Table Of Content2003. The Journal of Arachnology 31:122-130
AFRARCHAEA GRIMALDII, A NEW SPECIES OF ARCHAEIDAE
(ARANEAE) IN CRETACEOUS BURMESE AMBER
David Penney: Earth Sciences, The University of Manchester, Manchester, M13 9PL,
United Kingdom
ABSTRACT, Afrarchaea grimaldii new species (Archaeidae, Archaeinae) from 88-95 Ma (Cenoman-
ian-Turonian) Upper Cretaceous amber (Burmite) from Myanmar (Burma) is described. This is the first
spider to be described from this deposit and is the oldest known Archaeidae sensu stricto extending the
known range of the family by approximately 50 Ma from the previously oldest recorded specimens in
Baltic and Bitterfeld ambers, and provides further evidence that spiders were not severely affected by the
end-Cretaceous mass extinction event. It represents the oldest fossil record ofan araneophagic spider. This
species could be used to argue forboth the theory ofmobilistic biogeography and ousted relicts to explain
the zoogeography of the genus, but until new data become available, supports neither reliably.
Keywords: Myanmar, fossil spiders
Biological inclusions have been known which are distinguished from other spiders by
from Burmese amber or Burmite for almost a the combination ofpromarginal cheliceral peg
century (Cockerell 1916), but hitherto no spi- teeth and an abdomen-petiole stridulatory
&
ders have been described from this source system (Forster Platnick 1984). In addition,
(Ross & York 2000). Some ofthe spider fam- the three Recent genera: Afrarchaea Forster &
&
ilies present in the Burmese amber collections Platnick 1984, Austrarchaea Forster Plat-
in the Department of Palaeontology of the nick 1984 ^ndArchaea Koch & Berendt 1854,
Natural History Museum, London, were listed have their carapace with the pars cephalica el-
in Penney (2000) and Rasnitsyn & Ross evated above the pars thoracica, often con-
(2000); (the specimens listed under the fami- stricted between the head (which bears long,
lies Eusparassidae and Myrmeciidae by the slender chelicerae with a short fang) and tho-
&
latter authors (which are probably misidenti- rax to form a distinct neck (Forster Platnick
fications, pers. obs.) are no longer valid ar- 1984). Here, the first Cretaceous Archaeidae
achnological taxa and should read Sparassidae sensu stricto is described, from Burmese am-
and Corinnidae respectively [e.g. Platnick ber, and the systematics and biogeography of
A
2002]). Grimaldi et al. (2002) listed eleven the family are briefly discussed. checklist
families provisionally recorded from Burmese of fossil Archaeidae sensu lato is provided.
amber, including the specimen described here.
METHODS
&
Zherikhin Ross (2000) proposed a Late
—
Cretaceous age for Burmite, and based on the Preservation. Both specimens are pre-
shared insect taxa of this amber with other served in Burmese amber or Burmite (for de-
well-dated amber deposits it probably dates tails oflocality and stratigraphy, see Zherikhin
from the Cenomanian or Turonian (Grimaldi & Ross 2000; Grimaldi et al. 2002) and be-
et al. 2002). Cretaceous amber spiders have long to the Department of Entomology at the
previously been described from the Santonian American Museum of Natural History
of Siberia (Eskov & Wunderlich 1994), the (AMNH). The holotype, AMNH Bu-256 is
(3X4X5
Turonian of New Jersey (Penney 2002), the preserved in a small piece mm)
Barremian ofthe Isle ofWight (Selden, 2002) of clear yellow amber suffused with darker
and the Upper Neocomian-basal Lower Ap- bands, which represent layering ofthe resin at
tian of Lebanon (Penney & Selden 2002). the time ofexudation from the tree. This con-
The Archaeidae are small to medium-sized clusion is supported because only one region,
haplogyne, ecribellate araneomorph spiders between two of these darker bands, contains
122
— ———— — —
PENNEY—AF^^C//AEA IN CRETACEOUS BURMESE AMBER 123
air bubbles. The spider is preserved in a layer lin; NCA = National Collection ofArachnida,
without air bubbles; there are no syninclu- Plant Protection Research Institute, Pretoria;
sions. There is some fracture damage as a re- PIN = Palaeontological Institute ofthe USSR
sult of specimen preparation however, overall Academy of Sciences, Moscow; SGPIH =
this is an exq—uisitely preserved specimen. Geologisch-Palaontologisches Institut und
Methods* Prior to being received by the Museum, Hamburg.
author the amber had been set in a clear plas- SYSTEMATIC PALEONTOLOGY
tic resin and cut and polished to reveal the Family Archaeidae Koch & Berendt 1854
inclusioAn.MFNurHther preparation was carried out Subfamily Archaeinae Koch & Berendt 1854
at the as specified by the author to &
Afrarchaea Forster Platnick 1984
reveal further important taxonomic features.
