An orb-weaver spider (Araneae, Araneidae) from the early Eocene of India
Raman Patel,
1
Rajendra Singh Rana,
1
and Paul A. Selden
2,3
1
Department of Geology, Hemvati Nandan Bahuguna Garhwal University, Srinagar, 246174 Uttarakhand, India 〈ramanpatel2142@gmail.com〉,
〈rajendra.rana1@gmail.com〉
2
Paleontological Institute and Department of Geology, University of Kansas, 1475 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
〈selden@ku.edu〉
3
Natural History Museum, Cromwell Road, London SW7 5BD, UK
Abstract.—A new fossil spider is described from the early Eocene (Ypresian) Palana Formation (54 to 57 Ma) at the
Gurha opencast lignite mine, near Bikaner, western Rajasthan, India. It is the first report of a nonamber fossil spider
from India. The fossil is referred to the modern genus Nephila Leach, 1815, but with hesitation because, while its
habitus is similar to that genus, it lacks the behavioral synapomorphies that distinguish the genus.
Introduction
The golden orb-weaver genus Nephila Leach, 1815 is renowned
for its enormous orb webs constructed with distinctive gold-
colored silk, for its extreme sexual size dimorphism (females are
gigantic compared to the males), and for being a conspicuous
inhabitant of tropical forests (Kuntner et al., 2013). Some two
dozen species are recognized in the genus, together with several
subspecies (World Spider Catalog, 2018). Golden orb weavers
inhabit tropical and subtropical regions throughout the world,
and the enormous, permanent webs of the females serve as
microecosystems for a variety of kleptoparasites and other
cohabitants (Vollrath, 1987; Tso and Severinghaus, 1998;
Agnarsson, 2003, 2010; Harvey et al., 2007).
Nephila and related genera (presently including Clitaetra
Simon, 1889, Herennia Thorell, 1877, Nephilengys Koch, 1872,
and Nephilingis Kuntner in Kuntner et al., 2013) were placed in
the family Araneidae Clerck, 1757 by Simon (1894), together
with other orb weavers, and close to the tetragnathines (Kuntner
et al., 2008). They remained in Araneidae until Levi (1986),
with doubt, and then Coddington (1990) transferred the nephi-
lines and tetragnathines into the family Tetragnathidae Menge,
1866. The nephiline genera were raised to family status
(Nephilidae Simon, 1894) in the work of Kuntner (2006), where
they remained (but closer to araneids than tetragnathids, e.g.,
Pan et al., 2004; Álvarez-Padilla and Hormiga, 2011; Su et al.,
2011) until Dimitrov et al. (2017) returned these genera to the
family Araneidae as subfamily Nephilinae, a result also sup-
ported by Wheeler et al. (2017).
Despite the large size of the females, most fossil nephilines
described are males in amber, mainly because of the need of
adult males to wander from their webs to seek out the sedentary
females. The youngest fossil nepheline described is Minu-
tunguis silvestris Wunderlich, 2011, a male in Quaternary
Madagascan copal. Miocene Dominican amber contains five
species of Nephila, all males, described by Wunderlich (1982,
1986), and Wunderlich (2004) described nine male nephilines
from Eocene Baltic and Bitterfeld amber, which he referred to
three new genera: Eonephila Wunderlich, 2004, Luxurionephila
Wunderlich, 2004, and Palaeonephila Wunderlich, 2004. The
only female nephiline known hitherto from the Cenozoic Era is
Nephila pennatipes Scudder, 1885, from Eocene beds at Flor-
issant, Colorado. This species most closely resembles the one
described here in size and geological age. Mesozoic nephilines
include the males Cretaraneus vilaltae Selden, 1990 from the
Early Cretaceous of Spain, Geratonephila burmanica (Poinar in
Poinar and Buckley, 2012) from mid-Cretaceous Burmese
amber, C. liaoningensis Cheng, Meng, and Wang in Cheng
et al., 2008 from the Early Cretaceous of China, and C. mar-
tensnetoi Mesquita, 1996 from the Early Cretaceous of Brazil.
However, the age of Geratonephila was disputed by Wunder-
lich (2015), who synonymized the genus with Nephila. From his
long experience of working with Burmese amber, during which
time he had never seen a nephiline in the deposit, Wunderlich
(2015) considered that Geratonephila was more likely from the
Dominican Republic, of Miocene age, in which deposit the
modern genus is quite common; he suggested it might belong to
Nephila tenuis Wunderlich, 1986. Similarly, the two spiders
from the Early Cretaceous of China and Brazil are most likely
not nephilines but were placed in the genus Cretaraneus
because of their Cretaceous age.
The only female Cretaceous nephilines known are several
large, undescribed specimens from the Early Cretaceous Crato
Formation of Brazil, one of which was figured by Dunlop and
Penney (2012, fig. 93). A giant female spider from the mid-
Jurassic Daohugou Fossil-Lagerstätte of China was originally
described as Nephila jurassica Selden, Shih, and Ren, 2011.
However, shortly after its description, a giant male was dis-
covered in the same beds, which was considered to be con-
specific with N. jurassica; the species was placed in the new
genus Mongolarachne Selden, Shih, and Ren, 2013 and
removed from Nephilinae. Kuntner et al. (2013) had already
Journal of Paleontology, 93(1), 2019, p. 98–104
Copyright © 2018, The Paleontological Society
0022-3360/15/0088-0906
doi: 10.1017/jpa.2018.71
98
https://doi.org/10.1017/jpa.2018.71
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