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209
2003. The Journal of Arachnology 31:209–245
ON THE USE OF AMPULLATE GLAND SILKS BY WOLF
SPIDERS (ARANEAE, LYCOSIDAE) FOR ATTACHING THE
EGG SAC TO THE SPINNERETS AND A PROPOSAL FOR
DEFINING NUBBINS AND TARTIPORES
Mark A. Townley and Edward K. Tillinghast: Department of Zoology, Rudman
Hall, University of New Hampshire, Durham, New Hampshire 03824 USA. E-mail:
mtownley@cisunix.unh.edu
ABSTRACT. The means by which female wolf spiders attach an egg sac to their spinnerets was inves-
tigated using scanning electron microscopy. In four Pardosa species, we observed that silk fibers emerging
from ampullate gland spigots had been affixed to the surface of the egg sac. More specifically, primary
(1
8
) and secondary (2
8
) major ampullate (MaA) glands and 1
8
and 2
8
minor ampullate (MiA) glands all
contributed fibers for this purpose. The diameters of the 2
8
MaA and 2
8
MiA fibers were greater than
those of the 1
8
MaA and 1
8
MiA fibers and, correspondingly, the widths of the 2
8
ampullate spigots were
clearly greater than those of the 1
8
ampullate spigots. Larger 2
8
ampullate spigots were also observed in
adult females of species from three other lycosid genera. Thus, 2
8
ampullate glands, which in araneoids
function only in juveniles during proecdysis, are not only functional in adult female lycosids (and adult
females of several other families), but they appear to play a greater role than the 1
8
ampullate glands in
egg sac attachment. Observations made on the 1
8
and 2
8
ampullate spigots of adult females from species
belonging to several other families are also presented. Cuticular structures referred to as nubbins and
tartipores are present in some spinning fields on spinnerets. A proposal is made for defining these terms
by a criterion, namely their different origins, which differs from that applied previously.
Keywords: Ampullate silk gland, Pardosa,Hogna,Trochosa, Lycosoidea
With some exceptions, those spiders that
typically carry their egg sacs using only their
spinnerets belong to one of the following three
lycosoid taxa: the family Lycosidae, the fam-
ily Trechaleidae, or the subfamily Rhoicininae
(with Shinobius considered a member of the
latter taxon, Yaginuma 1991; Sierwald 1993).
The familial placement of Rhoicininae is un-
certain (Sierwald 1993; Carico 1993), but its
members are currently listed in Trechaleidae
(see Platnick 2002). The exceptions referred
to include other lycosoids (sensu Griswold et
al. 1999) (e.g., the ctenid Cupiennius, Barth
et al. 1991; Silva Davila in press) as well as
non-lycosoids (e.g., the nesticids Nesticus,
Eidmannella, Nielsen 1932:201; Bristowe
1958:223; Po¨tzsch 1963:30; the zorids Vor-
aptus,Neoctenus, Lawrence 1964:34; Silva
Davila in press, though in the latter paper
Neoctenus is transferred to Trechaleidae). The
spinnerets are also involved in carrying the
egg sac in at least some genera within the ly-
cosoid family Pisauridae (see Discussion), but
here the chelicerae play the principal role in
securing the egg sac, which is positioned be-
low the sternum. At times, lycosids hold the
egg sac in a similar attitude; e.g., during the
last phase of egg sac construction, when as-
sisting in spiderling emergence, or sometimes
when fleeing, after one has attempted to take
the egg sac away from the mother (e.g., Mont-
gomery 1903; Le´caillon 1905). And, con-
versely, some pisaurids occasionally ‘‘drag the
sac from the spinnerets alone in the same
manner as a lycosid’’ (Bishop 1924:28).
As described in Carico (1993), Sierwald
(1990a, 1993), Scheffer (1905) and literature
cited by the first two authors, differences exist
among lycosids, trechaleids, rhoicinines, and
pisaurids with regard to egg sac structure and
the maternal care afforded post-emergent spi-
derlings. Typically, the egg sacs of lycosids
and rhoicinines are spherical or lenticular,
those of pisaurids are spherical, while those of
trechaleids are hemispherical. A seam joining
the upper and lower valves of the egg sac is
apparent among lycosids, trechaleids, and
some rhoicinines (Shinobius), but not in some
210 THE JOURNAL OF ARACHNOLOGY
other rhoicinines (Rhoicinus) or pisaurids, and
only in trechaleids is a ‘skirt’ (Carico 1993)
produced at the seam.
Lycosid, trechaleid, and rhoicinine females
continue to carry their progeny for a number
of days after they have emerged from the egg
sac (lycosids about 2–14 days, e.g., McCook
1884; Montgomery 1903:72,76,82,90; Engel-
hardt 1964:303,387; Trechalea about 17–19
days, Carico et al. 1985; Shinobius about 4
days, Kaihotsu 1988). But while lycosid spi-
derlings climb onto the mother’s abdomen
during this period, trechaleid spiderlings and
at least Shinobius spiderlings (among rhoici-
nines) climb onto the outside of the egg sac,
which the female continues to carry. However,
Yaginuma (1991), specifying two Arctosa and
one Hygrolycosa species, reports that trans-
port on the egg sac, rather than on the moth-
er’s abdomen, also occurs in some lycosids. In
those instances in which trechaleid spiderlings
have been observed on the mother’s abdomen,
this appears to be due to crowding on the egg
sac with consequent spill-over (Carico 1993),
just as lycosid spiderlings may spill over onto
their mother’s cephalothorax.
Among pisaurids (where known), the egg
sac is carried by the female until shortly be-
fore the young emerge, or at latest when the
first spiderlings begin to emerge (Gertsch
1979:197). Typically, the egg sac is then se-
cured within a nursery web. The mother
builds the network of silk fibers that constitute
the nursery web before and/or after the end of
the egg-sac-carrying period, often on vegeta-
tion, with the spiderlings adding fibers after
their emergence. The post-emergent spider-
lings remain within the nursery web, guarded
by the mother (though see Montgomery 1909:
556; Forster 1967:84), for a period of time
that again varies considerably, sometimes
even within a species (e.g., Dolomedes fim-
briatus (Clerck 1757) spiderlings may remain
in the nursery web from 3 or 4 days (Bristowe
1958:191) to about 3 weeks (Nielsen 1932:
134)). After this period the spiderlings dis-
perse. Kaihotsu (1988) reports that Shinobius
females, after carrying their young on the out-
side of the egg sac for a few days, then hang
the egg sac with spiderlings in a nursery web,
where the spiderlings remain for about one
day.
This study is concerned with the specific
silk glands used by lycosids for attaching the
egg sac to the spinnerets. The impetus for the
study can be explained as follows. Observa-
tions made on spinnerets by a number of
workers indicated that a certain category of
ampullate silk glands, what we call secondary
(2
8
) ampullate silk glands, are functional in
juvenile spiders of most, if not all, entelegyne
taxa. However, in only some entelegynes are
2
8
ampullate glands also functional in adults
(sometimes only in the females, as in lycos-
ids). The only role assigned to these silk
glands that we were aware of when we began
this study is to produce silk fibers during one
specific period in the molt-intermolt cycle of
juveniles (detailed below). The question thus
arose, what are 2
8
ampullate silk glands used
for when they are retained in adults? It oc-
curred to us that if these silk glands are in-
volved in egg sac attachment in lycosoids, it
might be a situation in which we could, in
effect, catch spiders in the act of drawing fi-
bers from these glands and, thus, demonstrate
their use in at least one specific application in
certain adult spiders. At that time we were not
aware that Carico (1993) had already ob-
served certain 2
8
ampullate gland fibers being
used for egg sac attachment to the spinnerets
in trechaleids. As we describe below, trechal-
eids and the lycosids that we have examined
(primarily Pardosa) show similarities and dif-
ferences with respect to egg sac attachment,
including in their use of 2
8
ampullate silks. In
partial answer to the above question, all we
know at this time is that adult females of at
least some lycosid and trechaleid (Carico
1993) genera use silk from 2
8
ampullate
glands to help secure the egg sac to the spin-
nerets.
To provide a better overall perspective on
2
8
ampullate silk glands and their roles, the
following section reviews different categories
of ampullate silk glands. It is followed by four
sections dealing with nubbins and tartipores,
protuberances present in some spinning fields.
These sections are included because our in-
terpretations of spinneret micrographs ob-
tained during this study rely on and make ref-
erence to these protuberances. And because
our basis for distinguishing nubbins from tar-
tipores differs from that of earlier authors, and
also differs from our own earlier views, the
first of these four sections explains how we
arrived at our current definitions for nubbins
and tartipores. The last three sections review
211TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
some aspects of the occurrence of these pro-
tuberances. We hope this overview will be
useful in light of the growing importance of
these structures in phylogenetic studies. In ad-
dition to presenting observations made on the
spinnerets and egg sacs of some female ly-
cosids, this paper contains comparative obser-
vations made on the spinnerets of male lycos-
ids and the spinnerets of spiders from several
other families in which adult females retain
apparently functional 2
8
ampullate glands.
Categories of ampullate silk glands and
their roles.—The ampullate silk glands of
spiders within the Orbiculariae and some oth-
er taxa (e.g., Hersiliidae, Kovoor 1984; Se-
gestriidae, Clubionidae, Gnaphosidae, Thom-
isidae, Kovoor 1987; Oxyopidae, Kovoor &
Mun˜oz-Cuevas 1998) can be divided, on the
basis of histochemical differences, into major
ampullate silk (MaA) glands and minor am-
pullate silk (MiA) glands (reviewed in Kovoor
1977, 1987; also Kovoor & Peters 1988). The
ducts of the MaA glands connect to spigots
located on the anterior lateral spinnerets
(ALS) while MiA gland ducts connect to spig-
ots on the posterior median spinnerets (PMS).
In some other spiders, however, including the
Lycosidae, histochemical differences are not
readily apparent between those ampullate
glands with ducts that empty on the ALS ver-
sus those with ducts emptying on the PMS
(Kovoor 1976, 1987). One may therefore
question the validity of recognizing two dif-
ferent types of ampullate glands in such taxa.
Nevertheless, for clarity and in keeping with
precedents (e.g., Platnick et al. 1991:2), any
ampullate glands with ducts attached to the
ALS will be called MaA glands and any with
ducts attached to the PMS will be called MiA
glands (Table 1).
