Content uploaded by George Poinar
Author content
All content in this area was uploaded by George Poinar on May 04, 2015
Content may be subject to copyright.
This article was downloaded by: [George Poinar]
On: 12 March 2014, At: 07:18
Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
Historical Biology: An International Journal of
Paleobiology
Publication details, including instructions for authors and subscription information:
http://www.tandfonline.com/loi/ghbi20
Pulchellaranea pedunculata n. gen., n. sp. (Araneae:
Araneidae), a new genus of spiders with a review of
araneid spiders in Cenozoic Dominican amber
George Poinar Jra
a Department of Zoology, Oregon State University, Corvallis, OR 97331, USA
Published online: 26 Feb 2014.
To cite this article: George Poinar Jr (2014): Pulchellaranea pedunculata n. gen., n. sp. (Araneae: Araneidae), a new genus
of spiders with a review of araneid spiders in Cenozoic Dominican amber, Historical Biology: An International Journal of
Paleobiology, DOI: 10.1080/08912963.2013.869799
To link to this article: http://dx.doi.org/10.1080/08912963.2013.869799
PLEASE SCROLL DOWN FOR ARTICLE
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained
in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no
representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the
Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and
are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and
should be independently verified with primary sources of information. Taylor and Francis shall not be liable for
any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever
or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of
the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematic
reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any
form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://
www.tandfonline.com/page/terms-and-conditions
Pulchellaranea pedunculata n. gen., n. sp. (Araneae: Araneidae), a new genus of spiders with a
review of araneid spiders in Cenozoic Dominican amber
George Poinar Jr*
Department of Zoology, Oregon State University, Corvallis, OR 97331, USA
(Received 6 May 2013; accepted 24 November 2013)
Pulchellaranea pedunculata n. gen.,n. sp. (Araneae: Araneidae), a new genus of orb web spiders, is described from
Cenozoic Dominican amber. Fossil representatives of the Araneidae are uncommon, even though the family is widely
distributed today. An arboreal ant adjacent to Pulchellaranea pedunculata indicates the likely habit of the fossil spider in the
Dominican amber forest. The encounter with the ant may have resulted in both specimens falling into fresh resin on the trunk
of the Algarroba tree. A key to araneid spiders reported in Dominican amber is provided.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:D7F92A3C-46BB-48C0-9BA1-0496038F04A5
Keywords: fossil orb web spider; Dominican Republic; Hispaniolae
Introduction
Members of the spider family Araneidae are a globally
distributed clade of orb web spiders ranging from 2 to
15 mm in size (Levi 1996,2001,2002). They are probably
one of the most frequently observed spider groups because
their symmetrical webs, often with the spider positioned in
the middle waiting for potential prey, are quite noticeable.
Many araneids are festooned with colourful patterns of
yellow, orange, green, black or white while the opistho-
soma of especially many tropical forms bear various types
of projections, humps or hooks. Representatives of six
genera were previously described from Dominican
amber (Wunderlich 1988;Penney2008). The
present study describes a new genus of Araneidae from
this source.
Materials and methods
The specimen, which is well preserved and complete,
was obtained from mines in the Cordillera Septentrional of
the Dominican Republic. Dating of Dominican amber is
still controversial with the latest purposed age of 20–
15 mya based on Foraminifera (Iturralde-Vinent
and MacPhee 1996) and the earliest as 45 –30 mya based
on coccoliths (Ce
ˆpek in Schlee 1990). A range of ages for
Dominican amber is possible as the amber is associated
with turbiditic sandstones of the Upper Eocene to
Lower Miocene Mamey Group (Draper et al. 1994).
Dominican amber was produced by the leguminous tree,
Hymenaea protera Poinar, and a re-construction of the
Dominican amber forest based on amber fossils indicated
that the environment was similar to that of a present-day
tropical moist forest (Poinar and Poinar 1999). Obser-
vations and drawings were made with a Nikon SMZ-10
stereoscopic microscope (Nikon, Nippon Kogaku K. K.,
Tokyo, Japan). The amber piece containing the fossil is
sub-rectangular in outline, measuring 10 mm in greatest
length, 9 mm in greatest width and 6 mm in greatest
thickness.
