dominican amber spiders

Forthcoming title due to be
published May 2008
Dominican
AA comparative
mber Spiders
palaeontologicalneontological approach to identification,
faunistics, ecology and biogeography
Dr David Penney
Siri Scientific Press
Special Offer on
pre-publication orders!
(until 15th April 2008)
Please see below for further details and sample pages
Book details and ordering information
Title: Dominican Amber Spiders
Subtitle: A comparative palaeontological–neontological approach to
identification, faunistics, ecology and biogeography
Author: David Penney
Publisher: Siri Scientific Press
ISBN: Forthcoming
Publication date: Expected May 2008
Dimensions: 245 by 170 mm
Cover: Soft (300 gsm, gloss laminated)
Number of pages: 178 (130 gsm silk coated paper)
Number of illustrations: 300+ including high quality colour photos
Summary: see attached sample pages
Price: £40.00 (postage, packaging & handling extra, but free for prepublication orders*)
Methods of payment
Paypal: please make payment to spiderdavep
Direct bank transfer: for further details please email the author on
[email protected]
Cheque or international money order: please make payable (in £GBP
sterling) to David Penney and send to: Dr David Penney, 50 Burnside
Drive, Burnage, Manchester, M19 2LZ, United Kingdom
Please note that only payment in Sterling (Great British Pounds) can be accepted.
Whichever form of payment you use, please email the following information to
[email protected]: your name, delivery address and method of
payment (please put Amber Spider Book in the email subject field). Receipts will
be issued when the book is despatched. Thank you!
*refers to regular, surface mail only. The publisher and author accept no responsibility in the unlikely
event of orders lost in transit. Delivery by registered mail or courier can be arranged, with additional
costs paid by the purchaser. Please contact the author for details.
David Penney
A comparative palaeontologicalneontological approach to
identification, faunistics, ecology
and biogeography
Siri Scientific Press
Bar code here
Dominican Amber Spiders
The author is a Visiting Research Fellow at the University of Manchester,
UK and the leading world expert on fossil spiders preserved in amber and
in interpreting what they can reveal about the ecology of the extinct forests
in which they lived. In this book Dr Penney provides a comprehensive
synopsis of what is known about the Dominican Republic amber spider
fauna, much of which is based on his numerous scientific publications in
leading international journals. However, the book is not intended solely for
academics. It contains more than 300 illustrations including many colour
photographs, which should permit the identification of both the fossil and living
Hispaniolan spider fauna by both amber collectors and spider enthusiasts.
The introductory chapters provide full coverage of what is known about the
geological origins, chemistry and botanical source of Dominican amber and
the mining, preparation and distribution processes. which the author has
witnessed first hand. Previously unpublished data on historical biogeography
should make this book of interest to all those interested in the biogeography
of the Caribbean region. The volume also contains an extensive bibliography
of almost 350 entries providing a valuable resource for anybody interested in
fossil resins. This book far surpasses anything else availabe on this subject
and is expected to remain the leading reference work for many years to come.
Dominican
Amber Spiders
Dr David Penney
Foreword
It is not possible to fully appreciate the origins, evolutionary history or present biodiversity of any group
of organisms until one considers the fossil record. Yet despite this obvious statement, there often remains
a palpable void between the disciplines of palaeontology and neontology, and scientists from both camps
are to blame. More often than not, neontologists will not even consider fossils in systematic studies or when
describing new taxa. However, this is not a universal phenomenon and I am encouraged by the increasing
number of neontologists who do consider fossils. In some cases they may be unaware that fossils exist, but given
the electronic bibliographic databases that are currently available this is not really a credible excuse. Others
argue that fossils do not preserve sufficient detail for comparative studies with recent specimens. However,
as you will see in this volume, recent applications of high-resolution imaging technology to palaeontological
specimens render this argument one of semantics or methodology. Some neontologists have told me they do
not consider fossils in their studies because they are unimportant and they remain steadfast in this respect
despite my repeated attempts to convince them otherwise. A perfect example of just how important it is
to consider fossils relates to the spider family Archaeidae, which was first described in 1854 from fossils
preserved in Baltic amber, prior to being found in the extant fauna three decades later. Not only are fossils of
this family important from a taxonomic and nomenclatorial perspective, but fossils can also shed much light
on the historical biogeography of this family. The extant fauna is restricted to South Africa and Madagascar,
whereas fossils are known from the Baltic region, France, Myanmar, China and Kazakhstan.
Many palaeontologists are accidental zoologists. For some, having graduated from an Earth Sciences
degree course, most of their academic background lies in the physical, rather than biological sciences. Many
are unfamiliar with the finer details of taxonomic practices for extant members of the groups they then choose
to go on and study. There are many examples in the literature of new fossil taxa that have been erected solely on
the basis of age. In many cases, there is no indication that comparative extant specimens have been examined,
the diagnoses are often based on features not considered reliable for extant species, nor do the diagnoses serve
to differentiate the fossil taxa from living forms. Given that many groups of organisms demonstrate a high
degree of evolutionary stasis, such an approach is inappropriate. Using such an approach makes comparisons
of neontological and palaeontological faunistic datasets (e.g., in order to assess any degree of change over
time) impossible or highly misleading. As a zoologist turned palaeontologist I sit in both camps and suppose
I am an accidental geologist. I also have a lot of catching up to do! Ultimately however, researchers from
both these (and other) disciplines need to enter into each others’ realms in order to address effectively the
increasingly complex macro-scale evolutionary and biological questions that we currently pose in order to
secure research funding to understand the history of life on Earth.
This book has four main purposes. Firstly, to demonstrate a perfect example of how fossil and
extant faunas can be extremely similar and thus warrant a combined neontologicial and palaeontological
taxonomic approach. Secondly, to show how comparisons of faunas based on such an approach can permit
qualitative and quantitative investigations of interesting palaeo/biological questions, for example, relating to
palaeoecology and historical biogeography. Thirdly, to provide comprehensive information on our current
knowledge regarding the origins and formation of Hispaniola and Dominican Republic amber, including
methods for preparation and study. Finally, the book serves as an introduction to the fossil record of spiders,
and specifically, the identification of spiders preserved as fossils in Dominican Republic amber (and also
from the extant Hispaniolan fauna). It is not intended as an exhaustive guide for species identification, but
should permit the reader to identify their specimen at least to family, and in some cases even further. Readers
will then be directed to the relevant literature required to take their identifications to species level. Thus, this
volume is aimed at a broad audience and some will have a better knowledge of the subject than others. Rather
than include a comprehensive glossary, I have attempted to limit technical terms and explain them where they
appear in the text.