In more basal araneoids, including those in
the families Araneidae and Tetragnathidae
(Griswold et al. 1998), both the MaA and
MiA glands can be further subdivided into a
single pair of primary (1
8
) MaA/MiA glands
and two pairs of 2
8
MaA/MiA glands (Table
1) (Townley et al. 1993; Tillinghast & Town-
ley 1994). Observations made on spinnerets
(Coddington 1989; Forster et al. 1990; Peters
& Kovoor 1991; Hormiga 1994a,b, 2000;
Griswold et al. 1998) suggest a tendency for
the more derived araneoids to lack 2
8
MiA
glands. The 1
8
MaA and 1
8
MiA glands func-
tion in each juvenile stadium from immedi-
ately after ecdysis (even as the spider is hang-
ing by its ‘molting threads’) until about the
beginning of the following proecdysis (the
few days preceding ecdysis during which in-
ternal changes take place in preparation for
ecdysis), as well as throughout adulthood
(Townley et al. 1993). During proecdysis
these glands are remodeled, rendering them
temporarily nonfunctional (Townley et al.
1991). The task of producing ampullate silk
during each proecdysis is taken over by one
of the two pairs of 2
8
MaA glands and one of
the two pairs of 2
8
MiA glands. Each pair of
2
8
ampullate glands cycles through growth
and regression phases, reaching maximum
size and accumulation of luminal contents at
proecdyses in every other juvenile stadium,
with one pair of 2
8
MaA/2
8
MiA glands pro-
ducing silk during proecdysis in even-num-
bered stadia and the other pair functioning in
odd-numbered stadia (Townley et al. 1993).
Because only one of the two pairs of 2
8
MaA/
2
8
MiA glands produces silk in a given juve-
nile stadium (i.e., is ‘open’, see Table 1), there
is only one 2
8
MaA spigot on each ALS and
one 2
8
MiA spigot on each PMS of juveniles
(in addition to the single 1
8
MaA spigot and
single 1
8
MiA spigot on each ALS and PMS,
respectively). After the final molt, with no ad-
ditional proecdyses to pass through, both pairs
of 2
8
MaA/2
8
MiA glands degenerate (Seki-
guchi 1955b; Townley et al. 1991). Thus, 2
8
ampullate glands do not function in adults and
only nonfunctional vestiges, termed nubbins
(Coddington 1989; Yu & Coddington 1990),
of 2
8
ampullate spigots are present on adult
ALS and PMS (Sekiguchi 1955b; Peters 1955;
Mikulska 1966; Wilson 1969). This situation
exists in both sexes. External examinations of
spinnerets indicate that this description, out-
lined in Table 1, applies not only to basal ar-
aneoids, but to some non-araneoids (e.g., ox-
yopids, Kovoor & Mun˜oz-Cuevas 1998), and,
whatever their superfamilial placement, to
some mimetids (see data for Mimetus in Table
3; see also mimetid spinneret micrographs in
Platnick & Shadab 1993). Interestingly, it may
be that the above description applies to some
species within the mimetid genus Ero, but not
others (cf. figs. 29, 30 with figs. 41, 42 in
Platnick & Shadab 1993, noting especially the
presence of a MiA nubbin and MiA tartipore
(tartipore defined below) in fig. 42 and the
212 THE JOURNAL OF ARACHNOLOGY
Table 1.—The division of ampullate silk glands into different categories on the basis of which spinnerets receive their ducts (major vs. minor), whether
glands are functional in both juveniles and adults or just in juveniles during proecdysis (primary (1
8
) vs. secondary (2
8
)), and whether 2
8
ampullate glands
have an outlet to the extracorporeal environment in a given stadium or not (open vs. blocked). This scheme is based on observations made primarily on
Araneus (Townley et al. 1993; Townley 1993; Tillinghast & Townley 1994) and applies to both sexes. Lycosids and the species from the other families in
Table 2 deviate from this table only in that 2
8
MaA and 2
8
MiA glands of females apparently function not only in juveniles during proecdysis, but one pair
of each is functional in adults as well. The double-headed arrows between open and blocked 2
8
ampullate glands indicate that a given pair of glands is
alternately open and blocked; open throughout one stadium (i.e., from one ecdysis to the next), blocked throughout the following stadium, open again in the
stadium after that, and so on.
Ampullate Silk Glands
6 pairs
produce silk fibers used in a variety of applications including draglines and non-sticky structural elements of webs
Major Ampullate Silk (MaA) Glands Minor Ampullate Silk (MiA) Glands
3 pairs 3 pairs
ducts lead to anterior lateral spinnerets (ALS) ducts lead to posterior median spinnerets (PMS)
1
8
MaA Glands 2
8
MaA Glands 1
8
MiA Glands 2
8
MiA Glands
1 pair 2 pairs 1 pair 2 pairs
functional in juveniles
(from ecdysis to
start of next proec-
dysis) and adults
functional in juveniles only
(during proecdysis only) functional in juveniles
(from ecdysis to start of
next proecdysis) and
adults
functional in juveniles only
(during proecdysis only)
Open 2
8
^&
Blocked 2
8
Open 2
8
^&
Blocked 2
8
MaA Glands MaA Glands MiA Glands MiA Glands
1 pair 1 pair 1 pair 1 pair
ducts connect to spigots
that open to the out-
side environment
ducts do not connect to
spigots that open to
the outside environ-
ment
ducts connect to spigots
that open to the out-
side environment
ducts do not connect to
spigots that open to
the outside environ-
ment
213TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
absence of these protuberances in fig. 30; see
also Schu¨tt 2000:145).
In contrast to the situation just described for
basal araneoid taxa, examinations of spinner-
ets from spiders in certain amaurobioid (sensu
Griswold et al. 1999) and dionychan (sensu
Coddington & Levi 1991) families, including
the Lycosidae (Table 2), reveal a sexual di-
morphism wherein males appear to conform
to the above description, but females do not
(Fig. 1). Instead, adult females retain appar-
ently functional 2
8
MaA and 2
8
MiA spigots,
one pair of each, indicating that they use 2
8
,
as well as 1
8
, ampullate glands as adults. In
trechaleids, Carico (1993) has observed the
use of 1
8
and 2
8
MiA silks by adult females
for securing the egg sac to the spinnerets.
Here we report that adult female Pardosa use
1
8
and 2
8
MaA and 1
8
and 2
8
MiA silks to
attach their egg sacs to their spinnerets. The
2
8
ampullate fibers have greater diameters
than the 1
8
ampullate fibers, indicating a great-
er contribution from the 2
8
ampullate glands
to the support of the egg sac.
Terminology.—Nubbins and tartipores:
As mentioned, the term ‘nubbin’ has been ap-
plied to cuticular protuberances on adult ALS
and PMS that appear to be nonfunctional ves-
tiges of 2
8
MaA and 2
8
MiA spigots, respec-
tively. Other cuticular protuberances, scattered
within piriform and aciniform spinning fields
(Kovoor 1986; Platnick 1990; Yu & Codding-
ton 1990), as well as on the PMS and poste-
rior lateral spinnerets (PLS) of at least some
mygalomorphs (Glatz 1973; Palmer 1990),
have been called ‘tartipores’ (Shear et al.
1989; Yu & Coddington 1990). Originally, the
distinction between a nubbin and a tartipore
was based on whether the protuberance is a
morphological singular and can, therefore, be
uniquely designated (nubbin) or if it is one of
several, or many, such structures on a single
spinneret that are designated collectively (tar-
tipore) (Yu & Coddington 1990; see also Cod-
dington 1989:81). Both nubbins and tartipores
were tentatively interpreted to be vestigial
spigots. Additional observations, however, re-
vealed that the protuberances identified as tar-
tipores are not vestigial spigots. Instead, they
are the remains of collared openings that
formed in the cuticle when it was first being
laid down during proecdysis beneath the old
cuticle (Townley et al. 1993). The openings
allowed silk gland ducts to maintain their at-
tachments to spigots on the old cuticle during
proecdysis, despite the formation of the inter-
vening new cuticle (Fig. 1). Thus, while some
protuberances do seem to be vestigial spigots,
others have a very different origin, being rem-
nants of these openings.
After realizing that we were actually deal-
ing with two different categories of cuticular
protuberances, we made an ill-devised attempt
to both retain the original distinction between
nubbins and tartipores (singulars versus mul-
tiples) and distinguish vestigial spigots from
remnants of openings by use of the adjectives
‘vestigial-type’ and ‘non-vestigial-type’, re-
spectively (Townley et al. 1993). We soon
abandoned this approach in favor of another,
not previously published, that is concerned
only with the two different origins of the pro-
tuberances under consideration (for further ex-
planation see Townley 1993:7, 8). The latter
approach, which we will follow in this paper,
retains the terms nubbin and tartipore, but de-
fined as follows: Nubbin: a nonfunctional,
only partially formed, i.e. vestigial, spigot, ei-
ther morphologically singular or multiple.
Tartipore: a cuticular scar, morphologically
singular or multiple, that results, after ecdysis,
from a collared opening forming in the de-
veloping exoskeleton during proecdysis; the
opening accommodates a silk gland duct, al-
lowing the duct to remain attached to a spigot
on the old exoskeleton during proecdysis. By
these definitions the protuberances that were
initially called tartipores (those among piri-
form and aciniform spigots) are still called tar-
tipores (Fig. 1). However, only some of the
structures previously referred to as nubbins
are still called nubbins by our definition. For
example, as in earlier reports, we identify as
nubbins those nonfunctional protuberances in
some adults that occur where functional 2
8
MaA/2
8
MiA spigots would have formed if
the spider had instead molted to yet another
juvenile instar (see Figs. 1, 13, 15). But there
are other protuberances near ampullate spigots
in many adult and juvenile araneomorphs, pre-
viously called nubbins (e.g., Yu & Coddington
1990; Townley et al. 1991, 1993; Tillinghast
& Townley 1994), that we now identify as
ampullate tartipores, including the ‘‘second
nubbin’’ on the PMS of adult anapids and
some synotaxids (Griswold et al. 1998:41) and
the ‘‘second remnant’’ on the PMS of adult
oxyopids (Kovoor & Mun˜oz-Cuevas 1998:
214 THE JOURNAL OF ARACHNOLOGY
215TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
←
Figure 1.—Schematic diagram of the left anterior lateral spinneret (ALS) of a female lycosid during
proecdysis, shortly before the ecdysis that yields an adult. The upper ALS diagram represents the cuticle
of the penultimate instar which will be cast off at ecdysis. The ALS diagram below this represents the
underlying, newly-formed cuticle which will be part of the exoskeleton of the adult. Shown is the entire
distal segment (DS) of the ALS, to which the major ampullate (MaA) spigots (labeled as ‘1
8
’ and ‘2
8
’)
and piriform spigots (P) are restricted, atop the more distal portion of the ALS proximal segment (PS).To
aid orientation, a much less magnified depiction of the same part of the left ALS is enclosed by a box in
the ventral view of the spider, shown at top, in which the spinnerets are presented as if artificially spread.