All measurements were made using an ocular Type species, Archaea godfreyi Hewitt
graticule and are in mm. Drawings were done 1919 by original—designation.
under incident light with camera lucidas at- Distribution. Recent species in South Af-
tached to an Olympus SZH stereomicroscope rica and Madagascar, fossil species in Bur-
and a Nikon Optiphot stereo compound mi- mese amber, Myanmar (Burma).
croscope, and photographs were taken with a Remarks. Afrarchaea was erected as a
Nikon DIX digital camera attached to the Ni- monotypic genus by Forster& Platnick (1984)
kon microscope, using a 2.5X photoeyepiece for Archaea godfreyi from South Africa and
and a 2X objective lees then manipulated in Madagascar. It was distinguished from the
Adobe Photoshop. other genera by having a less constricted car-
Recent material examined. Afrarchaea apace “neck” and on the basis of the female
ngomensis Lotz 1996; 16,1$ from Ngome genitalia. Eskov (1992) considered Afrar-
State Forest, KwaZulu/Natal Province; NCA chaea ajunior synonym ofArchaea'^ however,
93/612 (coll. M. van der Merwe, Jan. 1993). this was not based on the examination of Re-
Abbreviations used in the text and fig- cent specimens and has not been accepted by
ures.—In the leg formula (e.g. 1423), the legs subsequent workers (Platnick 2002), Lotz
are ranked in order of length (longest first), (1996) described five new Afrarchaea species
Tm is the ratio of the distance that a tricho- from South Africa and provided new data for
bothrium is located from the base ofthe meta- A. godfreyi.
Tm =
tarsus (e.g. 0.8 indicates that the tri- Afrarchaea grimaldii new species
chobothrium is located eight-tenths ofthe way Figs. 1-5
along the metatarsus, from the proximal end
of the segment). Abbreviations used in the text Archaeidae: Grimaldi et al. 2002: 28, fig. 18c.
and figures are as follows: ALE = anterior lat- Material examined. -Holotype: AMNH
eral eye(s); AME = anterior median eye(s); b Bu-256, adult male, Burmese amber, Kachin:
= bulb; bs = blunt setae; car = carapace; cf Tanai Village (on Ledo Road 105 km NW of
“ clypeal foramen; cs = cheliceral seta; e = Myitkyna); coll. Leeward Capitol Corporation
embolus; ebl = extension ofbulb lip; f = fur- 2000, Non-types: AMNH Bu-706, degraded
row; F = flaw in amber; fe = femur; fg = specimen, same horizon and locality.
fang; LC ~ left chelicera; lot = lateral ocular Diagnosis.- Afrarchaea grimaldii can be
tubercle; mt = metatarsus; mx = maxilla; op distinguished from all other species by having
= opisthosoma; pa = patella; PEE = posterior a bent tegular apophysis and a spoon-shaped
lateral eye(s); PME = posterior mediae eye(s); embolus.
Pp, = pedipalp; pt = peg teeth; RC — right Etymology. The specific epithet is a pa-
chelicera; sp = spinneret region; T = tricho- tronym in honor of Dr. David Grimaldi
bothrium; ta = tarsus; TA = tegular apophy- (AMNH) for his contributions to the study of
sis; ti = tibia; 1-4 = walking le—gs 1-4. amber and for loaning and assisting in the
Repository abbreviations. AMNH = preparation of this material.
American Museum of Natural History; AP = Description of bolotype. Body length
Amber Museum ofPalanga, Lithuania; MCZ = 1.97; carapace 0.86 long, 0.43 wide, 0.64 high
Museum of Comparative Zoology, Harvard; in region of pars cephalica; region between
MfN = Museum fiir Naturkunde Institut fur caput and thorax strongly developed and little
Palaontologie, Humboldt-Universitat zu Ber- differentiated from head; indent at junction
124 THE JOURNAL OF ARACHNOLOGY
—
Figures 1-3. Afrarchaea ghmaldii new species. Holotype, AMNH Bu-256, Burmese Amber. 1, lateral
mm
view of whole specimen. 2, anterior view of chelicerae. Scale line 1.0 for both figures. 3, ventral
view of pedipalp. Scale line 0.1 mm.