Finally, it is easy to start an endeavour such as this, and indeed it was a labour of love. However, it
is not so easy to know when to stop. New fossil and extant species are being discovered and described all
Contents
1. Introduction................................................................................................................................................10
Caught in the act...................................................................................................................................11
What are spiders and how do they fit into the grand scheme of things?..............................................13
Spiders in the fossil record...................................................................................................................15
Age, radiations and extinction events...................................................................................................17
What is the importance of fossil spiders and why should we study them?..........................................19
2. Dominican Republic amber.......................................................................................................................21
Some history.........................................................................................................................................22
A note on terminology..........................................................................................................................23
Botanical source and age of Dominican amber....................................................................................23
The amber producing tree........................................................................................................23
Age of Dominican amber.........................................................................................................25
Physical and chemical properties.........................................................................................................26
Tissue and DNA preservation..............................................................................................................27
Authenticity (distinguishing amber from copal and fakes).................................................................28
The mining process..............................................................................................................................28
The amber mines......................................................................................................................29
The extraction process.............................................................................................................29
From mine to museum..............................................................................................................30
Methods of preparation and study........................................................................................................31
Preparation of raw amber inclusions......................................................................................31
Further preparation for closer scrutiny of inclusions..............................................................33
Light microscopy and photography.........................................................................................34
Advanced microscopy and computed tomography..................................................................35
Major collections of Dominican amber................................................................................................37
Conservation and curation of amber collections..................................................................................37
The diversity of Dominican amber inclusions.....................................................................................38
3. History of Hispaniolan araneology...........................................................................................................40
The extant fauna..................................................................................................................................40
The fossil fauna....................................................................................................................................41
Systematic checklist of fossil and extant Hispaniolan spiders............................................................42
4. Key to Hispaniolan spider families (fossil & extant)..............................................................................48
Morphology and terminology..............................................................................................................48
Key to the spider families of Hispaniola.............................................................................................52
Highly distinctive morphological features...............................................................................52
Dichotomous key to all Hispaniolan spider families...............................................................55
5. Family descriptions....................................................................................................................................65
Dipluridae.........................................................................................................................................................66
Cyrtaucheniidae................................................................................................................................................67
Microstigmatidae..............................................................................................................................................67
Barychelidae.....................................................................................................................................................68
Theraphosidae..................................................................................................................................................69
Filistatidae........................................................................................................................................................70
Sicariidae..........................................................................................................................................................71
Scytodidae........................................................................................................................................................72
Drymusidae......................................................................................................................................................73
Ochyroceratidae...................................................................................................................................73
Pholcidae..........................................................................................................................................................75
Caponiidae........................................................................................................................................................77
Tetrablemmidae................................................................................................................................................78
Segestriidae......................................................................................................................................................78
Oonopidae........................................................................................................................................................79
Palpimanidae....................................................................................................................................................81
Mimetidae.........................................................................................................................................................81
Oecobiidae........................................................................................................................................................83
Hersiliidae........................................................................................................................................................84
Deinopidae.......................................................................................................................................................86
Uloboridae........................................................................................................................................................87
Nesticidae.........................................................................................................................................................89
Theridiidae.......................................................................................................................................................90
Theridiosomatidae............................................................................................................................................94
Symphytognathidae..........................................................................................................................................95
Anapidae...........................................................................................................................................................96
Mysmenidae.....................................................................................................................................................96
Linyphiidae.......................................................................................................................................................97
Nephilidae........................................................................................................................................................99
Tetragnathidae................................................................................................................................................101
Araneidae.......................................................................................................................................................102
Lycosidae.......................................................................................................................................................106
Pisauridae...........................................................................................................................................107
Oxyopidae......................................................................................................................................................108
Zoridae...........................................................................................................................................................108
Ctenidae..........................................................................................................................................................109
Desidae...........................................................................................................................................................110
Dictynidae......................................................................................................................................................111
Amaurobiidae.................................................................................................................................................112
Miturgidae......................................................................................................................................................113
Anyphaenidae.................................................................................................................................................114
Liocranidae.....................................................................................................................................................116
Clubionidae....................................................................................................................................................116
Corinnidae......................................................................................................................................................117
Trochanteriidae...............................................................................................................................................119
Prodidomidae.................................................................................................................................................120
Gnaphosidae...................................................................................................................................................120
Selenopidae....................................................................................................................................................122
Sparassidae.....................................................................................................................................................122
Philodromidae................................................................................................................................................124
Thomisidae.....................................................................................................................................................124
Salticidae........................................................................................................................................................125
6. Aspects of palaeoecology & historical biogeography............................................................................129
The Miocene Dominican amber forest..............................................................................................130
Resin as a trap: taphonomy and bias of amber spider inclusions......................................................130
The site of resin secretion.......................................................................................................130
The entrapment process..........................................................................................................131
Do different ambers trap organisms in the same way?.........................................................133
Bias in the amber fauna..........................................................................................................135
Comparison of the fossil and extant spider faunas............................................................................137
Origins of the Hispaniolan spider fauna............................................................................................142
Predictions for the extant fauna based on the fossil fauna................................................................150
7. Other fossil arachnids in Dominican amber.........................................................................................152
References cited............................................................................................................................................156
Index.............................................................................................................................................................171
10
Introduction
When most people hear the word ‘fossil’ they tend to conjure up images of giant dinosaurs such as Tyrannosaurus
rex or shelled marine molluscs. Prior to the Hollywood blockbuster movie Jurassic Park, which was based on
recreating dinosaurs through extracting their DNA from blood in the guts of fossil mosquitoes preserved in
amber, few people would entertain the notion that insects occur in the fossil record. However, insects, spiders
and other terrestrial arthropods are common as fossils (Grimaldi & Engel, 2005) and particularly so in amber,
where they are often preserved with life-like fidelity.
Amber has properties similar to amorphous, polymeric glass and is the fossilized form of tree resin,
which consists of a complex mixture of terpenoid and/or phenolic compounds. It is derived from numerous
different tree families (e.g., Pinaceae, Araucariaceae, Cupressaceae, Leguminoseae, Combretaceae to name
but a few), most of which can be determined by comparing the infrared spectra of the amber with those of
resins from extant tree species. Such techniques also serve to differentiate one amber type from another. For
example, Baltic amber has a highly characteristic plateau off one side of one of the spectrograph peaks, which
is referred to as the ‘Baltic shoulder’.
The oldest ambers that contain fossil arthropods are from the Lower Cretaceous of Lebanon (Poinar
& Milki, 2001) and Jordan (Kaddumi, 2005), although there is no reason not to expect future discoveries of
inclusions in older (e.g., Jurassic) fossil resins. Various deposits from around the world (Figure 1.1) fill in
the gaps from the Cretaceous until the present, although hardened resins younger than 40,000 years of age
are considered to be sub-fossil and are called copal (not shown in Figure 1.1). Amber is a prime example of
a Konservat-Lagerstätte (an occurrence of exceptional preservation, which permits detailed morphological
comparisons with living relatives) and so in contrast to most fossils preserved in sediments, amber inclusions
are particularly important from a phylogenetic perspective and can be used to investigate micro- as well as
macro-evolutionary processes.