Preservation
The quality of preservation of the fossil spider is excellent
and many morphological features have been preserved in
lifelike condition. The pristine condition of the present
fossil is thought to be the result of fixation of the tissues
from chemicals contained in the original resin as it flowed
from the tree source. While dehydration is also an
important part of the preservation process, it can cause
structural distortions, especially in some soft-bodied
arthropods. Fortunately, the fossil spider shows little
sign of distortions, only possibly some partial collapsing of
the legs. The most common alterations of arthropods in
amber are colour changes, especially a darkening of the
integument.
Systematics
Characters that align the fossil with the family Araneidae
are three claws, eight eyes in two rows with the lateral eyes
adjacent yet some distance from the medians, a clypeal
height under two diameters of the anterior median eyes, a
globose abdomen overhanging the carapace and legs with
macrosetae (Levi 2002).
q2014 Taylor & Francis
*Email: poinarg@science.oregonstate.edu
Historical Biology, 2014
http://dx.doi.org/10.1080/08912963.2013.869799
Downloaded by [George Poinar] at 07:18 12 March 2014
Pulchellaranea Poinar, n. gen.
Description. Carapace subequal in length and width, sides
convex, cephalic portion not raised; sternum roundish, wider
than long, lateral margins project between coxae; anterior
border of sternum surrounds basal portion of triangular-
shaped labium; labium twice as wide as long; clypeus length
less than two diameters of the anterior median eyes; anterior
median eyes prominent, one diameter apart, nearly twice
diameter of posterior median eyes; posterior median eyes
facing dorsad, twice diameter of lateral eyes; lateral eyes
contiguous, positioned at tip of peduncles arising from the
upper edge of carapace; peduncles swollen slightly at tip;
posterior row of eyes straight; palps with median and
terminal apophyses subequal in length; opisthosoma pale
yellow, wider than long, attached one-quarter distance from
anterior end and overhanging spinnerets; anterior edge of
opisthosoma rounded, with three pairs of sigilla, posterior
edge notched, with four pairs of posteriorly directed hook-
like protrusions.
Etymology. The generic name is from the Latin
‘pulcher’ for beautiful and the Latin ‘aranea’ for spider.
Type species.Pulchellaranea pedunculata Poinar
(Figures 1– 8).
Diagnosis. The new genus is distinguished from extant
genera of araneids by the following combinations of
characters: the presence of pedunculated, terminal lateral
eyes, the shape of the carapace, sternum, labium and
opisthosoma and the structure of the palps. The fossil shows
some similarities with members of the following extant
genera. Members of Enacrosoma Mello-Leita
˜o, 1932 also
have a wider than long opisthosoma with pairs of lateral
humps, but the opisthosoma is rectangular and the humps
occur on the anterior and lateral as well as the posterior
surfaces. The opisthosoma is also marked with paired,
indistinct darker patches and spots, and there is a sclerotised
ring round the spinnerets. The males have a narrow cephalic
region set off from the remainder of the carapace and the
shape of the palps is quite different from those of the present
fossil (Levi 2001).
The fossil shows some resemblance to the Domin-
ican amber Enacrosoma verrucosa (Wunderlich, 1988)
that was originally tentatively assigned to the genus
Cyclosa (Wunderlich 1988) but then transferred to the
genus Enacrosoma by Penney (2001). However, the
lateral eyes of E. verrucosa are juxtaposed on a slight
rise on the carapace while those of the fossil are on
distinct peduncles. Also the opisthosoma of E. verrucosa
has only three pairs of hooks while that of Pulchellar-
anea pedunculata has four pairs. In addition, E.
verrucosa has only one pair of sigilla while Pulchellar-
anea pedunculata has three pairs. The long, slender,
median apophysis of E. verrucosa is also quite different
from the short, thick median apophysis of Pulchellar-
anea pedunculata.
Figure 1. Dorsal view of Pulchellaranea pedunculata n. gen.
et sp. adjacent toan ant of the genus Pseudomyrmex (Hymenoptera:
Formicidae) in Dominican amber. Scale bar ¼1.3 mm.
Figure 2. Dorsal view of Pulchellaranea pedunculata n. gen.
et sp. in Dominican amber. Scale bar ¼1.1 mm.
2G. Poinar Jr.