with pars thoracica 0.29 high, 0.46 long; with liceral foramen, thickened in proximal half
numerous distinct tubercles, each bearing a when viewed laterally and tapering slightly at
single seta lying flat against the carapace their tips along fang furrow, lacking triangular
(Figs. 1, 5); narrow furrow running down mid- projections on apical promargin; each has a
line from cheliceral foramen, visible when single erect dorsal seta close to proximal con-
viewed anteriorly through the chelicerae (Fig. striction; lacking cheliceral dentition, but nu-
2). ALE and PLE of equal size and contigu- merous peg-teeth along promargin and long
ous, on a tubercle (Eig. 2), AME larger, PME strong hairs along promargin in fang region
not visible but presumably smallest as in Re- (Figs. 2, 4); lateral stridulatory ridges not vis-
cent species (e.g. Lotz 1996). Clypeus slightly ible but presumed to be present; fang short
greater than diameter of AME. Chelicerae and curved backwards. Sternum 0.50 long,
0.79 long, slightly divergent and project out 0.16 wide, possibly tuberculate, lateral mar-
from the body at approximately 45 degrees gins appear to project slightly between coxae.
when viewed laterally (Figs. 1, 5), strongly Labium longer than broad, maxillae consid-
constricted basally where they insert into che- erably longer than broad, slightly convergent.
PENNEY—AF/?A/?C//AEA IN CRETACEOUS BURMESE AMBER 125
—
Eigures 4-5. Afrarchaea grimaldii new species. Holotype, AMNH Bu-256, Burmese Amber. 4, an-
mm
terior view of chelicerae. 5, lateral view of whole specimen. Scale line 1.0 for both figures.
projecting from body at similar angle to che- the diagnostic characters of the genus given
iicerae (Fig. 1). Opisthosoma 1.11 long, by Forster & Platnick (1984). It can be ex-
height and width uncertain, wrinkled surface cluded from the remaining archaeid genera as
(Figs. 1, 5), presumably a taphonomic artifact follows: Archaea C.L. Koch & Berendt 1854,
resulting from dehydration process associated because it lacks the distinctive slender neck
with amberpreservation, presumably subglob- between the head region and the pars cephal-
ular in life; no dorsal scutum. Opisthosoma ica; Baltarchaea Eskov 1992, because it lacks
covered with small patches ofchitinous tissue the cephalic posterior angular projections, the
each ofwhich bears short, fat, blunt seta; spin- abdomen does not extend beyond the spinner-
nerets and anal tubercle not clear and sur- ets and the legs and chelicerae are not com-
rounded by chitinous ring (Figs. 1, 5). paratively short; Mimetarchaea Eskov 1992,
Leg formula 1423; leg 1 fe 1.14, pa 0.29, because it lacks the mimetid-like metatarsal
ti 1.00, mt 0.49, ta 0.34, total 3.26; leg 2 fe spines on legs 1 and 2; Austrarchaea Forster
0.86, pa 0.19, ti 0.74, mt 0.36, ta 0.29, total & Platnick 1984, because the neck in the fos-
2.44; leg 3 fe 0.57, pa 0.19, ti 0.40, mt 0.29, sil specimen is too short, as is the embolus of
ta 0.24, total 1.69; leg 4 fe 0.93, pa 0.19, ti the male palp; Jurarchaea Eskov 1987, be-
0.71, mt 0.40, ta 0.31, total 2.64. All leg seg- cause although Eskov (1987) was somewhat
ments without spines, but with pubescence of ambiguous with his diagnosis, in that he did
fine setae; each metatarsus with single tricho- not provide any distinct autapomorphies for
bothrium (Tm 1-4 = 0.8-0.9); tibiae 1-3 with his new taxa, but provided a list of general
at least one dorsal trichobothrium, none visi- morphological descriptions that he later em-
ble on ti 4 (Fig. 1). Some leg segments show phasized may be somewhat speculative, he
evidence of annulations and darker markings, placed this genus closer to the families Par-
particularly in distal region; three tarsal claws archaeidae and Holarchaeidae than Archaei-
on onychium; paired claws toothed, unpaired dae sensu stricto. There are no spine-like
claw simple. Pedipalp has relatively long fe- horns sensu Lotz (1996) visible on pars ce-
mur, large rounded bulb, bent tegular apoph- phalica of the fossil, but these are small in
ysis and sp—oon-shaped embolus (Fig. 3). Recent specimens and may be present in the
Female. Unknown. — fossil but obscured by the legs or flaws in the
Distribution and age. Burmese amber, amber. Specimen Bu-706 is preserved in a
Myanmar (Burma); probably Upper Creta- clear piece of amber with a spider syninclu-
&
ceous (see Zherikhin Ross 2000): Ceno- sion (possibly Oonopidae). It is severely de-
manian-Turo—nian (see Grimaldi et al. 2002) graded, barely visible, and it is only with a
Remarks. This specimen conforms with reasonable degree of imagination that the
126 THE JOURNAL OF ARACHNOLOGY
raised pars cephalica and elongated chelicerae (1854) and Menge (1854) were considered
can be seen. For this reason it is very tenta- lost for many years (e.g. Forster & Platnick
tively assigned to this species. Ecological ob- 1984), however, many of Koch & Berendt's
servations of Recent archaeids are sparse, but (1854) types are kept in the Institut fiir Pa-
all evidence suggests that they are araneo- laontologie. Museum fiir Naturkunde, Zen-
phagic, free-moving, cryptozoic hunters (For- tralinstitut der Humboldt-Universitat zu Berlin
ster & Platnick 1984). Most Afrarchaea in but those of Menge are still considered lost
collections have been caught using pitfall (Table 1).