11
1.1 Amber deposits of the world. Data primarily from Martínez-Delclòs et al., 2004) with updates
Caught in the act
Syninclusions (two or more inclusions in the same piece of amber) often preserve interactions between
organisms, for example, behaviours such as mating (Figure 1.2), mate guarding, commensalism, parasitism
(e.g., nematode worms exiting their arthropod host, hymenopteran larvae and mites preserved in situ, still
penetrating the cuticle of their host organism), disease (e.g., pathogenic fungi), defecation, egg laying, phoresy
(one organism being transported by another—pseudoscorpions are sometimes found attached to the legs of
insects), predation and maternal care (e.g., ants carrying larvae and pupae). Even delicate spider webs are
occasionally preserved, sometimes with prey attached (Figure 1.3). So excellent is the degree of preservation
in amber that even the tiny glue droplets of spider viscid silk are often clearly visible (Figure 1.4), even
in Cretaceous ambers dating back to in excess of 100 million years (Zschokke, 2003, 2004). Amber may
also preserve the death throes of the entombed arthropods as they struggled to escape the sticky exudates,
for example, in the form of wing movements and disarticulation of body parts. Fossilized spider blood was
reported recently, exuding from joints where the resin had caused the legs to autospasize (detach as a result of
an external force) from the body (Penney, 2005a; Figure 1.5).
Such observations as those listed above are rarely, if ever, encountered in the non-amber fossil record
because of the different taphonomic processes (what happens to the organism between death and becoming
a fossil) that control the preservation of organisms in carbonate rocks and amber (Martínez-Delclòs et al.,
2004). However, I do not wish to imply here that the non-amber fossil record is any less important. An
additional consequence of this variation is that, on the whole, sediments and amber preserve different subsets
of the arthropod community. Thus, the fossil record of amber is biased towards preserving certain groups,
as are all other fossil deposits. In addition, the fossil record in general is notoriously incomplete, but not
27
exuded, the volatile components of the resin, such as sesquiterpene hydrocarbons which function to control
viscosity and flexibility, are gradually lost to produce hardened resin. In general, the processes involved in
fossilization seem to be progressive oxidation and polymerization via free-radical mechanisms. In contrast to
Baltic amber, Dominican amber does not contain succinic acid. Thus, the new species name Staphylococcus
succinus erected by Lambert et al. (1998) for a newly discovered species of bacterium in Dominican amber is
slightly inappropriate.
Tissue and DNA preservation
The mode of tissue preservation in amber appears to be a rapid and thorough fixation and dehydration, which
may be sufficient for preserving deoxyribonucleic acid (DNA) more consistently than any other kind of
fossilization process (Grimaldi et al., 1994a). Henwood (1992) reported excellent preservation of soft tissues,
including locomotory, digestive, respiratory, nervous and sensory tissues from cantharid and nitulid beetles
preserved in Dominican Republic amber. However, not all ambers preserve internal tissues in such detail.
For example, the recently discovered amber from western Amazonia appears to preserve only the cuticular
exoskeleton of the arthropod inclusions (Antoine et al., 2006).
During the early 1990s numerous researchers claimed that they had managed to extract small strings
of DNA from Dominican amber inclusions, such as the stingless bee Propblebeia dominicana (Cano et al.,
1992), a termite Mastotermes electrodominicus (DeSalle et al., 1992, 1993), a wood gnat Valeseguya disjuncta
(DeSalle, 1994) and the amber producing tree itself (Poinar et al., 1993). Even more impressive were claims
by Cano et al. (1993) of DNA extraction from a beetle Libanorhinus succinus in Lebanese amber, dating back
as far as 135 million years. A number of publications rode the crest of the amber DNA wave (e.g., Poinar,
1994), including a whole book by Poinar & Poinar (1994), which is a very pleasant read and particularly useful
for those intending to visit amber producing areas.
However, an excellent study by Austin et al. (1997) at the Natural History Museum, London, which
employed a rigorous experimental method with appropriate control groups, was unable to replicate the abovementioned studies. DNA was retrieved (including from the control samples, which should not have contained
any!) but all sequences detected were derived from obvious non-insect contaminants, such as vertebrates
and fungi. Thus, they concluded that the likelihood of DNA being preserved in amber was minimal, but
such negative results cannot disprove its existence. Kimberly et al. (1997) were unable to replicate DNA
extraction from Dominican amber Propblebeia, despite using nine specimens and nested amplifications,
which greatly enhanced the sensitivity of the assay process. Stankiewicz et al. (1998) found that relatively
resistant macromolecules, such as lignin, cellulose and chitin were extremely degraded in Dominican amber
plant and insect inclusions, concluding that the persistence of fragile molecules such as DNA would be highly
improbable.
This century, more recent claims (Rogers et al., 2000) have proposed that rapidly dehydrated inclusions
may contain undamaged DNA. However, Hebsgaard et al. (2005) pointed out that many research groups
currently accept one hundred thousand to one million years as the maximum age range for DNA survival,
based on both theoretical and empirical data, and that despite the problems associated with modelling the
long-term survival of DNA in the geosphere, the claims of DNA from amber (1,000-fold older than theoretical
predictions for maximal DNA survival) are cause for concern, regardless of how good a preservative the natural
resin is. According to these authors, all claims for amber DNA to date suffer from inadequate experimental
design and insufficient authentification criteria. Some of the earlier proponents for DNA extraction have now
held up their hands and admitted that it looks like the end of the road for fossil DNA, whereas others still
support their findings; the burden of proof now lies with the latter. The jury is still out on whether or not it
is possible to extract DNA from amber, or indeed whether it remains in tact in fossilized resin, but given the
evidence to hand it would seem unlikely.
A particularly interesting study by Koller et al. (2005) used a new technique for internal fixing of
amber inclusions in association with electron microscopy and histochemical staining. They noted that cellular
29
the principal amber deposits of the eastern Baltic (Wu,
1997). Under the close scrutiny of Rafael Trujillo,
the then dictatorial head of the government, a West
German company began extracting and exporting
unprocessed amber in large quantities. However,
it wasn’t long before the Dominican government
withdrew the mining rights for political and economic
reasons and the company collapsed shortly thereafter.
The government also prohibited the export of
unprocessed amber, permitting only finished goods to
leave the country (Wu, 1997).
The amber mines
The mines in the Cordillera Septentrional are located 2.5 A La Toca amber mine in the Cordillera Septentrional
slightly north of Santiago (Figure 2.1) and are
often named after local villages. They include: Palo
Quamado, Palo Alto, Juan de Nina, La Toca, Las
Cacaos, Los Aguitos, La Cumbre, Los Higos, La
Bucara, Pescado Bobo, El Naranjo, Las Auyamas,
El Arroyo, Aquacate, Carlos Diaz and Villa Trina.