Downloaded by [George Poinar] at 07:18 12 March 2014
Males of the genus Hypognatha Gue
´rin-Me
´neville,
1840 also have lateral eyes on the tips of projections but
the projections are positioned on the front of the head
region beneath the median eyes. This genus is also
characterised by a pattern of scutes on the dorsum of the
opisthosoma and a posterior notch in the sternum, neither
of which occurs in the present fossil.
Figure 3. Ventral view of Pulchellaranea pedunculata n. gen.
et sp. in Dominican amber. S, sternum. Scale bar ¼1.0 mm.
Figure 4. Dorsal view of opisthosoma of Pulchellaranea
pedunculata n. gen. et sp. Note four pairs of hooks (arrows) and
three pairs of sigilla. Scale bar ¼500 mm.
Figure 5. Dorsal view of head of Pulchellaranea pedunculata
n. gen. et sp. showing anterior median eyes, posterior median
eyes and lateral eyes positioned at the tips of peduncles (arrows).
Scale bar ¼170 mm.
Figure 6. ‘En face’ view of Pulchellaranea pedunculata n. gen.
et sp. showing anterior median eyes, clypeus and chelicerae.
Scale bar ¼140 mm.
Historical Biology 3
Downloaded by [George Poinar] at 07:18 12 March 2014
In Encyosaccus Simon, 1895, the carapace is as wide in
the head region as it is in the thoracic region and the dorsum
of the opisthosoma is covered with sclerotised discs.
Members of the genus Xylethrus Simon, 1895 have a
wide sternum as occurs in the fossil but also have the
opisthosoma covered with sclerotised disks and the
paramedian process on the male palp is drawn out into a
thread (Levi 1996). Species of Gasteracantha Sundwall,
1833 also have a wide sternum, but males have the median
eyes set off from the rest of the carapace, and the latter is
narrowed anteriorly with straight sides. In addition, the
dorsum of the opisthosoma has dark markings and the palp
has a circular paramedian apophysis, which is not the case in
the present fossil (Levi 1996).
The fossil shares some characters with Cyclosa Menge,
1866 but members of this genus have adjacent posterior
median eyes, non-pedunculated lateral eyes, and the thoracic
region is raised and often set off from the cephalic portion.
The opisthosoma can be spherical but is usually longer than
wide, pointed and often bearing tubercles (Levi 2002).
The fossil can be differentiated from members of the
genus Molinaranea Mello-Leitao, 1940 that have a setose
abdomen longer than wide with at most two humps and an
anterior median or posterior median bulge (Levi 2002). The
genus Acanthepeira Marx, 1883 is characterised by lateral
eyes situated on the sides of the projections, not at the apex
as in Pulchellaranea pedunculata. Also the abdomen of
Acanthepeira has large spines and the palpal sclerites are
partly covered by a large cymbium, which is not the case
with Pulchellaranea pedunculata. Species of Parawixia F.
O.P. Cambridge, 1904 have soft or setose sides of the
carapace and abdomen, and the latter is as long as wide or
longer than wide. Also, the abdomen is attached at the
anterior end and does not overlap the spinnerets. In addition,
the posterior median eye (PME) eye diameter is subequal to
that of the other eyes. None of these characters resemble the
condition on Pulchellaranea pedunculata.
Pulchellaranea pedunculata Poinar, n. sp.
(Figures 1– 8)
Holotype male. Body length, 2.9 mm: legs, palps and
carapace reddish brown; opisthosoma pale yellow.
Carapace. Length (dorsal), 1.5 mm, width, 1.3 mm;
clypeus length 110 mm, less than the diameter of anterior
median eye; anterior median eyes large, protruding
anteriorly, diameter, 170mm; length of peduncles support-
ing contiguous lateral eyes, 160 mm; all coxae bearing from
two to four tubercles; all femora spined; tibia without spurs,
femur I longer than femur IV; tibia IV with 1/1/1 bristles
(Figures 1– 3). See Table 1 for lengths of leg segments.
Palps. Palpal lengths (from dorsal view) 950 and
900 mm; palpal widths, 280 and 300 mm; setae not
observed on palpal patella; palp with elongate cymbium
extending from tibia; median and terminal apophyses
subequal in length (Figures 2, 7 and 8).
Opisthosoma. Wider than long, length, 1.5 mm, width,
1.6 mm; pale yellow, covered with minute tan tubercles,
each bearing short tan or white seta; attached to carapace at
anterior fourth, rounded anteriorly, overhanging spinnerets,
sclerotised ring around spinnerets absent (Figures 1 – 4).