By late proecdysis, the duct of the primary (1
8
) MaA gland, previously connected to a spigot on the old
cuticle (that labeled ‘1
8
’), has just been re-modeled (Townley et al. 1991, 1993) and is now connected to
a spigot on the new cuticle (again labeled ‘1
8
’) (silk gland ducts are indicated by dashed lines). Thus, the
1
8
MaA gland, nonfunctional during proecdysis, will again be functional immediately after ecdysis. Col-
lared openings (tartipore progenitors) form in the new cuticle to accommodate the ducts of any silk glands
that are to remain functional throughout proecdysis. The ducts of two such silk glands are shown connected
to spigots on the old cuticle. After ecdysis the collapsed forms of these openings (tartipores) will remain
evident in the new cuticle. A single MaA tartipore progenitor (MaATP) forms on each ALS of the new
cuticle to accommodate the duct of a secondary (2
8
) MaA gland. Multiple piriform tartipore progenitors
(PTP) also form on each ALS, one per piriform gland duct. (For clarity only a few piriform spigots are
shown, and of those on the old cuticle, the duct connected to only one is shown. In reality, more piriform
spigots are present and it appears that each piriform spigot on the old cuticle remains connected to a
functioning duct, thus requiring the formation of one PTP on the new cuticle for each piriform spigot on
the old cuticle.) The 2
8
MaA gland identified as ‘open’ will become ‘blocked’ at ecdysis (because its
outlet, the spigot, will be lost along with the rest of the old cuticle), and, conversely, that identified as
‘blocked’ will become ‘open’ (since the 2
8
MaA spigot it is connected to will be open to the outside
environment after ecdysis). The portion of the female new cuticle shown within a box differs from the
situation in males (depicted at lower right) because 2
8
ampullate spigots do not form in adult males. Only
ampullate nubbins (MaA nubbin (MaAN) on ALS, MiA nubbin on PMS) form in the positions occupied
by 2
8
ampullate spigots in adult females and, thus, all 2
8
ampullate glands are ‘blocked’ and nonfunctional
in adult males. Structures not drawn precisely to scale. BL, book lung; ALS, anterior lateral spinneret;
PMS, posterior median spinneret; PLS, posterior lateral spinneret; DS, distal segment of anterior lateral
spinneret; PS, proximal segment of anterior lateral spinneret; P, piriform gland spigot; PT, piriform tarti-
pore; PTP, piriform tartipore progenitor; 1
8
, primary major ampullate gland spigot; 2
8
, secondary major
ampullate gland spigot; MaAT, major ampullate tartipore; MaATP, major ampullate tartipore progenitor;
MaAN, major ampullate nubbin.
136). This is not the first time such protuber-
ances have been referred to as tartipores (Plat-
nick & Forster 1993:7, 9; Griswold et al.
1998:11), but in these earlier instances the dis-
tinction made between tartipores and nubbins
was not stated. The term tartipore was perhaps
applied solely because of the resemblance be-
tween ampullate tartipores and the more well
known tartipores in piriform and aciniform
spinning fields, rather than because of recog-
nition of what tartipores, as here defined, rep-
resent. As indicated above, ampullate tarti-
pores mark the sites where 2
8
ampullate gland
ducts passed through the cuticle during the
most recent proecdysis, enabling 2
8
ampullate
glands to function throughout proecdysis (Fig.
1). Note that Fig. 1 depicts only the ALS from
a lycosid and so only spigots, tartipores, and
ducts of MaA and piriform glands are shown.
Bear in mind that a comparable situation ex-
ists on the PMS with the spinning apparatus
of MiA and aciniform glands, respectively.
Ampullate gland nubbins versus ampul-
late gland tartipores.—When examining
spinnerets, care must be taken if one wishes
to determine whether ampullate nubbins and
tartipores are present or not, as well as distin-
guish the former category from the latter.
Viewing the spinnerets at various angles and
from different directions is sometimes re-
quired. In some adult araneoids, for example,
the MiA nubbin and MiA tartipore on a PMS
often occur side by side (e.g., Coddington
1989:fig 16; Platnick et al. 1991:fig. 271, low-
er black arrow, tartipore on left, nubbin right;
Townley et al. 1991:fig. 24; lower arrow to
tartipore, upper to nubbin; Hormiga et al.
1995:fig. 16C, nubbin left, tartipore right) and
can be interpreted as a single structure if
viewed at too low a magnification or from an
216 THE JOURNAL OF ARACHNOLOGY
inopportune angle, or if the MiA nubbin is
especially small. In describing the PMS of
adult males of two anapid species, Platnick et
al. (1991:60) noted the presence of ‘‘a large
posterior minor ampullate gland spigot ac-
companied by a vestigial remnant bearing a
short lobe on its medial side’’. The ‘‘vestigial
remnant’’ is a MiA tartipore, the ‘‘short lobe’’
is a MiA nubbin. Even with careful observa-
tion it can sometimes be difficult, especially
with certain species, to discern a given tarti-
pore or nubbin. We were puzzled for a time
by our inability to spot a MaA tartipore on the
ALS of juvenile and adult Araneus cavaticus
(Keyserling 1882) until it became clear that
this tartipore occurs at a site where, in this
species, the cuticle is typically compressed or
overhung by the piriform spinning field and
the tartipore is obscured (Townley et al.
1993). In contrast, single or multiple MaA tar-
tipores are often clearly visible in many other
araneomorphs as a number of published mi-
crographs attest (several were cited in Town-
ley et al. 1993:36 as ‘‘non-vestigial-type MaA
nubbins’’; other examples include Platnick et
al. 1991: fig. 16, multiples in Gradungula, fig.
39, a single in Thaida, fig. 277, a round single
in Pachygnatha next to smaller oblong MaA
nubbin; Harvey 1995: fig. 11, a single in Am-
bicodamus posterolateral to the 2
8
MaA spig-
ot; Davies 1998a:fig. 68, a single in Jalkabur-
ra lateral to the 2
8
MaA spigot; Platnick 1999:
fig. 3, a single in Liocranoides between and
lateral to 1
8
and 2
8
MaA spigots; Hormiga
2000:plate 42B, a single in Laminacauda pos-
terolateral to MaA nubbin, larger than the
multiple piriform tartipores). In this paper,sin-
gle MaA tartipores can be seen in Figs. 1, 8,
9, 12, 13, 16, 18, 22, 24, 26, 28, 30, 32, 34,
36, 40 & 41 and single MiA tartipores can be
seen in Figs. 10, 11, 14, 15, 19–21, 23, 25,
27, 29, 31, 33, 35, 37–39 & 42.
Given that the occurrence and number of
ampullate ‘‘nubbins’’ (i.e., ampullate nubbins
and/or tartipores) are being used as characters
in cladistic analyses (Coddington 1990; Hor-
miga et al. 1995; Scharff & Coddington 1997;
Griswold et al. 1998, 1999; Hormiga 2000),
accurately determining the presence/absence
of ampullate tartipores and nubbins, and mak-
ing a clear distinction between the two, can
only aid phylogenetic studies.
Occurrence of tartipores.—Tartipores can
occur in the exoskeletons of adults and juve-
niles, at least as early as second instars (see
Methods for the definition of the first instar
used in this paper). We have not seen and are
not aware of any reports of tartipores in first
instars. However, given the occurrence of
functioning silk glands and spigots in postem-
bryos of at least some mygalomorph taxa
(Bond 1994), the possibility of tartipores in
first instars of such taxa cannot be dismissed.
But at least for those araneomorphs in which
functional silk glands and spigots first appear
in first instars, tartipores do not need to form
until deposition of the second instar cuticle
begins. Consequently, for such spiders, tarti-
pores would first be seen in second instars.
(The presence of one, presumably 1
8
, MaA
spigot base per ALS in Nephila (Tetragnathi-
dae) postembryos has been described by Ble-
her (2000), but their ability to produce silk is
uncertain and only the ducts of 2
8
ampullate
glands are known to be accommodated by am-
pullate tartipores.)
It is of interest, therefore, that protuberanc-
es, reminiscent of but recognizably different
from tartipores, are sometimes evident on the
ALS and PMS of first instars, in positions
consistent with those of ampullate tartipores
in later instars. We have seen them near 2
8
ampullate spigots on the ALS and PMS of
first instar Pardosa xerampelina (Keyserling
1877) (Figs. 2–5) and Octonoba sinensis (Si-
mon 1880) (Uloboridae), and on the PMS of
first instar Argiope aurantia Lucas 1833 (Ar-
aneidae) and Herpyllus ecclesiasticus Hentz
1832 (Gnaphosidae). We tentatively refer to
them as ‘pre-tartipores’ (not to be confused
with the tartipore progenitors referred to in
Fig. 1). If they truly are precursors of the tar-
tipores in later instars, their occurrence sug-
gests that the epithelial cells that are capable
of forming tartipores, at least ampullate tarti-
pores, are already determined by the postem-
bryo stage.
Occurrence of nubbins.—In general, nub-
bins as here defined occur in adults, being
more abundant in males (largely since silk
glands used solely or primarily in prey capture
tend to regress in adult males), and are onto-
genetically vestigial. That is, they are located
in adults in positions where functional spigots
would have formed if the spider had remained
a juvenile after its most recent molt. In addi-
tion to the MaA and MiA nubbins present in
a variety of adult male and female araneocla-
217TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 2–5.—ALS and PMS from a first instar Pardosa xerampelina (removed from dorsum of its
mother’s abdomen) showing protuberances, tentatively termed ‘pre-tartipores’, in positions that are held
by ampullate tartipores in later instars: 2. Right ALS, entire spinning field shown (two MaA and three
piriform spigots), pre-tartipore in box; 3. Higher magnification of pre-tartipore from Fig. 2; 4. Right PMS,
entire spinning field shown (two MiA and three aciniform spigots), pre-tartipore in box; 5. Higher mag-
nification of pre-tartipore from Fig. 4. Posterior at top, lateral at right in all four figures. Scale bars (2, 4)
5
5
m
m; (3, 5)
5
1
m
m.