Most of these mines are located 800–1,000 m above
sea level and can only be accessed by hiking along
steep mountain trails. Those in the Cordillera Oriental
are situated at elevations of less than 200 m above
sea level and include: La Medita and Ya Nigua near
El Valle. Two additional sites (Comatillo and Sierra
de Agua) are located at Bayaguana in the lowlands
near Cotui (Poinar, 1992: table 4). The mines consist
primarily of tunnels leading deep into the sides of
mountains and occasionally they form deep, vertical
pits dug into the ground.
2.6 The author descending into the La Toca amber mine
The extraction process
The amber mines at La Toca are particularly noted for
producing vertebrate fossils. They are little more than
hand-carved tunnels dug into the side of the rock,
that follow a vein of the amber-containing blue-grey
sediments (Figure 2.5). None of the mining process
is mechanized. The tunnels are excavated using
hammers and chisels, with illumination provided
by candlelight; far from the industrialized process
depicted in the movie Jurassic Park. The tunnels
are damp, slippery, unlit, extremely cramped and
may extend up to 200 m into the hillside. It is not
possible to stand in the tunnels, which have barely
any supporting structures built in (Figure 2.6). The 2.7 Miners holding a drainage pipe for pumping out water
trunk posts visible in Figures 2.5 and 2.6 are to assist following flooding of the mine
36
2.18 VHR-CT scans of a spider in Paris amber (family Micropholcommatidae) from Penney et al. (2007). (a) dorsal
view; (b) ventral view; (c) anterior view; (d) posterior view; (e) lateral view; (f) lateral sectioned view; (g) right pedipalp
dorsal view; (h) right pedipalp posterior view; (i) left pedipalp retrolateral view; (j) right pedipalp prolateral view (body
length approx. 1 mm)
48
Key to
Hispaniolan
spider families
The spider family keys presented here are modified primarily from Jocqué & Dippenaar-Schoeman (2006)
and should permit the identification of both the fossil and extant spider fauna recorded from Hispaniola to
date. Some families are currently known either from the fossil or the extant fauna alone, which is not to say
that they will not be discovered from the other fauna at a later date. Jocqué & Dippenaar-Schoeman (2006)
also provided an identification key based on spider web structure; not particularly useful for amber-preserved
spiders, but a fabulous aid to identifying living spiders in the field.
Morphology and terminology
The basic body plan of spiders is shown in Figure (4.1), with more specific features illustrated throughout
the key and in the family description pages. They have two major tagma (body parts) joined by a narrow
pedicel through which the circulation, nervous and feeding systems are canalized: the anterior prosoma and
the posterior opisthosoma. The prosoma is covered dorsally by the carapace, which is divided into two regions:
the anterior cephalic region (where the eyes are situated) and the posterior thoracic region (often containing
a fovea—a distinct depression which serves as an internal point for muscle attachment). In some species the
division of these regions is distinct, demarcated by a cervical groove, whereas in others it is not noticeable;
both situations may occur in a single family. The shape and surface texture of the carapace (including pitting
and arrangement of setae, etc.) can be important taxonomic features. For example, the spider family Scytodidae
(spitting spiders) are recognizable immediately by their high, convex carapace that has evolved to contain the
enlarged posterior lobe of the venom gland, which occupies much of the prosoma.
Most spiders have eight eyes in two rows, although some families have fewer, or even no eyes at all.
The basic arrangement consists of two rows of four eyes each. These eye rows may be straight, recurved or
49
DORSAL
VIEW
VENTRAL
VIEW
pedipalp
chelicera
fang
pedipalp
chelicera
carapace
fovea
pedicel
sigilla
cardiac mark
abdomen
spinnerets
procurved eye row
MOQ: median ocular
quadrangle
coxa
trochanter
femur
book lung
maxilla
labium
sternum
patella
epigyne
spiracle
tibia
metatarsus
tarsus
tarsal claws
recurved eye row
AME: anterior median eye
ALE: anterior lateral eye
PLE: posterior lateral eye
PME: posterior median eye
cheliceral stridulatory
file
cribellum
trichobothria
spinnerets &
anal tubercle
spine
tarsal scopula
simple haplogyne pedipalp
4.1 General spider morphology
tarsal claws
with claw tuft
complicated entelegyne
pedipalp
epigyne
calamistrum
on leg 4
metatarsus
53
– Vertical orb-web decorated with stabilimenta: Araneidae
– Horizontal orb-web of ecribellate silk: Tetragnathidae
– Horizontal orb-web of cribellate silk: Uloboridae
Leg morphology:
– Tarsi and metatarsi of first legs with distinctive prolateral scopula (4.3): Palpimanidae
– Front leg with distinctive tibial and metatarsal clasping spurs (4.4): Mysmenidae
– Femur of first pair of legs with sclerotized spot ventro-subdistally (4.5): Mysmenidae
– Third pair of legs directed forwards rather than backwards: Segestriidae
– Legs very long and slender, with few or no spines: Pholcidae, Ochyroceratidae (some Theridiidae and
Scytodidae)
– Tarsi of fourth pair of legs with comb of serrated setae (4.6): Theridiidae, Nesticidae
– Tibiae and metatarsi of first two pairs of legs with distinct prolateral spination consisting of a series of short
spines, interspersed with a series of longer, slightly curved spines (4.7): Mimetidae
– Legs without spines, first femur enlarged, tibia, metatarsus and tarsus of first two pairs of legs with ventral
cusps (4.8): Trachelas (Corinnidae)
– Tarsi distinctly longer than metatarsi: Anapidae
– Legs with many, extremely long prominent spines: Oxyopidae
– Tiny spiders with femur of fourth pair of legs distinctly thicker than the other femora: Orchestina
(Oonopidae)
– Apex of metatarsi with soft, trilobate membrane (very difficult to see) (4.9): Sparassidae
– Claw tufts consisting of several rows of lamelliform setae (4.10): Anyphaenidae
– First pair of legs long and directed forward, femora with long trichobothria; metatarsus four with a calamistrum
set in a depression: Uloboridae
– Tibia of third pair of legs only, with double row of trichobothria: Mangora (Araneidae)
– All tibiae with many long (2–4× leg diameter) trichobothria: Theridiosomatidae
– Legs laterigrade with anterior two pairs distinctly thicker and longer than posterior two (4.11):
Thomisidae