Material examined. Holotype male (accession # A-10-
267) from the northern mountain range in the Dominican
Republic deposited in the Poinar amber collection
maintained at Oregon State University. Female unknown.
Etymology. The specific epithet is from the Latin
‘pedunculus’ for foot in reference to the pedunculated
lateral eyes.
Diagnosis. As for the genus.
Figure 7. Dorsal view of head of Pulchellaranea pedunculata
n. gen. et sp. showing palps. Scale bar ¼410 mm.
Figure 8. Right palp of Pulchellaranea pedunculata n. gen. et
sp. CY, cymbium; F, femur; MA, median apophysis; P, patella;
PC, paracymbium; ST, subtegulum; T, tibia; TA, terminal
apophysis; TG, tegulum. Scale bar ¼280 mm.
4G. Poinar Jr.
Downloaded by [George Poinar] at 07:18 12 March 2014
Discussion
Male spiders of various groups may have one or more pairs
of eyes on projections, presumably related to finding,
mating with and then escaping from females. The length of
the projections can vary considerably on different genera.
The lateral eyes on males of Hypognatha can extend to
half the head width (Levi 1996) while the Mexican amber
Mirometa valdespinosa Petrunkevitch, 1963 has all eight
eyes on separate projections (turrets) oriented in different
positions (Petrunkevitch 1963).
Presently, there are some 325 species of extant spiders
in 182 genera and 45 families in Hispaniola and already
some 170 fossil spider species in 99 genera and 45 families
from Dominican amber. All of the amber species and 26%
of the genera are extinct (Penney 2008; Wunderlich 1988).
Fossil representatives of the Araneidae are uncommon,
even though the family dates back to the Early Cretaceous
(Penney and Ortun
˜o2006) and is widely distributed today.
While 58 species of extant araneids in 27 genera occur in
Hispaniola, only representatives of three of these genera
occur in Dominican amber. Representatives of the
following extant and extinct genera have been reported
from Dominican amber: Araneus Clerck, 1757; Araneo-
meta Wunderlich, 1988;Enacrosoma Mello-Leita
˜o, 1932;
Fossilaraneus Wunderlich, 1988;Molinaranea Mello-
Leita
˜o, 1940; Pycnosinga Wunderlich, 1988 and Pulchel-
laranea Poinar n. gen. (Wunderlich 1988; Penney 2008).
The following key separates the males (corresponding
females remain unknown) of the nine species of araneids
thus far described from Dominican amber:
1. Cephalic portion of carapace raised (expanded) in
area of posterior median eyes ... ... ... ... ... .....2
– Cephalic portion of carapace not raised in area of
posterior median eyes ... ... ... ... ... ... ... ..... 4
2. Both pairs of lateral eyes directed downwards
... ... ... ... ... ... ... ....... Araneometa excelsia
Wunderlich, 1988
–Both pairs of lateral eyes not directed downwards
... ... ... ... ... ... ... ... ... ... ... ... ... ...... 3
3. Both pairs of lateral eyes directed
laterally; opisthosoma with long bristles; second pair
of sigilla spheroid ... ... ... Araneometa herrlingi
Wunderlich, 1988
– Posterior pair of lateral eyes directed laterally;
anterior pair of lateral eyes directed downward;
opisthosoma with short bristles; second pair of
sigilla elongate, transverse ... ... ... ... ... ... ...
Araneometa spirembolus Wunderlich, 1988
4. Carapace and abdomen setose; median apophysis
long, thin, with subequal terminal prongs
... ... ... ... ... ... Molinaranea mitnickii Saupe,
Selden and Penney, 2010
– Not as above ... ... ... ... ... ... ... ... ... ..... 5
5. Each pair of lateral eyes on peduncles protruding
outward from carapace; opisthosoma with four pairs of
hooks and three pairs of sigilla ... ... ... ... ... ...
Pulchellaranea pedunculata n. gen.,n. sp.
