENTS
symphytognathoids, or their ancestors approx-
imately a further 35 Ma back in the fossil re- We thank Dany Azar (MHNP) for provid-
cord, from the oldest known described fossil ing the specimen, Mark Harvey (Western
Linyphiidae in Turonian New Jersey amber Australian Museum), Gustavo Hormiga
A
(Penney in press) (Fig. 4). number of au- (George Washington University), Nikola]
thors e.g. Wunderlich (1986), Millidge (1993) Scharff (Zoologisk Museum, Copenhagen) for
and Hormiga (1994a, 2000) have provided their comments on the manuscript, and Lev-
—
various hypotheses regarding the subfamilial erhulme Trust grant F/00 1201 Mesozoic
phylogenetic relationships within the Linyphi- Arachnids.
idae. These were compared and contrasted by
LITERATURE CITED
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