– Metatarsus of second leg with a retrolateral excavation in the distal half, bearing a single, prominent
retrolateral spine (4.12): Misionella (Filistatidae)
– Metatarsi with flexuous zones (very difficult to see): Hersiliidae
– Metatarsus of legs three and/or four with a preening brush (4.13): Gnaphosidae
– Tarsi two-clawed, with prolateral claw distinctly more pectinate: Selenopidae
Prosoma (excluding leg) morphology:
– Anterior median eyes very large (4.14): Salticidae
– Posterior median eyes very large (4.15): Deinopidae
– Posterior median eyes of irregular shape (4.16, 4.17): Gnaphosidae, Prodidomidae
– Spider with only two eyes: Nops (Caponiidae)
– Spider with only four eyes: Tetrablemmidae, some Uloboridae
– Eight eyes in two rows: anterior with six, posterior with two (4.18): Selenopidae
– Lateral eyes on tubercles: Thomisidae
– Eight eyes in a hexagonal arrangement, anterior medians smallest (4.19): Oxyopidae
– Cephalic region of carapace ‘the head’ strangely modified: Linyphiidae (Erigoninae) and some Theridiidae
(e.g., Faiditus)
– Thoracic region of carapace distinctly domed (six eyes) (4.20): Scytodidae
– Clypeus forming a conical projection terminating in a sphere, both fringed with setae: Floricomus
(Linyphiidae)
– Chelicerae large with distinct projections (4.21): Tetragnathidae, some Corinnidae (e.g., Megalostrata)
55
– Sternum with pit organs on promargin (4.22): Theridiosomatidae
– Male palpal tibia widened distally and rim with a single row of long setae (4.23): Theridiidae
Opisthosoma morphology:
– Tiny spiders with conspicuous large, hairy anal tubercle (4.24): Oecobiidae
– Abdomen with large spiny outgrowths: some Araneidae (e.g., Gasteracantha, Micrathena)
– Tracheal spiracle broad, positioned well forward of the spinnerets: Anyphaenidae
– Abdomen extending beyond the spinnerets: Filistatidae, some Theridiidae and Pholcidae
– Abdomen flat and wedge-shaped, wider behind (long spinnerets) (4.25): Hersiliidae
– Anterior spinnerets as long as abdomen, long distal segment with spigots all along its median side (4.25):
Hersiliidae
– Anterior spinnerets as long as abdomen and widely separated, distal segment without spigots along its
median side (4.26): Dipluridae
– Anterior lateral spinnerets advanced on ventral side of abdomen and widely separated, bearing elongated
piriform gland spigots (4.27): Zimiris (Prodidomidae)
– Spinnerets tubular in appearance: Gnaphosidae
Dichotomous key to all Hispaniolan spider families
1. – Fangs close parallel to longitudinal axis of the body (paraxial), two pairs of booklungs (4.28): Section
1: Mygalomorphae
– Fangs close at 90o to longitudinal axis of the body (diaxial), one pair of booklungs (or maybe absent)
(4.29): Araneomorphae 2
2. – Calamistrum (metatarsus of fourth leg) and cribellum present (sometimes absent in males) (4.30): Section
2: Cribellate spiders
– Calamistrum and cribellum absent: Ecribellate spiders 3
3. – Less than eight eyes: Section 3
– Eight eyes: 4
4. – Tarsi with two claws: Section 4: Dionycha
– Tarsi with three claws: Section 5: Trionycha
Sometimes the third (unpaired) claw can be difficult to see in fossils
Section 1: Mygalomorphae
1. – Tarsi with two claws: 2
– Tarsi with three claws: 3
2. – Leg tarsi with clavate trichobothria along their length (4.31); apical segment of posterior spinnerets
long and finger-like; anterior lobe of endites well developed (4.32); clypeus often wide: Theraphosidae
– Leg tarsi with no, or only 4–6 clavate trichobothria basally; apical segment of posterior spinnerets short
and dome-shaped; anterior lobe of endites not well developed: Barychelidae
3. – Body covered with blunt-tipped clavate setae; booklung openings small and oval (4.33):
Microstigmatidae
– Body without blunt-tipped or clavate setae; booklung openings slit-like: 4
62
4.78
4.79
4.88
4.89
4.80
4.90
4.91
4.81
4.92
Araneid palp
4.82
4.93
4.94
4.83
?
4.95
4.84
4.96
4.85
4.97
4.87
4.86
4.98
4.99
65
Family
descriptions
The primary aim of this chapter is to permit the identification of Dominican amber fossil (and extant Hispaniolan)
spiders to family level. In some cases it may also facilitate identification to genus or even species. I have tried
to avoid the use of overly spider-specific technical terms wherever possible, because I want general amber
collectors to be able to use it as easily as more experienced arachnologists. For this reason, I have concentrated
only on the morphological features important for identification and have not provided extensive, detailed
accounts of all parts of spider anatomy for each family. For those interested, this information can be found in
Jocqué & Dippenaar-Schoeman (2006). Also, in many instances the identification information is specific to the
Hispaniolan taxa, rather than to the family as a whole. The relevant publications include primarily those that
record taxa from Hispaniola for the first time, rather than global or continental revisionary studies of particular
groups. Such works can be located easily using Platnick’s online World Spider Catalogue. Similarly, the
additional notes relate primarily to the Hispaniolan fauna and again I have tried to avoid too much information
regarding complicated systematic and phylogentic studies, but in places have included interesting snippets of
such.
66
INFRAORDER MYGALOMORPHAE
FAMILY DIPLURIDAE Simon, 1889
Funnel web mygalomorphs
Dominican amber, extinct taxa
Masteria sexoculata (Wunderlich, 1988); Ischnothele? sp.; Masteria sp.
Hispaniola, extant taxa
Ischnothele jeremie Coyle, 1995; Ischnothele garcia Coyle, 1995
Identification
The Hispaniolan diplurids are small for mygalomorph spiders, having a carapace length of approximately
5 mm in the extant species and only 1.5 mm in the fossil species. Chelicerae porrect, long fangs, furrows with
distinct teeth. Eight eyes (six in the fossil species) in a compact group; ocular area twice as wide as long
(Figure 5.1). Four spinnerets widely spaced; posterior pair very long (Figure 5.2). Tarsi with three claws,
scopulae absent. Male pedipalps as in Figures 5.3 and 5.4.
5.3 Masteria sexoculata
5.4 Ischnothele jeremie
5.1 Extant Ischnothele
carapace
5.2 Extant Ischnothele in lateral view
5.5 Masteria sp. in amber
Natural history
These spiders spin funnel webs that consist of two functionally distinct parts: a tubular retreat hidden
in an existing crevice, such as between rocks or the roots of a tree, and an exposed fan-like capture web.
Additionally, the webs often serve as homes to kleptoparasitic spiders (see Mysmenidae).