traps or through sifting leaf litter (Lotz 1996). The taxonomic composition and systematic
There is no evidence to suggest that the close- placement of the Archaeidae sensu lato con-
ly related families Holarchaeidae, Pararchaei- tinues to stimulate lively debate. Since its
dae and Mecysmaucheniidae are also araneo- original description, ten Recent (two with fos-
&
phagous (Forster Platnick 1984). Using the sil representatives) and four strictly fossil gen-
premise of behavioral fixity, which states that era have at one time or another, been placed
fossil organisms can be expected to have be- within the Archaeidae; these are now distrib-
haved in a similar manner to their Recent rel- uted among the six families: Archaeidae sensu
atives at genus and often at family level, the stricto, Holarchaeidae, Mecysmaucheniidae,
specimen described above represents the old- Pararchaeidae, Tetragnathidae and Salticidae
est known occurrence of araneophagy in the (e.g. Eskov 1992; Platnick 2002). As currently
spider fossil record (Jurarchaea belongs ei- delimited, Archaeidae sensu stricto contains
ther in the family Pararchaeidae or Holar- 18 Recent species in three genera (Platnick
chaeidae [see Eskov 1987]). This is the first 2002; Harvey 2002) and ten fossil species
occuiTence ofAfrarchaea in the fossil record, (Archaeidae sensu lato) in five genera (Table
taking the genus back 88-95 Ma, and is also 1). Holl (1829) described the new genus and
the oldest record of the Archaeidae sensu species Entomocephalus formicoides from
stricto, extending the known range of this Baltic amber. This was listed as belonging in
family by approximately 50 Ma from the pre- the Archaeidae by Petrunkevitch (1958) and
vious oldest records in Baltic and Bitterfeld to my knowledge this is the only mention of
ambers. These fossils extend the known range this taxon in the literature since its descrip-
of yet another Recent spider family through tion. HolFs figure of this specimen (plate 8:
and beyond the end Cretaceous mass extinc- fig. 68a) is almost certainly a salticid probably
tion event, suggesting that this catastrophe had belonging to the genus Myrmarachne Mac-
little effect on the araneofauna, and provides Leay 1839, even though the figure and de-
further evidence for the great longevity for scription have the specimen with only six
many Recent spider families (Selden & Pen- eyes. If indeed this is the case, then under the
ney 2001) and genera (e.g. Penney 2002). ICZN law of priority, Entomocephalus Holl
1829, precedes Myrmarachne MacLeay 1839.
DISCUSSION
However, the location of the specimen on
The spider family Archaeidae is unique in which the description was based is unknown,
that it was first described from three fossil and the description of the genus consisted of
species in Baltic amber (Koch & Berendt only one sentence. The name Myrmarachne is
1854) in a paper published posthumously by well established, in common usage and should
Menge (1854) who added three more new spe- probably be maintained unless HolTs fossil
cies. The first Recent species was discovered specimen can be located. Eoarchaea Forster
in Madagascar a quarter of a century later (O. & Platnick 1984 was erected based on a single
Pickard-Cambridge 1881) and subsequently immature amber spider (Forster & Platnick
they have been found in Africa and other re- 1984). No mature specimens of this species
gions ofthe southern hemisphere (e.g. Harvey are known and the fossils attributed to it prob-
2002). It is also, to my knowledge, the only ably belong to various other Archaea species
family to have received paleontological treat- (Eskov 1992).