218 THE JOURNAL OF ARACHNOLOGY
Table 2.—Spider species in which adult females are known to have two MaA spigots (1
8
and 2
8
)on
each ALS and two MiA spigots (1
8
and 2
8
) on each PMS while adult males have one MaA spigot (1
8
)
on each ALS and one MiA spigot (1
8
) on each PMS. The families listed here are almost certainly not the
only ones that contain species fitting this description. Note that Neoramia (Agelenidae) apparently do not
fit this description (Griswold et al. 1999); nor do some salticid genera (see ‘Ampullate gland spigot,
nubbin, tartipore complements’ in Results) or several amaurobiid genera, including Amaurobius (see ‘Com-
parative ampullate gland spigot morphology’ in Discussion). Also, this description may not extend to all
Coras species (see ‘Ampullate gland spigot, nubbin, tartipore complements’ in Results).
Family Species References
Lycosidae Gladicosa gulosa (Walckenaer 1837) this study
Pardosa amentata (Clerck 1757)
Pardosa lapidicina Emerton 1885
Pardosa lugubris (Walckenaer 1802)
Pardosa modica (Blackwall 1846)
Pardosa moesta Banks 1892
Pardosa saxatilis (Hentz 1844)
Richter 1970
this study
Wa¸sowska 1977
this study
this study
this study
Pisauridae
Agelenidae
Dolomedes scriptus Hentz 1845
Pisaurina mira (Walckenaer 1837)
Agelena labyrinthica (Clerck 1757)
Agelenopsis naevia (Walckenaer 1842)
Agelenopsis potteri (Blackwall 1846)
this study
this study
Kokocin´ski 1968
this study
this study
Amaurobiidae
Thomisidae
Philodromidae
Coras aerialis Muma 1946
Misumenops asperatus (Hentz 1847)
Xysticus cristatus (Clerck 1757)
Tibellus oblongus (Walckenaer 1802)
this study
this study
Wa¸sowska 1977
Wa¸sowska 1967, 1977; this study
Clubionidae Clubiona phragmitis C.L. Koch 1843 Mikulska 1969; Wa¸sowska 1969;
Wis´niewski 1986a,b
Miturgidae
Salticidae Cheiracanthium mildei L. Koch 1864
Salticus scenicus (Clerck 1757) this study
this study
dans, flagelliform and aggregate nubbins form
on the PLS of many adult male araneoids
(Sekiguchi 1955a; Peters & Kovoor 1991:fig.
3b; Platnick et al. 1991:fig. 275; Townley et
al. 1991:fig. 16; Townley 1993:fig. 16; Gris-
wold et al. 1998:figs. 25D, 39D, 43D), though
a number of males within the ‘reduced piri-
form clade’ of Griswold et al. (1998) (see also
Hormiga 2000) and the Micropholcommatidae
(Schu¨tt 2000) retain the aggregate/flagelliform
spigot triad. Other examples include aciniform
nubbins on the PMS (Mu¨ller & Westheide
1993) and PLS (pers. obs.) of adult male Ar-
giope and on the PMS of some adult male
uloborids (Kovoor & Peters 1988:53), pseu-
doflagelliform nubbins on the PLS and para-
cribellar nubbins on the PMS of some adult
male deinopoids (Kovoor & Peters 1988; Pe-
ters 1992), paracribellar nubbins on the PMS
of adult male austrochilids (Peters 1983; Plat-
nick et al. 1991:fig. 33), and nubbins of un-
certain gland type on the ALS of adult male
Hypochilus, next to the single, large ampullate
spigot (Platnick et al. 1991:fig. 4; Townley
1993:figs. 17D,E). By the interpretation of
Platnick et al. (1991:51) the latter would be
MaA nubbins.
Among the examples of exceptions to the
general rule are nubbins occasionally seen in
early instar Cyrtophora (Araneidae) that sug-
gest phylogenetic vestiges of either aggregate
or flagelliform spigots. In an examination of
six first instar Cyrtophora citricola (Forska˚l
1775), on one PLS Peters (1993:figs. 11b, c)
observed a single ‘‘shaft-like structure’’ on the
vestigial plate of the aggregate-flagelliform
triad. (On nine PLS one or two ‘‘knobs with
pores’’ were seen on these vestigial plates, but
we do not interpret these as nubbins.) Nubbins
that are also apparently phylogenetic vestiges
of aggregate spigots are often retained right
up to maturity in female Drapetisca socialis
(Sundevall 1833) (Linyphiidae); functional
aggregate spigots are absent throughout the
ontogeny of these spiders (Schu¨tt 1995). The
occurrence of a MaA nubbin on the ALS of
penultimate instar female Malala lubinae Da-
219TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 6–7.—Right PLS on the last exuvium shed
by a female Hogna sp. killed while a penultimate in-
star (i.e. the cuticle of the antepenultimate instar is
shown): 6. Three aciniform nubbins, among aciniform
(and cylindrical?) spigots, can be seen within the box;
7. Higher magnification of the three aciniform nubbins
(*) from Fig. 6, arrow to example of an aciniform
tartipore. Posterior at left, lateral at top in both figures.
Scale bars (6)
5
100
m
m; (7)
5
25
m
m.
vies 1993 (tentatively Amaurobiidae, Platnick
2002) (Davies 1993) is also atypical.
In the course of the present study we ob-
served a few consistently-located aciniform
nubbins on the PMS (Figs. 19–21) and PLS
(Figs. 6, 7) of some juvenile and adult lycos-
ids, the details of which are given in the Re-
sults. The reason these nubbins form with as
much regularity as we have seen, particularly
in Hogna, remains to be explained. Certainly,
it is not unusual in an aciniform or piriform
spinning field to encounter occasional incom-
pletely formed spigots (i.e., nubbins). But
such nubbins are presumably teratological,
given their random and, typically, asymmet-
rical occurrence (present on one spinneret but
not its pair). The occurrence of aciniform nub-
bins in Hogna, on the other hand, is neither
random nor asymmetrical. Because they are
present in juvenile females, they are another
example of atypical nubbins.
Nubbins resulting from different gland
types and, in some cases, nubbins of the same
gland type in different taxa vary considerably
in the extent to which their development pro-
ceeds before it is aborted. Thus, nubbins may
range from being small mounds or small
spherical or oblong protuberances, e.g., in the
case of some MaA and MiA nubbins, to being
essentially normal spigot bases on which
shafts never develop, e.g., in the case of many
aciniform and paracribellar nubbins of adult
males, as well as the aforementioned nubbins
of adult male Hypochilus. It may even be with
these more developed nubbins that a shaft
does sometimes form on the spigot base, but
the junction of the base and shaft appears mal-
formed, suggesting that the spigot and/or the
silk gland it serves actually are not functional.
At the opposite extreme, we have indica-
tions that in some taxa (the three examples
seen thus far are dionychans) it is common for
certain nubbins not to form at all. In an adult
male Salticus scenicus (Clerck 1757) (Salti-
cidae), we observed single 2
8
ampullate nub-
bins on both ALS and the left PMS, but not
on the right PMS (Table 3). That this individ-
ual had 2
8
MiA spigots when it was a penul-
timate instar, on the right PMS as well as the
left PMS, is indicated by the MiA nubbin on
the left PMS and the MiA tartipores on both
PMS. The same situation was seen in an adult
male Tibellus oblongus (Walckenaer 1802)
(Philodromidae) and an adult male Misumen-
220 THE JOURNAL OF ARACHNOLOGY
Figures 8–11.—Portions of the ALS and PMS containing the ampullate spigots, from an adult female
Pardosa saxatilis and the last exuvium shed by this individual (i.e. the cuticle of the penultimate instar):
8, 10. Penultimate instar; 9, 11. Adult; 8, 9. Left ALS (posterior at right, lateral at top); 10, 11. Right
PMS (posterior at left, lateral at top). Note the relative increase in diameter of the bases of the 2
8
MaA
and 2
8
MiA spigots in the adult cuticle and that the MiA tartipore and 2
8
MiA spigot switch positions
from one instar to the next. Unlabeled arrows point to examples of piriform (Figs. 8–9) or aciniform (Fig.
10) tartipores. Scale bars
5
10
m
m.
ops asperatus (Hentz 1847) (Thomisidae), ex-
cept that for the latter it was on the left PMS
that a MiA nubbin was lacking (Table 3). The
one MiA nubbin that was present on the S.
scenicus and M. asperatus individuals was
very small, as was the right MiA nubbin on a
second adult male M. asperatus. But the left
MiA nubbin on the latter spider was much
larger (likewise in the T. oblongus individual),
clearly showing cuticular sculpturing in the
form of longitudinal ridges like those on the
bases of the 1
8
MiA spigots.
Finally, in adult males it is sometimes the
case with multiple nubbins that not all of the
spigots of a given gland type are represented
by nubbins; some appear to develop into func-
tional spigots (at least they have shafts and the
base-shaft junctions do not look malformed).
METHODS
Spiders examined.—Spinnerets with at-
tached egg sacs were examined by scanning
electron microscopy (SEM) in Pardosa moes-
ta Banks 1892 (6 specimens), Pardosa lapi-
dicina Emerton 1885 (2 specimens), Pardosa
modica (Blackwall 1846) (1 specimen), Par-
dosa littoralis Banks 1896 (1 specimen), and
Trochosa ruricola (De Geer 1778) (1 speci-
men). The numbers of specimens given in-
clude only those that yielded useful informa-
tion. The P. littoralis was collected in central
South Carolina. The others were collected in
southeastern (se) New Hampshire (NH).