Relevant publications
Schawaller (1982a), Wunderlich (1988, 2004), Coyle (1995), Raven (1985, 2000)
Additional notes
The strictly fossil genus Microsteria Wunderlich, 1988 was synonymized with the extant genus
Masteria L. Koch, 1873 by Raven (2000). Raven examined the holotype of M. sexoculata Wunderlich, 1988
and was unable to determine the characters proposed by Wunderlich (1988) as diagnostic for the genus, thus
he synonymized the genus with Masteria (Raven, 2000). Wunderlich (2004: 619) doubted Raven’s synonymy
and provided additional characters in validation of Microsteria. Given Raven’s expertise with mygalomorph
spiders the synonymy was retained by Penney (2006b). The specimen described as Ischnothele? sp. (Figure
5.5) by Schawaller (1982a) was transferred to the genus Masteria by Raven (1985: 161). Wunderlich (1988)
also described a specimen as Ischnothele? sp. However, according to Coyle (1995: 29) the carapace and palpal
92
5.69 Male Faiditus crassipatellaris; note the raised ocular
region and the abdomen extending beyond the spinnerets
5.67 Photograph and drawing of male Styposis pholcoides
5.70 Male Dipoena altioculata; note the raised ocular
region with an extremely high clypeus (DG)
5.68 Male Lasaeola vicinoides in lateral (top) and dorsal 5.71 Female Spintharus longisoma; note the kite-shaped
(bottom) views; note the cylindrical carapace (DG)
abdomen with two distinct humps (KL)
101
FAMILY TETRAGNATHIDAE Menge, 1866
Four-jawed spiders
Dominican amber, extinct taxa
Azilia hispaniolensis Wunderlich, 1988; Cyrtognatha weitschati Wunderlich, 1988; Homalometa
fossilis Wunderlich, 1988; Leucauge sp.; Tetragnatha pristina Schawaller, 1982
Hispaniola, extant taxa
Agriognatha argyra Bryant, 1945; A. espanola Bryant, 1945; A. rucilla Bryant, 1945; Antillognatha
lucida Bryant, 1945; Azilia montana Bryant, 1940? [specimen was a juvenile female]; Chrysometa bigibbosa
(Keyserling, 1864); C. conspersa (Bryant, 1945); C. cornuta (Bryant, 1945); C. maculata (Bryant, 1945);
C. obscura (Bryant, 1945); C. sabana Levi, 1986; Glenognatha mira Bryant, 1945; Hispanognatha guttata
Bryant, 1945; Leucauge argyra (Walckenaer, 1842); L. regnyi (Simon, 1897); L. venusta (Walckenaer, 1842);
L. venustella Strand, 1916; Metabus ebanoverde Álvarez-Padilla, 2007; Tetragnatha elongata Walckenaer,
1842; T. nitens (Audouin, 1826); T. orizaba (Banks, 1898); T. pallescens F.O.P.-Cambridge, 1903; T. tenuissima
O.P.-Cambridge, 1889
Identification
Small to large spiders with eight eyes in two rows. The basic body plan of this family is highly variable,
but quite distinct in some genera (e.g., Tetragnatha spp.; Figure 5.90). The carapace is longer than wide and
the chelicerae vary from short and stout to highly modified (especially in mature males), with rows of large
teeth and large projecting spurs (Figures 5.91 & 5.92). Legs often long and spinose, with three tarsal claws.
Leucauge has a distinctive double row of trichobothria in the proximal half of the prolateral surface of the
femur of the fourth leg (not to be confused with uloborids which also possess a calamistrum). The opisthosoma
also varies from elongate in Tetragnatha (Figure 5.90) to globular in Glenognatha. In Leucauge and Azilia
it extends caudally beyond the spinnerets, the latter sometimes has small dorsal humps (Figure 5.93). The
pedipalp of mature males is relatively simple, with a large paracymbium and the conductor and embolus coiled
distally (Figure 5.94); often cymbial processes are present.
5.94 Pedipalp of Tetragnatha pristina
5.90 Tetragnatha body plan (male)
5.92 Chelicerae of Cyrtognatha weitschati
5.91 Chelicerae of Tetragnatha pristina
5.93 Abdomen of Azilia
127
5.144 Lyssomanes viridis; note the eyes are in four rows
(WM)
5.142 Anterior view of Salticidae showing the enlarged
anterior median eyes
AME
ALE
PME
PLE
5.143 Dorsal view of Salticidae in Dominican amber
showing the typical eye arrangement: AME (anterior
median eyes), ALE (anterior lateral eyes), PME (posterior
median eyes), PLE (posterior lateral eyes)
5.145 Menemerus bivittatus on tree trunk
Lyssomanes pristinus, a
large and relatively
common species in
Dominican amber;
note the elongate
cymbium
Corythalia scissa, sometimes the
retrolateral tibial apophysis lies
flush with the cymbium and
can be difficult to see
5.146 Some of the variation in fossil salticid pedipalps
129
Aspects of
palaeoecology
& historical
biogeography
The life histories and behaviour of spiders are relatively well differentiated at the family level, and spider
assemblages can be indicative of particular climatic conditions. The principal of behavioural fixity in the fossil
record is well documented and states that the behaviour, ecology, and climatic preferences of fossil organisms
will be similar to those found in their present-day descendants at the generic and often family level (e.g.,
Boucot, 1989). The close similarity of the Dominican Republic amber spider fauna at supraspecific (i.e., genus
and family) level to the extant Neotropical spider fauna (see later and Penney, 1999, 2005d, 2007b; Penney &
Pérez-Gelabert, 2002) supports the idea that the climate of Hispaniola at the time of the Dominican Republic
amber resin formation was tropical. This chapter assesses how the fossil and extant Neotropical araneofaunas
can be used together to investigate various interesting questions relating to the palaeoecology and historical
biogeography of the region.
138
PALEOZOIC
MISS.
MESOZOIC
PENN.
PERMIAN
TRIASSIC
CENOZOIC
JURASSIC
CRETACEOUS
PALEOCRIBELLATAE
ARANEOMORPHAE
HAPLOGYNAE
Triassaraneus
+
Argyrarachne
ERESOIDEA
KEY
PALPIMANOIDEA
DIONYCHA
RTA CLADE
DIVIDED CRIBELLUM CLADE
CANOE TAPETUM CLADE
range extension based on phylogenetic relationships
period of Dominican Republic amber resin secretion
range based on published record of described specimen(s)
fossil spiders described from Dominican Republic amber
hypothesised relationships
ghost lineage based on phylogenetic relationship
possible ancestral relationship
Unplaced families include Chummidae, Cybaeidae, Cycloctenidae, Hahniidae, Homalonychidae,
Synaphridae, Ephalmatoridae (fossil) and Insecutoridae (fossil)
AMAUROBIOIDS
Lycosoidea indet.