– Lateral eyes not on erect peduncles protruding
outward from carapace; opisthosoma with three pairs
of hooks and one pair of sigilla or lacking both hooks
and sigilla ... ... ... ... ... ... ... ... ... ... ......6
6. Femora lacking spines ... ... ... ... ... ... ...
Pycnosinga fossilis Wunderlich, 1988
– Femora with spines ... ... ... ... ... ... ... ..... 7
7. Opisthosoma with three pairs of hooks and one pair
of sigilla ... ... ... ... .. . ... ........ E. verrucosa
(Wunderlich, 1988)
– Opisthosoma lacking hooks and sigilla
... ... ... ... ... ... ... ... ... ... ... ... ... ...... 8
8. Tibia IV ventral with 1/1 bristles, paracymbium
short ... ... ... ... ... Araneus nanus Wunderlich,
1988
– Tibia IV ventral lacking bristles, paracymbium long
... ... ... ... ... ... ... ...... Fossilaraneus incertus
Wunderlich, 1988.
The ant (Figure 1) adjacent to Pulchellaranea
pedunculata indicates the habit of the fossil spider in the
amber forest. Members of Pseudomyrmex (Hymenoptera:
Formicidae) are small ants that move quickly over tree
surfaces. Therefore, it is likely that Pulchellaranea
pedunculata was an arboreal species that possibly
encountered the ant, with both falling into some fresh
resin on the trunk of the Algarroba tree.
Acknowledgements
The author thanks Joerg Wunderlich, Herbert Levi and Pat Craig
for discussions regarding morphological characters of the fossil,
Cesare Baroni Urbani for identifying the amber ant and Paul
Seldon, Art Boucot and Roberta Poinar for comments on earlier
drafts of the manuscript.
References
Draper G, Mann P, Lewis JF. 1994. Hispaniola. In: Donovan S, Jackson
TA, editors. Caribbean geology: an introduction. Kingston: The
University of the West Indies Publishers’ Association. p. 129 – 150.
Iturralde-Vinent MA, MacPhee RDE. 1996. Age and paleogeographic
origin of Dominican amber. Science. 273:1850 – 1852.
Table 1. Lengths of leg segments of Pulchellaranea peduncu-
lata.
Leg segment Leg 1 Leg 2 Leg 3 (mm) Leg 4 (mm)
Femur 1.4 mm 1.1 mm 370 700
Patella 430 mm 430 mm 200 330
Tibia 900 mm 670 mm 470 730
Metararsus 830 mm 600 mm 430 600
Tarsus 370 mm 370 mm 233 370
Historical Biology 5
Downloaded by [George Poinar] at 07:18 12 March 2014
Levi HW. 1996. The American orb weavers Hypognatha, Encyosaccus,
Xylethrus,Gasteracantha, and Enacrosoma (Araneae, Araneidae).
Bull Mus Comp Zool. 155:89–157.
Levi HW. 2001. The orbweavers of the genera Molinaranea and
Nicolepeira, a new species of Parawixia and comments on orb-
weavers of temperate South America (Araneae: Araneidae). Bull
Mus Comp Zool. 155:445–476.
Levi HW. 2002. Keys to the genera of araneid orbweavers (Araneae,
Araneidae) of the Americas. J Arachnol. 30:527 – 562.
Penney D. 2008. Dominican amber spiders. Manchester: Siri Scientific
Press, 176 p.
Penney D, Ortun
˜o VM. 2006. Oldest true orb-weaving spider (Araneae:
Araneidae). Biol Lett. 2:447–450.
Petrunkevitch A. 1963. Chiapas amber spiders. In: Studies of fossiliferous
amber arthropods of Chiapas. Vol. 31. Mexico. Univ. Calif. Publ.
Entomol. p. 1–40.
Poinar GOJr, Poinar R. 1999. The amber forest. Princeton, NJ: Princeton
University Press, 239 p.
Saupe EE, Selden PA, Penney D. 2010. First fossil Molinaranea
(Araneae: Araneidae), from Middle Miocene Dominican amber,
with a phylogenic and paleobiogeographic analysis of the genus.
Zool J Linn Soc. 158:711–725.
Schlee D. 1990. Das Bernstein-Kabinett. Stuttg Beitr Naturk (C). 28:100.
Wunderlich J. 1988. Die Fossilen Spinnen im Dominikanischen
Bernstein. Beitr Araneol. 2:1–378.
6G. Poinar Jr.
Downloaded by [George Poinar] at 07:18 12 March 2014