Other spinnerets were also examined, both
from several lycosid species (without attached
egg sacs) and from species belonging to other
families (mostly those in which adult females
221TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
retain functional 2
8
MaA/2
8
MiA glands) (see
Table 3). These were also collected in se NH
with the following exceptions: P. xerampelina
(se NH and southwestern Maine), Pardosa
hortensis (Thorell 1872) (Luxembourg), Par-
dosa lugubris (Walckenaer 1802) (Luxem-
bourg), Gladicosa gulosa (Walckenaer 1837)
(southern NH and central Virginia), Hogna
helluo (Walckenaer 1837) (se NH and central
Virginia), Agelenopsis naevia (Walckenaer
1842) (central Virginia and western North
Carolina), Coras aerialis Muma 1946 (se NH
and southwestern Maine), Coras lamellosus
(Keyserling 1887) (southwestern Maine), Cor-
as montanus (Emerton 1890) (southwestern
NH), Misumenops oblongus (Keyserling
1880) (southwestern NH), Thanatus rubicellus
Mello-Leita˜o 1929 (central Virginia), and Phi-
dippus audax (Hentz 1845) (western Pennsyl-
vania and se NH). Some spiders were collect-
ed as juveniles and several antepenultimate or
penultimate instar lycosids were prepared for
SEM immediately. The others were raised to
the adult stage with shed exuvia saved for lat-
er examination. Spinnerets on exuvia from a
few of the lycosids were prepared for SEM,
as described below, in order to compare ju-
venile and adult spinning fields within the
same individual. There were two reasons for
examining spinnerets other than lycosid spin-
nerets to which egg sacs were attached: first,
to gain a more complete view of spinning field
morphology in lycosids, since attached egg
sacs usually make clear viewing of spinning
fields difficult or impossible; and, second, to
compare spinning fields, especially ampullate
spigots, between males and females, between
juveniles and adults, and among different ly-
cosid and non-lycosid species.
Spiders were identified using keys and de-
scriptions given, primarily, in Chamberlin &
Ivie (1941), Carico (1972, 1973), Dondale &
Redner (1978, 1982, 1990), Brady (1979,
1986), Kaston (1981), Roberts (1985), Heimer
& Nentwig (1991), Roth (1993), and Prentice
(2001). Family assignments and taxonomic ci-
tations follow Platnick (2002). Voucher spec-
imens for this study are deposited in the Mu-
seum of Comparative Zoology, Harvard
University. Most consist only of cephalotho-
raxes and isolated epigyna since the spinnerets
of nearly all collected specimens were pro-
cessed for SEM.
SEM.—Spinnerets without attached egg
sacs were artificially spread in preparation for
SEM using the forceps squeeze technique of
Coddington (1989), which is a modified ver-
sion of a technique suggested to that author
by J. Kovoor. Carbon dioxide anesthetized
spiders were severed at the pedicel, the tines
of fine forceps were placed on either side of
the spinnerets, one dorsad and one ventrad,
and the forceps were squeezed. Any fecal ma-
terial ejected from the stercoral sac through
the anal tubercle was either absorbed with a
tissue or rinsed off with distilled water. In-
spired by Coddington’s (1989) recommenda-
tion that live spiders be killed by immersion
in boiling water or fixative to spread spinner-
ets, the spread spinnerets were immersed in
boiling water (about 2–5 sec depending on the
size of the abdomen) as a kind of first fixation.
The forceps were then held closed using a
snug-fitting rubber O-ring (Fine Science
Tools, Inc.) and their tips with the held ab-
domen were inserted through a hole, just large
enough to accommodate the forceps, made in
the cap of a 20 ml scintillation vial filled with
a modified version of Karnovsky’s (1965) fix-
ative containing only 1% glutaraldehyde / 1%
formaldehyde in 0.1 M cacodylate buffer, pH
7.2. Abdomens were kept refrigerated in the
fixative from overnight to several days, al-
lowed to come to room temperature, trans-
ferred to distilled water for about 20 min, and
then taken through an ethanol series (30%,
50%, 70%, 85%, 95%, 100% used once,
100% fresh; 1–2 hr in each
,
70%, 2 hr-over-
night in each
$
70%). Finally, the samples
were immersed in hexamethyldisilazane
(HMDS) (Nation 1983) overnight and then al-
lowed to air dry. All solutions/solvents were
also in scintillation vials so transfers were
made by moving cap, forceps, and abdomen
as a unit from one vial to the next. Abdomens
were mounted on pin-type SEM specimen
mounts (stubs) with carbon adhesive tabs
(Electron Microscopy Sciences) and carbon
paste (Structure Probe, Inc.), sputter-coated
with gold/palladium (about 20 nm), and ex-
amined on an AMR Model 3300 FE field-
emission SEM operated with a 7 kV acceler-
ating voltage.
Some lycosid spinnerets with an attached
egg sac were prepared for SEM using the
same protocol just described, except that the
specimen was not immersed in boiling water
and the spinnerets were only partially spread
222 THE JOURNAL OF ARACHNOLOGY
Table 3.—Ampullate spigot (spig), nubbin (nub), and tartipore (tart) complements in examined spiders belonging to the families listed in Table 2, as well
as in examined Mimetus (see ‘Categories of ampullate silk glands etc.’ in the introductory section). Where these complements differed between the left and
right spinnerets of an individual, both numbers are shown, with the left spinneret value placed on the left. With the exception of spinnerets from P. mira
juveniles (identified by P. Sierwald), the spinnerets from those juveniles that are identified to species were obtained either from exuvia shed by spiders raised
to adults or from the progeny of identified females. n
5
number of individuals examined. Ad
5
adult, ALS
5
anterior lateral spinneret, An
5
antepenultimate,
F
5
female, M
5
male, MaA
5
major ampullate, MiA
5
minor ampullate, P
5
penultimate, PMS
5
posterior median spinneret, U
5
unknown.
Family
Species Instar Sex n
Number of
MaA spig
per ALS
Number of
MaA nub
per ALS
Number of
MaA tart
per ALS
Number of
MiA spig
per PMS
Number of
MiA nub
per PMS
Number of
MiA tart
per PMS
Lycosidae
Gladicosa gulosa (Walckenaer 1837)
Hogna sp.
Hogna aspersa (Hentz 1844)
Hogna helluo (Walckenaer 1837)
Ad
Ad
An
P
Ad
P
Ad
F
M
F
F
F
F
F
4
2
1
1
1
1
2
2
1
2
2
2
2
2
0
1
0
0
0
0
0
1
1
1
1
1
1
1
2
1
2
2
2
2
2
0
1
0
0
0
0
0
1
1
1
1
1
1
1
Pardosa sp.
Pardosa hortensis (Thorell 1872)
Pardosa lapidicina Emerton 1885
Pardosa littoralis Banks 1896
Pardosa lugubris (Walckenaer 1802)
An or P
P
P
Ad
Ad
Ad
Ad
P
F
M
F
F
F
M
F
M
6
3
1
1
2
2
1
1
2
2
2
2
2
1
2
2
0
0
0
0
0
1
0
0
1
1
1
1
1
1
1
1
2
2
2
2
2
1
2
2
0
0
0
0
0
1
0
0
1
1
1
1
1
1
1
1
Pardosa modica (Blackwall 1846)
Pardosa moesta Banks 1892
Pardosa saxatilis (Hentz 1844)
Ad
Ad
Ad
Ad
Ad
P
Ad
Ad
F
M
F
M
M
F
F
M
1
1
8
1
1
1
2
1
2
1
2
1
1
2
2
1
0
1
0
1
1
0
0
1
1
1
1
1
1
1
1
1
2
1
2
1,2
1
2
2
1
0
1
0
1,0
1
0
0
1
1
1
1
1
1
1
1
1
Pardosa xerampelina (Keyserling 1877)
Trochosa ruricola (De Geer 1778)
Trochosa terricola Thorell 1856
1st
2nd
4th
Ad
3rd
Ad
Ad
U
U
U
F
U
F
M
4
1
1
1
1
1
1
2
2
2
2
2
2
1
0
0
0
0
0
0
1
0
1
1
1
1
1
1
2
2
2
2
2
2
1
0
0
0
0
0
0
1
0
1
1
1
1
1
1
223TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Table 3.—Continued.
Family
Species Instar Sex n
Number of
MaA spig
per ALS
Number of
MaA nub
per ALS
Number of
MaA tart
per ALS
Number of
MiA spig
per PMS
Number of
MiA nub
per PMS
Number of
MiA tart
per PMS
Pisauridae
Dolomedes scriptus Hentz 1845
Dolomedes tenebrosus Hentz 1844
Pisaurina mira (Walckenaer 1837)
Ad
Ad
Ad
An or P
Ad
An
Ad
F
M
F
F
F
M
M
3
1
1
1
2
1
2
2
1
2
2
2
2
1
0
1
0
0
0
0
1
1
1
1
1
1
1
1
2
1
2
2
2
2
1
0
1
0
0
0
0
1
1
1
1
1
1
1
1
Agelenidae
Agelenopsis naevia (Walckenaer 1842)
Agelenopsis potteri (Blackwall 1846)
Ad
Ad
Ad
Ad
F
M
F
M
2
2
6
2
2
1
2
1
0
1
0
1
1
1
1
1
2
1
2
1
0
1
0
1
1
1
1
1
Amaurobiidae
Coras aerialis Muma 1946
Coras lamellosus (Keyserling 1887)
Coras montanus (Emerton 1890)
Ad
Ad
Ad
Ad
F
M
F
M
1
2
1
1
2
1
2,1
1
0
1
0,1
1
1
1
1
1
2
1
2
2
0
1
0
0
1
1
1
1
Thomisidae
Misumena vatia (Clerck 1757)
Misumenops asperatus (Hentz 1847)
Misumenops oblongus (Keyserling 1880)
Ad
Ad
Ad
Ad
Ad
F
F
M
M
F
1
2
1
1
1
2
2
1
1
2
0
0
1
1
0
1
1
1
1
1
2
2
1
1
2
0
0
1
0,1
0
1
1
1
1
1
Xysticus emertoni Keyserling 1880
Xysticus ferox (Hentz 1847)
Xysticus punctatus Keyserling 1880
3rd
Ad
Ad
4th
5th
U
M
M
U
U
1
1
2
1
2
2
1
1
2
2
0
1
1
0
0
1
1
1
1
1
2
1
1
2
2
0
1
1
0
0
1
1
1
1
1
224 THE JOURNAL OF ARACHNOLOGY
Table 3.—Continued.
Family
Species Instar Sex n
Number of
MaA spig
per ALS
Number of
MaA nub
per ALS
Number of
MaA tart
per ALS
Number of
MiA spig
per PMS
Number of
MiA nub
per PMS
Number of
MiA tart
per PMS
Philodromidae
Philodromus vulgaris (Hentz 1847)
Thanatus rubicellus Mello-Leita˜o 1929
Tibellus oblongus (Walckenaer 1802)
Ad
Ad
Ad
Ad
M
M
F
M
1
1
1
1
1
1
2
1
1
1
0
1
1
1
1
1
1
1
2
1
1
1
0
1,0
1
1
1
1
Miturgidae
Cheiracanthium mildei L. Koch 1864 Ad
Ad F
M2
32
10
11
12
10
11
1
Salticidae
Phidippus audax (Hentz 1845)
Salticus scenicus (Clerck 1757)
Sitticus pubescens (Fabricius 1775)
Ad
Ad
Ad
Ad
Ad
F
M
F
M
M
2
3
1
1
1
2
2
2
1
?