DEINOPOIDEA
ORBICULARIAE
JURARANEIDAE
ARANEOIDEA
6.5 Evolutionary tree of spiders highlighting the fossil (red circles) and Recent (blue text) Hispaniolan faunas
RASTELLOIDINA
MYGALOMORPHAE
SYMPHYTOGNATHOIDS
1.6
THERAPHOSODINA
OPISTHOTHELAE
NEOGENE
23
ATYPOIDEA
ARANEAE
65
95
135
152
180
205
230
250
240
260
290
300
310
325
355
375
410
Attercopus
PALEOGENE
Age (Ma)
DEVONIAN
MESOTHELAE
ATYPIDAE
ANTRODIAETIDAE
MECICOBOTHRIIDAE
HEXATHELIDAE
DIPLURIDAE
NEMESIIDAE
MICROSTIGMATIDAE
BARYCHELIDAE
PARATROPIDIDAE
THERAPHOSIDAE
CYRTAUCHENIIDAE
IDIOPIDAE
CTENIZIDAE
ACTINOPODIDAE
MIGIDAE
HYPOCHILIDAE
GRADUNGULIDAE
AUSTROCHILIDAE
FILISTATIDAE
CAPONIIDAE
TETRABLEMMIDAE
SEGESTRIIDAE
DYSDERIDAE
ORSOLOBIDAE
OONOPIDAE
PHOLCIDAE
DIGUETIDAE
PLECTREURIDAE
OCHYROCERATIDAE
LEPTONETIDAE
TELEMIDAE
SCYTODIDAE
PERIEGOPIDAE
DRYMUSIDAE
SICARIIDAE
OECOBIIDAE
HERSILIIDAE
ERESIDAE
UNPLACED ENTELEGYNES*
MIMETIDAE
MALKARIDAE
LAGONOMEGOPIDAE
SPATIATORIDAE
HUTTONIIDAE
STENOCHILIDAE
PALPIMANIDAE
MICROPHOLCOMMATIDAE
HOLARCHAEIDAE
PARARCHAEIDAE
MECYSMAUCHENIIDAE
ARCHAEIDAE
NICODAMIDAE
TITANOECIDAE
PHYXELIDIDAE
DICTYNIDAE
ZODARIIDAE
CRYPTOTHELIDAE
SPARASSIDAE
ANYPHAENIDAE
CLUBIONIDAE
CORINNIDAE
ZORIDAE
LIOCRANIDAE?
PHILODROMIDAE
SALTICIDAE
SELENOPIDAE
THOMISIDAE
CITHAERONIDAE
AMMOXENIDAE
TROCHANTERIIDAE
GALLIENIELLIDAE
LAMPONIDAE
GNAPHOSIDAE
PRODIDOMIIDAE
NEOLANIDAE
STIPHIDIIDAE
AGELENIDAE
AMPHINECTIDAE
DESIDAE
AMAUROBIIDAE
TENGELLIDAE
ZOROCRATIDAE
PISAURIDAE
TRECHALEIDAE
LYCOSIDAE
PARATTIDAE
PSECHRIDAE
SENOCULIDAE
OXYOPIDAE
CTENIDAE
ZOROPSIDAE
MITURGIDAE
DEINOPIDAE
ULOBORIDAE
NEPHILIDAE
ARANEIDAE
TETRAGNATHIDAE
LINYPHIIDAE
PIMOIDAE
BALTSUCCINIDAE
CYATHOLIPIDAE
SYNOTAXIDAE
NESTICIDAE
THERIDIIDAE
PROTHERIDIIDAE
THERIDIOSOMATIDAE
MYSMENIDAE
ANAPIDAE
SYMPHYTOGNATHIDAE
with 39 shared families (Figure 6.5), equating to a similarity value of 75%. The similarity values for genera
(Figure 6.7) are considerably lower at 15%; 26 genera are currently considered strictly fossil. However, such
genera have, in the past, been subsequently discovered in the extant fauna (see later) or synonymized with
recent genera. For example, Raven (2000: 573) synonymized Microsteria Wunderlich, 1988 with Masteria L.
Koch, 1873 (Dipluridae), so this value is considered an upper limit for the taxa described to date. The species
similarity is 0% because all described Dominican amber fossil and sub-fossil spiders are currently considered
144
The poor quality of this image is a result of reducing the file size for
distribution by email. The published version is of much higher quality
6.8 Continental affiliations of fossil and extant Hispaniolan spiders. (a) Extant genera recroded from Dominican
Republic amber. (b) Genera recorded from the Recent Hispaniolan fauna. (c) Extant genera from the amber and Recent
Hispaniolan faunas combined. (d) Continental species distributions for extant genera recorded in Dominican Republic
amber. (e) Continental species distributions for genera recorded from the Recent Hispaniolan fauna. (f) Continental
species distributions for both faunas combined.
effects). Current geological and palaeontological evidence suggests that all on-island lineages forming the
extant fauna must be younger than Middle Eocene (<40 million years; Iturralde-Vinent & MacPhee, 1999).
So what can the above analyses of fossil and extant spider faunas reveal about the continental origins
of this relatively young Neotropical fauna? Firstly, based on the relative geographical distributions of shared
genera, the Caribbean fauna probably did not originate from North America. In fact, some of the shared spider
genera recorded from North America result from Caribbean originators, not vice versa (Reiskind, 2001).
The shared values for Central and South America are reasonably similar when the faunas are compared at
generic level, with slightly higher values for the former (Figure 6.8a–c). However, it must be remembered
that the Panamanian isthmus arose long after the Dominican amber was formed. The great biotic interchange
(GBI) (Stehli & Webb, 1985) following the establishment of this land connection between North and South
America masks the former distributions of taxa and accounts for the observed similarity of the Central and
South American faunas. Additional complications result from the fact that the North American fauna is better
known than either the Central, South or West Indian faunas. Indeed, the low percentage similarity between
the fossil amber fauna and the on-island fauna would tend to suggest the latter is poorly known, a fact recently
demonstrated by Penney (2004b) and Hormiga et al. (2007). Within the last two years alone, five spider families
have been recorded from the extant Hispaniolan fauna for the first time: Prodidomidae (Platnick & Penney,
2004), Hersiliidae (Rheims & Brescovit, 2004) and Ochyroceratidae, Mysmenidae and Symphytognathidae
(Hormiga et al., 2007). Further similarities presumably result from more recent anthropogenic introductions
and through emigration and immigration of highly dispersive taxa to and from all neighbouring regions,
masking the earlier achievements of taxa with low dispersal capabilities (see later).
Thus, although we see patterns emerging at generic level, we need higher resolution in order to
generate a clearer picture. The comparative analyses of the continental species distributions for genera shared
with both the fossil and extant Hispaniolan faunas clearly indicate greatest affinities with South America
165
Biodiversity Research. Nova Science Publishers Inc., New York.
Penney, D. & Drew, B. 1996. A novel technique for extracting Segestria spp. from their retreats. Newsl. Br.
Arachnol. Soc., 76: 2–3.
Penney, D. & Langan, A.M.L. 2006. Comparing amber fossils across the Cenozoic. Biol. Letts., 2: 266–270.
Penney, D. & Ortuño, V.M. 2006. Oldest true orb-weaving spider (Araneae: Araneidae). Biol. Letts., 2: 447–
450.
Penney, D. & Pérez-Gelabert, D.E. 2002. Comparison of the Recent and Miocene Hispaniolan spider faunas.
Rev. Ibér. Aracnol., 6: 203–223.