ment by that most eminent of arachnologists, Archaeidae was divided into four families:
Eugene Simon, who described a new species Archaeidae sensu stricto, Mecysmaucheni-
preserved in Baltic amber (Simon 1884). The idae, Holarchaeidae and Pararchaeidae by For-
specimens described by Koch & Berendt ster & Platnick (1984) in a review of the su-
'
PENNEY— IN CRETACEOUS BURMESE AMBER
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128 THE JOURNAL OF ARACHNOLOGY
perfamily Palpimanoidea. They also placed a of east and west Gondwanaland was initiated
number of disparate families (Mimetidae, Mi- by seafloor spreading between Africa and
cropholcommatidae, Textricellidae) alongside Madagascar in the Somali basin during the Ju-
the archaeoids, increasing the size of the Pal- rassic quiet interval (c. 165 Ma), with the
pimanoidea considerably, which had previous- landmass of Madagascar + India eventually
ly consisted of only three families: Palpiman- separating from Africa during the late Jurassic
idae, Stenochilidae and Huttoniidae. However, (152 Ma) (McLoughlin 2001). Madagascar
few subsequent authors agreed with these au- separated from the Seychelles-India block
thors’ concept of the Palpimanoidea (see dis- 95-84 Ma and India migrated rapidly north
&
cussions in Eskov 1987, 1992; Coddington reaching equatorial latitudes by the Eocene
Levi 1991), the monophyly of which was and combining with southern Asia (including
questioned. The superfamily Palpimanoidea West Burma) only about 43 Ma (McLoughlin
was cut back to its original size by Schiitt 2001). West Burma had separated from north-
(2000), based on a reanalysis of the autapo- eastern Gondwana in the late Triassic-late Ju-
&
morphies proposed by Forster Platnick rassic during the formation of the Neotethys
(1984). However, the correct systematic place- Ocean and was accreted to southeast Asia by
ment of the archaeids remains uncertain the late Cretaceous (McLoughlin 2001).
(Schiitt 2000). Therefore, the occurrence of this genus in
Fossils are often considered to be less use- Burmese amber could be used to support the
ful than Recent specimens for systematic stud- theory of mobilistic biogeography for its Re-
ies because of their imperfect preservation. cent distribution only if it existed throughout
However, they are of paramount importance Gondwanaland during the late Triassic-late
in studies of historical biogeography, and can Jurassic. Three spiders have been described
&
play a decisive part in the falsification of pro- from the Triassic (Selden Gall 1992; Selden
posed hypotheses (e.g. Eskov 1990). For ex- et al. 1999) but none were placed in Recent
ample, the current Gondwanan distribution of genera. The currently more plausible expla-
the Recent species of the spider family Ar- nation for the presence of Afrarchaea in this
chaeidae supports the theory ofmobilistic bio- locality is that it is an ousted relict from a
geography i.e. that the fragmentation ofGond- formerly pancontinental distribution in north-
wanaland and the subsequent continental drift ern paleolattitudes as is the case forArchaea,
can explain their current distribution. How- another Recent archaeid genus, and presum-
ever, because fossils of this family occur in ably also the sister taxon ofAfrarchaea.
Baltic amber (Koch & Berendt 1854) and ACKNOWLEDGMENTS
from the Jurassic of Kazakhstan (Eskov
1987), the paleontological data contradict this I thank D. Grimaldi of the American Mu-
hypothesis and a different explanation is re- seum of Natural History, New York for pre-
quired (the specimen reported from Domini- paring and providing the Burmese amber
can amber by Wunderlich [1999] is actually specimens for research purposes, J. Wunder-
preserved in Madagascan copal [Wunderlich, lich, Germany, forproviding rare manuscripts,
NCA
pers. comm. 2000]). The theory ofousted rel- A. Dippenaar-Schoeman, South Africa,
icts (e.g. Eskov & Golovatch 1986) proposes for providing comparative Recent material,
that austral disjunctions result from a formerly and R Selden, University of Manchester, for
pancontinental distribution followed by the his comments on the manuscript. This work
extinction of ‘intermediate links’ from the was compl—eted under Leverhulme Trust grant
northern continents. There is a considerable F/00 1201 Mesozoic Arachnids.
amount ofpaleontological data, in the form of LITERATURE CITED
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