0
0
0
1
?
1
1
1
1
?
2
2
2
1
1
0
0
0
1,0
1
1
1
1
1
1
Mimetidae
Mimetus notius Chamberlin 1923
Mimetus puritanus Chamberlin 1923 Ad
Ad
Ad
M
F
M
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
225TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
by placing the forceps more anteriorly on the
abdomen with the tines extending only part-
way across the width of the abdomen. While
some spiders were severed at the pedicel, oth-
ers were left intact. With other specimens, Pe-
ters’ (1982) paraffin technique was applied af-
ter anesthetization, with or without squeezing
the abdomen. Hot paraffin was added either
dorsally or ventrally to the spinnerets and the
adjacent part of the egg sac, fusing the egg
sac to the spinnerets. The abdomen with at-
tached egg sac was then immersed directly in
70% ethanol, taken to 100% ethanol as above,
then transferred to benzene, which was
changed twice over the course of at least a
few weeks. The specimen was then air dried,
mounted, sputter-coated, and examined. We
never settled on a uniform protocol since no
one of the variants emerged as clearly superior
to the others with all specimens or species,
though squeezing the abdomen is most often
warranted.
As mentioned, spinnerets on exuvia shed by
a few lycosids were also examined by SEM.
These were prepared by isolating the spinneret
group and sticking it to a carbon adhesive tab
on a SEM stub. A small volume of an aqueous
solution containing a detergent (to reduce sur-
face tension and increase wettability) was then
applied to the spinnerets prior to attempting
to uncrumple the spinnerets. We used Lae-
mmli’s (1970) sodium dodecyl sulfate electro-
phoresis sample buffer, diluted to about half
strength, for this purpose, though other com-
positions would no doubt serve at least as
well. While immersed in this solution, an in-
sect pin or a tine on a pair of fine forceps was
inserted into each spinneret in order to re-ex-
pand it and allow its spinning field to be
viewed. The spinneret group was then fixed
(though this is probably not necessary), de-
hydrated, treated with HMDS, air dried,
mounted, sputter-coated, and examined as de-
scribed above.
Distinguishing 1
8
ampullate spigots from
2
8
ampullate spigots.—In the results present-
ed below, differences between 1
8
and 2
8
am-
pullate fibers and spigots of adult female ly-
cosids are noted. Our interpretation of the
relative contributions made by 1
8
and 2
8
am-
pullate glands to egg sac attachment relies on
having identified 1
8
and 2
8
ampullate spigots
correctly. We considered four lines of evi-
dence in making these identifications and, in
the manner of Coddington (1989), the same
kind of reasoning can be applied to many oth-
er spider families. (1) Since 2
8
ampullate spig-
ots are represented only by ampullate nubbins
in adult male lycosids (Figs. 13, 15), their po-
sitions relative to the 1
8
ampullate spigots can
be used as a key to distinguishing 1
8
from 2
8
ampullate spigots in adult females and juve-
niles. (2) Because only 2
8
ampullate glands
are functional during proecdysis (right up un-
til the old cuticle is shed from the spinnerets),
the only ampullate fibers emerging from spig-
ots on the old cuticle during proecdysis are 2
8
ampullate fibers (Townley et al. 1993). Con-
sequently, the only ampullate fibers on the
exuvium after ecdysis pass through 2
8
ampul-
late spigots (Townley et al. 1991:figs. 14–15;
Townley et al. 1993:fig. 4). While 2
8
ampul-
late fibers do not invariably remain attached
to exuvia, they do so with enough regularity
that examinations of exuvia can be used to
determine which ampullate spigots are 1
8
and
which are 2
8
. Thus, in our scans of lycosid
exuvia, we have only observed ampullate fi-
bers emerging from those spigots that we have
identified as 2
8
ampullate spigots, never from
those identified as 1
8
ampullate spigots (Figs.
8, 10, 19, 20). (3) In general, ampullate tar-
tipores, resulting from openings made to ac-
commodate 2
8
ampullate gland ducts (see Ter-
minology section above), occur closer to 2
8
ampullate spigots than 1
8
ampullate spigots.
The ampullate tartipores on lycosid ALS and
PMS likewise occur closer to the spigots iden-
tified as 2
8
ampullate spigots (e.g., Figs. 18,
21). (4) The arrangement of MaA and piri-
form spigots on the ALS of lycosids is essen-
tially the same as in A. cavaticus (the same
cannot be said of the arrangement of spigots
on the PMS where the MiA spigots are locat-
ed). In this araneid we have observed by dis-
section that the 2
8
MaA ducts lead to the more
posteriorly placed ampullate spigot on each
ALS (Townley et al. 1991, 1993). The spigot
identified as the 2
8
MaA spigot in lycosids
likewise occurs posterior to that identified as
the 1
8
MaA spigot.
Definition of first instar.—Downes’
(1987) definition for the first instar is followed
in this report with subsequent instars num-
bered accordingly. A spider becomes a first
instar as a result of the first ecdysis that pro-
duces an exuvium that both has legs and does
not remain attached to the spider. In the period
226 THE JOURNAL OF ARACHNOLOGY
Figures 12–15.—Entire spinning fields on ALS and PMS from a penultimate instar male Pardosa sp.
and an adult male Pardosa modica: 12, 14. Penultimate instar; 13, 15. Adult; 12, 13. Left ALS (posterior
at right, lateral at top); 14, 15. Right PMS (posterior at left, lateral at top). Note that 2
8
MaA and 2
8
MiA
spigots are represented in the adult male only by MaA and MiA nubbins, respectively. Unlabeled arrows
point to examples of piriform (Figs. 12–13) or aciniform (Figs. 14–15) tartipores. Scale bars (12, 14)
5
10
m
m; (13, 15)
5
20
m
m.
between hatching and the ecdysis that yields
a first instar, the spider is a postembryo
(Downes 1987).
RESULTS
Abbreviations on micrographs.—AC
5
attachment cone; C
5
cylindrical gland spig-
ot; ES
5
egg sac; N
5
MaA nubbin (if the
spinneret is an ALS) or MiA nubbin (if the
spinneret is a PMS); 1
85
1
8
MaA spigot (on
ALS) or 1
8
MiA spigot (on PMS); 2
85
2
8
MaA spigot (on ALS) or 2
8
MiA spigot (on
PMS); T
5
MaA tartipore (on ALS) or MiA
tartipore (on PMS); l
5
left; r
5
right.
Ampullate gland spigot, nubbin, tarti-
pore complements.—Spinnerets from male
and female representatives of three lycosid
genera (Pardosa,Gladicosa,Trochosa) were
examined, as were those from females only of
the genus Hogna (Table 3). With one clearly
anomalous exception (see Table 3), no varia-
tion was seen with regard to the number of
ampullate gland spigots/nubbins/tartipores for
a given sex at a given stage of development.
Assuming these genera present the typical, if
not invariable, lycosid condition, adult fe-
males and juveniles of both sexes that are at
least second instars have two MaA spigots and
one MaA tartipore on each ALS (Figs. 8, 9,
12, 16–18, 22, 40, 41) and two MiA spigots
and one MiA tartipore on each PMS (Figs. 10,
11, 14, 19–21, 23, 38, 39, 42) (first instars
lack tartipores, Figs. 2–5), while adult males
have one MaA spigot, one MaA nubbin, and
one MaA tartipore on each ALS (Fig. 13) and
one MiA spigot, one MiA nubbin, and one
MiA tartipore on each PMS (Fig. 15). The 2
8
MaA and 2
8
MiA spigots of juvenile males
are vestigial in adult males, being represented
only by 2
8
MaA/2
8
MiA nubbins.
227TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 16–21.—Portions of the ALS and PMS containing the ampullate spigots, from an adult female
Hogna helluo and the last exuvium shed by this individual (i.e. the cuticle of the penultimate instar), as well
as from the last exuvium shed by a female Hogna sp. killed while a penultimate instar (i.e. the cuticle of the
antepenultimate instar; the same exuvium from which the PLS in Figs. 6–7 was taken): 16, 19. Antepenultimate
instar; 17, 20. Penultimate instar; 18, 21. Adult; 16, 17, 18. Left ALS (posterior at right, lateral at top); 19,
21. Left PMS (posterior at right, lateral at top); 20. Right PMS (posterior at left, lateral at top). The MaA
tartipore is largely obscured in Fig. 17. Unlabeled arrows point to examples of piriform (Figs. 16, 18) or
aciniform (Figs. 19, 21) tartipores. Asterisks (*) identify the two aciniform nubbins on each PMS. Scale bars
5
25
m
m.
228 THE JOURNAL OF ARACHNOLOGY
With regard to ampullate spigot/nubbin/tar-
tipore complements, apart from the absence of
a MiA nubbin on one PMS in single adult
male specimens of M. asperatus and T. ob-
longus, no departures from the lycosid con-
dition were observed when viewing spinnerets
from the pisaurid (Figs. 24–27), agelenid
(Figs. 28, 29), thomisid (Figs. 32, 33), phil-
odromid (Figs. 30, 31), and miturgid (Figs.
34, 35) species listed in Table 3. The same is
true of the C. aerialis examined. But in an
adult female C. lamellosus, the number of
MaA spigots/nubbins differed between the
right and left ALS (Table 3). The formation
of a MaA nubbin rather than a 2
8
MaA spigot
on one ALS in this individual again seems to
be an example of an anomaly. More signifi-
cant was the presence of 2
8
MiA spigots on
both PMS (and the absence of 2
8
MaA spigots
on both ALS) of an adult male C. montanus
(Table 3). This indicates that, in Coras, com-
plements differ either interspecifically or in-
traspecifically. In P. audax, adult males, as
well as adult females (Figs. 36, 37), retain 2
8
MaA and MiA spigots (Table 3). This is also
the case in the salticid Philaeus chrysops
(Poda 1761) (Millot 1935) and is consistent
with Millot’s (1935:509) statement that sexual
dimorphism in the spinning apparatus of sal-
ticids is negligible. It is, therefore, worth not-
ing that adult male specimens of S. scenicus
and Sitticus pubescens (Fabricius 1775) (one
of each species) lacked 2
8
ampullate spigots
(Table 3), though for the latter we have only
PMS data. These spigots were represented by
nubbins and, in the case of one PMS on the
S. scenicus individual, it appears that even the
nubbin did not form (see ‘Occurrence of nub-
bins’ in the introductory section). As an adult
female S. scenicus did have 2
8
ampullate spig-
ots (Table 3), it appears that some salticids
match the lycosid condition (Table 2) while
others do not.