Penney, D. & Selden, P.A. 2006. Assembling the Tree of Life—Phylogeny of Spiders: a review of the strictly
fossil spider families. Acta Zool. Bulgarica, supplement 1: 25–39.
Penney, D. & Selden, P.A. 2007. Fossils explained: Spinning with the dinosaurs: the fossil record of spiders.
Geol. Today, 23: 231–237.
Penney, D., Wheater, C.P. & Selden, P.A. 2003. Resistance of spiders to Cretaceous–Tertiary extinction events.
Evolution, 57: 2599–2607.
Penney, D., Dierick, M., Cnudde, V., Masschaele, B., Vlassenbroeck, J., Van Hoorebeke, L. & Jacobs, P. 2007.
First fossil Micropholcommatidae (Araneae), imaged in Eocene Paris amber using x-ray computed
tomography. Zootaxa, 1623: 47–53.
Pérez-Asso, A.R. & Pérez-Gelabert, D.E. 2001. Checklist of the millipedes (Diplopoda) of Hispaniola. Bol.
S.E.A., 28: 67–80.
Pérez-Gelabert, D.E. 1999. Catálogo sistemático y bibliografía de la biota fósil en ámbar de al República
Dominicana. Hispaniolana (N. Ser.), 1: 1–65.
Pérez-Gelabert, D.E. 2001. Preliminary checklist of the Orthoptera (Saltatoria) of Hispaniola. J. Orthoptera
Res., 10: 63–74.
Pérez-Gelabert, D.E. 2008. Arthropods of Hispaniola (Dominican Republic and Haiti): a checklist and
bibliography. Zootaxa, in press.
Pérez-Gelabert, D.E. & Flint, O.S. Jr. 2000. Annotated list of the Neuroptera of Hispaniola, with new faunistic
records of some species. J. Neuropterol., 3: 9–23.
Perkovsky, E.E., Zosimovich, V.Yu & Vlaskin, A.Yu 2003. Rovno amber insects: first results of analysis.
Russian Entomol. J., 12: 119–126.
Perrichot, V. 2004. Early Cretaceous amber from south-western France: insight into the Mesozoic litter fauna.
Geol. Acta, 2: 9–22.
Perrichot, V. 2005. Environnements paraliques à amber et à végétaux du Crétacé Nord-Aquitain (Charentes,
Sud-Ouest de la France. Mém. Géosci. Rennes, 118: 1–213.
Petrunkevitch, A. 1911. A synonymic index-catalogue of spiders of North, Central and South America with all
adjacent islands, Greenland, Bermuda, West Indies, Terra del Fuego, Galapagos, etc. Bull. Amer. Mus.
Nat. Hist., 29:1–791.
Petrunkevitch, A. 1928 The Antillean spider fauna—a study in geographic isolation. Science, 68: 650.
Petrunkevitch, A. 1942. A study of amber spiders. Trans. Connect. Acad. Arts Sci., 34: 119–464. Pls I–LXIX.
Petrunkevitch, A. 1963. Chiapas amber spiders. Uni. Calif. Publ. Entomol., 31: 1–40.
Petrunkevitch, A. 1971. Chiapas amber spiders, 2. Uni. Calif. Publ. Entomol., 63: 1–44.
Piel, W.H. 2001. The systematics of neotropical orb-weaving spiders in the genus Metepeira (Araneae,
Araneidae). Bull. Mus. Comp. Zool., 157: 1–92.
Pike, E.M. 1993. Amber taphonomy and collecting biases. Palaios, 8: 411–419.
Pike, E.M. 1994. Historical changes in insect community structure as indicated by hexapods of Upper
Cretaceous Alberta (Grassy Lake) amber. Can. Entomol., 126: 695–702.
Platnick, N.I. 2008. The world spider catalogue (version 8.5). Online at, http://research.amnh.org/entomology/
spiders/catalog/INTRO2.html.
Platnick, N.I. & Nelson, G. 1978. A method of analysis for historical biogeography. Syst. Zool., 27: 1–16.
Platnick, N.I. & Penney, D. 2004. A revision of the widespread spider genus Zimiris (Araneae, Prodidomidae).
174
physical properties of fluids 133
Pisauridae 41, 45, 63, 106, 107, 138, 140, 141
Plio-Pleistocene cooling/glaciations 142
Prodidomidae 41, 46, 51, 53, 55, 61, 120, 121, 138, 140, 141, 144, 150
Pseudochiridiidae 155
pseudoscorpions 11, 13, 14, 152–154
redeposition of amber 26
ricinuleids 13, 14
Russian amber 16, 38
Salticidae 19, 23, 40, 41, 47, 50, 53, 59, 106, 108, 125, 127, 128, 131, 134, 135, 137, 138–141, 145
Samoidae 154
schizomids 13, 14, 152
scorpions 13, 14, 153, 155
Scutoverticidae 154
Scytodidae 42, 48, 50, 53, 59, 72, 73, 138, 140, 141, 150
Segestriidae 43, 53, 59, 78, 79, 138, 140, 141
Selenopidae 46, 53, 59, 122, 137, 138, 140, 141
sesquiterpenes 26, 27
Sicariidae 42, 50, 59, 71–73, 138, 140, 141
size of spider inclusions 134–136, 139
Smarididae 154
solifuges 13, 14, 153, 155
solubility of amber 26, 34
South American amber 19, 39
Spanish amber 16, 19, 34
Sparassidae 47, 53, 59, 63, 120, 122, 124, 138, 140, 141, 145
spider family richness 137–139
spider–insect co-radiations 19
Sternophoridae 155
Symphytognathidae 41, 44, 57, 94, 95, 138, 140, 141, 144, 150
synanthropic spiders 14, 75, 76, 84, 93, 94, 120, 122, 149
syninclusions 11, 131
taphonomy 11, 90, 130, 131, 133, 137, 139
terpenoid compounds 10, 26, 27
Tetrablemmidae 43, 50, 51, 53, 57, 78, 134, 138, 140, 141, 145, 150
Tetragnathidae 40, 41, 44, 53, 64, 100, 101, 134, 138–141, 145, 146
Theraphosidae 15, 42, 52, 55, 69, 138, 140, 141, 146
Theridiidae 15, 41, 43, 50, 51, 53, 55, 57, 64, 82, 89, 90, 94, 98, 100, 134, 135, 138–141, 150
Theridiosomatidae 40, 44, 51, 53, 55, 64, 94, 134, 138, 140, 141
Thomisidae 47, 53, 53, 59, 124, 125, 138, 140, 141
Triassic spiders 17
trigonotarbids 13, 14
Trochanteriidae 46, 61, 118, 119, 120, 138, 140, 141
Trypochthonidae 154
Ukranian amber 16, 42
Uloboridae 43, 51, 53, 57, 87, 134, 138, 140, 141, 150
uropygids 13, 14, 152
vicariance 25, 142, 145, 146, 149
viscosity (of resin) 27, 132, 133, 137