Relative sizes of 1
8
and 2
8
ampullate
gland spigots.—In early instar lycosids (first
and second instar P. xerampelina, third instar
T. ruricola), 1
8
and 2
8
MaA spigots are rough-
ly comparable in size, with the bases and
shafts of the 1
8
MaA spigots a little greater in
diameter than those of the 2
8
MaA spigots
(Fig. 2). Typically, MiA spigots more clearly
differ in size, again with the shafts and bases
of 1
8
MiA spigots wider than those of 2
8
MiA
spigots (Fig. 4). In Pardosa of both sexes, the
1
8
ampullate spigots, especially the 1
8
MiA
spigots (Fig. 14), may retain marginally to
moderately greater size through the penulti-
mate instar, or 1
8
and 2
8
ampullate spigots
may have about the same diameter (Figs. 8,
12). In contrast, in female Hogna, it is the 2
8
ampullate spigots that tend to be larger in the
antepenultimate (Figs. 16, 19) and penultimate
(Figs. 17, 20) instars. The difference is more
pronounced on the ALS, with the bases of the
2
8
MaA spigots clearly larger than those of
the 1
8
MaA spigots. Also, the difference in
size between ampullate spigots and nearby
aciniform or piriform spigots is greater in
these Hogna juveniles than in juvenile Par-
dosa, due, seemingly, to disproportionately
larger ampullate spigots (rather than smaller
aciniform and piriform spigots) (cf. Figs. 8,
10, 12, 14 with Figs. 16, 17, 19, 20).
With the final molt in female Pardosa, the
bases of the 2
8
ampullate spigots become de-
cisively wider than those of the 1
8
ampullate
spigots (cf. Fig. 8 with Fig. 9, and Fig. 10 with
Fig. 11, all taken from the same individual).
The degree to which this occurs varies notice-
ably within a species. Nevertheless, it has
been apparent in all adult female Pardosa that
we have examined (Table 3). The relative dis-
parity between 1
8
and 2
8
ampullate spigots
may also increase after the last molt in female
H. helluo so that it is perhaps more obvious
in adults than in penultimate instars that the
bases of the 2
8
ampullate spigots have greater
diameters than those of the 1
8
ampullate spig-
ots (cf. Fig. 17 with Fig. 18, and Fig. 20 with
Fig. 21). However, the changes seen following
the last molt in Hogna females are certainly
not as dramatic as those observed in Pardosa.
In adult female G. gulosa (Figs. 22, 23) and
T. ruricola (Fig. 43), the bases of the 2
8
am-
pullate spigots, especially the 2
8
MaA spigots,
are likewise of greater diameter than the 1
8
ampullate spigot bases.
Among the adult female pisaurids examined
(Table 3), noticeably (but not greatly) wider
2
8
ampullate spigot bases were observed on
the ALS and/or PMS of all three Dolomedes
scriptus Hentz 1845 (Figs. 24, 25), though one
of these had 1
8
and 2
8
MaA spigots of essen-
tially the same size while a second spider had
1
8
and 2
8
MiA spigots of similar size. There
was also little difference in size between 1
8
and 2
8
ampullate spigots on the one D. tene-
brosus Hentz 1844 examined, with 2
8
spigots
229TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 22–27.—Portions of the ALS and PMS containing the ampullate spigots, from adult females:
22, 23. Gladicosa gulosa (Lycosidae); 24, 25. Dolomedes scriptus (Pisauridae); 26, 27. Pisaurina mira
(Pisauridae); 22, 24. Right ALS (posterior at left, lateral at top); 26. Left ALS (posterior at right, lateral
at top); 25, 27. Right PMS (posterior at left, lateral at top); 23. Left PMS (posterior at right, lateral at
top). Unlabeled arrows point to examples of piriform (Figs. 22, 26) or aciniform (Figs. 23, 25, 27)
tartipores. An arrowhead in Fig. 23 points to an aciniform nubbin. Scale bars
5
25
m
m.
230 THE JOURNAL OF ARACHNOLOGY
only marginally larger. In Pisaurina,1
8
and 2
8
MaA spigots were of about the same size (Fig.
26) and 1
8
MiA spigots were larger than 2
8
MiA spigots (Fig. 27).
Among the non-lycosoid adult females ex-
amined (Table 3, Figs. 28–37), larger-diameter
2
8
ampullate spigot bases were observed only
on the ALS (and not the PMS) of two thom-
isids, M. oblongus (Figs. 32, 33) and Misu-
mena vatia (Clerck 1757). The difference was
small and, moreover, in two M. asperatus fe-
males the 1
8
MaA spigots were larger than or
about the same size as the 2
8
MaA spigots (not
shown). Likewise, in Agelenopsis (Figs. 28,
29), Coras (not shown), Tibellus oblongus
(Walckenaer 1802) (Figs. 30, 31), Cheiracan-
thium mildei L. Koch 1864 (Figs. 34, 35), P.
audax (Figs. 36, 37), and S. scenicus (not
shown), 2
8
ampullate spigots were either
about the same size as the 1
8
ampullate spigots
or smaller (though in adult P. audax, male and
female, the 2
8
ampullate spigots were longer
and in the one examined S. scenicus adult fe-
male, 2
8
MiA spigots were longer than 1
8
MiA
spigots while 1
8
and 2
8
MaA spigots were of
about equal length).
Egg sac attachment.—In the four Pardosa
species examined with attached egg sacs,
eight silk fibers emanating from the 1
8
and 2
8
MaA spigots and the 1
8
and 2
8
MiA spigots
were attached to the egg sac (Figs. 38–42).
Likewise, in a single specimen of T. ruricola,
1
8
and 2
8
MaA fibers at least were used to
secure the egg sac to the spinnerets (Fig. 43).
We were unable, however, to determine if this
individual was also using MiA fibers. In some
preparations, single to several piriform fibers
accompanied MaA fibers, but in no instances
were fibers observed coming from aciniform
or cylindrical (
5
tubuliform) gland spigots,
including those on the PLS.
In all five species, 2
8
ampullate fibers had
greater diameters than 1
8
ampullate fibers
(Figs. 38–43, 49). Measurements obtained on
some of these fibers from four of these species
are presented in Table 4 and, though the data
are few, provide an indication of the disparity
between 1
8
and 2
8
ampullate fibers. In con-
trast, on the left ALS of one of the examined
adult female D. scriptus, the 1
8
MaA spigot
was only slightly smaller than the 2
8
MaA
spigot and silk fibers emerging from these
spigots had diameters of 2.40
m
m and 2.50
m
m, respectively. On the left ALS of an adult
female M. asperatus, the 1
8
and 2
8
MaA spig-
ots were about the same size and fibers emerg-
ing from them had diameters of 2.47
m
m and
2.00
m
m, respectively.
Ampullate fibers from the spinnerets are typ-
ically attached to the surface of the egg sac by
groups of fibers that often take the form of a
cone (Figs. 38, 40, 44, 46, 48). The fibers in
these ‘‘attachment cones’’ have smaller diam-
eters than the 1
8
ampullate fibers (Fig. 47 and
many of them appear to be fused to one an-
other (Figs. 45, 47). They extend beyond the
cone for a short distance on the surface of the
egg sac. A single attachment cone secures one
or more ampullate fibers to the egg sac. Thus,
one (Fig. 44) to several (Figs. 38, 48) cones in
close proximity affix the eight ampullate fibers.
There appears to be a generic difference with
regard to the number of cones that are typically
produced (one or two in Gladicosa and Tro-
chosa, several in Pardosa), but more obser-
vations are needed to verify this.
Aciniform nubbins.—Though not an objec-
tive of this study, a small number of aciniform
nubbins were noticed on several lycosid spec-
imens and we think their seemingly non-ran-
dom occurrence warrants mention. In four fe-
male Hogna individuals (two adult H. helluo,
one adult H. aspersa (Hentz 1844), one pen-
ultimate instar Hogna sp.), two aciniform nub-
bins were observed on each PMS in the vicin-
ity of the MiA spigots. (No male Hogna have
been examined.) They were also present on the
most recent exuvium shed by the penultimate
instar (i.e., the cuticle of the antepenultimate
instar) and on the last exuvium shed by one of
the adult H. helluo. One of these nubbins oc-
curs anterior to the 1
8
MiA spigot, the other
lateral or posterolateral to the 2
8
MiA spigot
(Figs. 19–21). In addition, three aciniform nub-
bins were observed on each PLS, roughly in
the middle of the spinning field, in the penul-
timate instar and its most recent exuvium (Figs.
6, 7), as well as in one adult H. helluo. On the
other examined Hogna cuticles, either one or
two aciniform nubbins were found per PLS,
though this could be because these PLS prep-
arations were not fully expanded and additional
nubbins were obscured.
We have not seen these aciniform nubbins
in Pardosa or Trochosa. In an adult male G.
gulosa, two aciniform nubbins were on the left
PMS in the same positions as in Hogna, but
only the more lateral of the two was present
231TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 28–33.—Portions of the ALS and PMS containing the ampullate spigots, from adult females:
28, 29. Agelenopsis naevia (Agelenidae); 30, 31. Tibellus oblongus (Philodromidae); 32, 33. Misumenops
oblongus (Thomisidae); 28, 32. Right ALS (posterior at left, lateral at top); 30. Left ALS (posterior at
right, lateral at top); 29, 31, 33. Right PMS; 29, 33 Posterior at right, lateral at bottom; 31 Posterior at
top, lateral at right. Unlabeled arrows point to examples of piriform (Figs. 30, 32) or aciniform (Fig. 31)
tartipores. Scale bars (28)
5
25
m
m; (29)
5
50
m
m; (30–32)
5
15
m
m; (33)
5
10
m
m.
on the right PMS, and none were found on the
PLS. Of four adult female G. gulosa, one had
a single lateral aciniform nubbin on one PMS
(Fig. 23) but none on the other PMS or either
PLS, while on a second female we could find
only one nubbin on one of the PLS. No acin-
iform nubbins were seen on the spinnerets of
the other two females (though on one of these
our views of the PMS and PLS were limited
as the spinnerets were not well spread).
DISCUSSION
The observations presented in this paper
demonstrate that adult females of at least
some species of lycosids use 1
8
and 2
8
MaA/
MiA gland fibers to connect the egg sac to the
232 THE JOURNAL OF ARACHNOLOGY
Table 4.—Diameters (in
m
m) of ampullate fibers produced by adult female lycosids for attaching the
egg sac to the spinnerets. Measurements were made by SEM at magnifications
.
10,000
3
.n
5
number
of individuals from which fibers were measured. For each spider, corresponding fibers from both ALS/
PMS were measured and averaged. For P. moesta and P. lapidicina, the means so obtained for each
individual were then used to calculate the overall means
6
their standard errors.
Species n1
8
MaA fibers 2
8
MaA fibers 1
8
MiA fibers 2
8
MiA fibers
Pardosa moesta Banks 1892
Pardosa lapidicina Emerton 1885
Pardosa littoralis Banks 1896
Trochosa ruricola (De Geer 1778)
5
2
1
1
0.83
6
0.038
0.99
6
0.026
0.90
1.31
2.56
6
0.120
3.93
6
0.319
2.86
3.06
0.76
6
0.020
0.93
6
0.090
0.50
2.57
6
0.171
3.86
6
0.181
2.53
Figures 34–37.—ALS and PMS from adult females: 34, 35. Cheiracanthium mildei (Miturgidae); 36,
37. Phidippus audax (Salticidae); 34, 36. Portion of right ALS containing the MaA spigots (posterior at
left, lateral at top); 35. Portion of left PMS containing the MiA spigots (posterior at right, lateral at top);
37. Left PMS, entire spinning field shown (two MiA and two aciniform spigots) (posterior at right, lateral
at top). Unlabeled arrows point to examples of piriform (Figs. 34, 36) or aciniform (Fig. 37, the only one)
tartipores. Scale bars
5
25
m
m.
spinnerets. Given the greater diameter of the
2
8
ampullate fibers (Table 4), they presumably
constitute the more indispensable part of this
tether.
Egg sac attachment in lycosids versus
trechaleids and rhoicinines.—In contrast to
Pardosa females that use both MaA and MiA
fibers (1
8
and 2
8
) for attaching the egg sac, for
a total of eight ampullate fibers, Carico (1993)
reports that trechaleid females use only MiA
fibers (but, again, both 1
8
and 2
8
; see his fig.
4) for this purpose, for a total of four ampul-
late fibers. And while 2
8
ampullate fibers are
considerably wider than 1
8
ampullate fibers in
233TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Pardosa and Trochosa (at least on the ALS
of the latter, Table 4), the 1
8
and 2
8
MiA fibers
of Hesydrus, shown in fig. 4 of Carico (1993),
do not appear to differ substantially in diam-
eter. Carico (1993:230) describes these tre-
chaleid MiA fibers as ‘strong’ and this seems
apt considering that our measurements of 2
8
ampullate fiber diameters in Pardosa,Trocho-
sa, and Dolomedes were in the range of about
2.3–4.4
m
m, whereas the MiA fibers in Cari-
co’s fig. 4 have diameters of about 9–12
m
m.
Thus, what the trechaleid tether lacks in num-
ber of fibers is perhaps more than made up for
in strength per fiber. Additional comparisons
between these two families are made in the
appropriate sections below.
In contrast to lycosids and trechaleids, it
has been reported that Rhoicinus and Rhoici-
naria (the latter currently placed in Amauro-
biidae, see Platnick 2002) carry their egg sacs
attached to the posterior spinnerets; i.e., the
PLS (see Exline 1950, 1960). If true, attach-
ment is presumably accomplished using fibers
from aciniform and/or cylindrical glands, rath-
er than ampullate gland fibers.
Ampullate glands in lycosids versus ar-
aneoids.—To us, the female lycosid’s (or tre-
chaleid’s) use of 2
8
ampullate silk is most in-
teresting when compared with the araneoid
condition. The evidence to date indicates that
2
8
ampullate glands produce silk in araneoids
only during proecdyses (Townley et al. 1993)
and, therefore, these glands are not needed
and not functional in adults of either sex.
What occurs in adult female lycosids (and
adult females from some other families, Ta-
bles 2, 3) appears to be a variation on this
scheme. As in araneoids, the 2
8
ampullate
glands of juvenile lycosids apparently produce
silk during proecdyses. This is indicated by
the presence of ampullate tartipores in second
instar through adult lycosids and is consistent
with the replacement of 2
8
ampullate spigots
by 2
8
ampullate nubbins in adult males (Table
3; Figs. 1, 12–15). In araneoids, the final molt
differs from the preceding molts, with regard
to the ampullate glands, in that the blocked 2
8
ampullate glands present in the last juvenile
instar (see Table 1) remain blocked and do not
re-develop in the adult. And because the open
2
8
ampullate glands present in the last juvenile
instar become blocked and regress in the
adult, as they do after each molt, the adult
contains two sets of blocked 2
8
ampullate
glands, rather than one blocked set and one
open set (as in juveniles). From our observa-
tions on spinnerets, we infer that this differ-
ence between the final molt and all preceding
molts does not exist in female lycosids
(among others). That is, the blocked 2
8
am-
pullate glands present in a penultimate instar
female lycosid do become open 2
8
ampullate
glands in the adult and re-develop accordingly
(Fig. 1). Presumably, they are not functional
immediately after the last ecdysis, requiring at
least a day or two to re-develop and accu-
mulate luminal contents, but would thereafter
be able to produce silk fibers concurrently
with the 1
8
ampullate glands.
On the other hand, it would seem that the
2
8
ampullate glands of female lycosids (Par-
dosa at least) are not completely unresponsive
to the hormonal changes that culminate in an
adult being produced, as evidenced by the en-
larged 2
8
ampullate spigots of adults (e.g.,
Figs. 9, 11). And given the greater diameters
of 2
8
ampullate fibers in adult females (rela-
tive to the 1
8
ampullate fibers), there may also
be internal changes to the 2
8
ampullate glands,
such as substantial increases in the calibers of
their ducts, that occur at the same time.
Other functions of 2
8
ampullate gland fi-
bers.—Though we have only observed 2
8
am-
pullate fibers being used for egg sac attach-
ment, we are not suggesting that this is the
only role the 2
8
ampullate glands play in the
adult female. Instead, it may be that when
these spiders are not carrying egg sacs, fibers
from these glands are used for other purposes.
One possibility is that they contribute to the
dragline. Such an application of 2
8
ampullate
silk may be especially significant for species
in which adult female draglines stimulate
courtship behaviors in adult males (reviewed
in Tietjen & Rovner 1982; also, e.g., Stratton
& Uetz 1983; Lizotte & Rovner 1989; Hebets
et al. 1996; Ferna´ndez-Montraveta & Ruano-
Bellido 2000). As in egg sac attachment, the
greater diameter of these fibers might also
make them better suited to this role than the
1
8
ampullate fibers. Mechanically, the 2
8
am-
pullate fibers would present a more substantial
trail that might be more easily discerned by
males and, from a chemosensory perspective,
the greater surface area of a 2
8
ampullate fiber
has greater potential for presenting phero-
mones to males. Considering that 2
8
ampullate
spigots (implying functional 2
8
ampullate
234 THE JOURNAL OF ARACHNOLOGY
Figures 38–40.—Spinnerets with attached egg sac in adult female Pardosa modica: 38. 1
8
and 2
8
MaA
fibers from both ALS and 1
8
and 2
8
MiA fibers from both PMS are attached to the egg sac; 39. Left PMS
from the same preparation, showing more clearly the emergence of fibers from the MiA spigots, as well
as 1
8
and 2
8
MaA fibers from the left ALS in the upper left corner; 40. Right ALS from the same
preparation, showing the emergence of fibers from the MaA spigots. The MaA fibers from this ALS are
attached to the egg sac by a well-defined (and undamaged) attachment cone. Note that the 2
8
ampullate
fibers have considerably greater diameters than the 1
8
ampullate fibers. The PLS are out of the field of
view in Fig. 38 (no fibers were observed coming from the PLS). Scale bars (38)
5
100
m
m; (39, 40)
5
50
m
m.
235TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 41–42.—Ampullate fibers used for egg sac attachment in Pardosa littoralis (posterior at right,
lateral at top): 41. Portion of left ALS showing 1
8
and 2
8
MaA fibers; 42. Portion of left PMS showing
1
8
and 2
8
MiA fibers. Both micrographs were taken after the egg sac was removed from the spinnerets.
In preparing the spider for SEM (before the egg sac was removed), the shafts (*) of both MaA spigots
and the 2
8
MiA spigot became detached from the bases. Scale bars
5
25
m
m.
glands) are present in adult females belonging
to several other families in which egg sacs are
not carried on the spinnerets (Tables 2, 3), and
assuming fibers from these spigots play a use-
ful role(s) in such females, it would seem like-
ly that adult female lycosids use 2
8
ampullate
fibers for purposes in addition to egg sac
transport. On the other hand, the greater ma-
terial and energetic cost of producing the larg-
er-diameter 2
8
ampullate silk may limit its use
in other applications.
Comparative ampullate gland spigot
morphology.—After observing the relatively
large 2
8
ampullate spigots of adult female ly-
cosids, we were curious to know if this feature
is unique to lycosids, or perhaps to lycosids,
trechaleids, and rhoicinines. Such a limited
occurrence would more strongly suggest that
the mode of egg sac transport used by these
spiders may have been made possible by or
facilitated by selection for enlarged 2
8
ampul-
late spigots from which relatively large-di-
ameter 2
8
ampullate fibers are drawn. And in
this context, might this feature extend to pi-
saurids as well? Several authors have noted
that adult female pisaurids transport their egg
sacs, held under the sternum, using not only
their chelicerae and palps, but also silk from
the spinnerets (e.g., Le´caillon 1905:137; Bish-
op 1924:27–28; Nielsen 1932:133, 135; Bris-
towe 1958:187, 190; Dondale & Redner 1990:
322; Carico 1976:63, 1993:235–236), and
Carico (1993) has suggested that egg sac
transport using the spinnerets is plesiomorphic
for lycosids, trechaleids, and pisaurids. This
raises the possibility that ampullate silk may
also play a role, albeit a less crucial one, in
egg sac transport in the Pisauridae. On the
other hand, Roberts (1995:236) has ‘‘. . . never
seen any threads running between the sac and
the spinners ...’’in pisaurids, which points
up the desirability of investigating the extent
and nature of silk use in egg sac transport
within this family. Or are relatively large 2
8
ampullate spigots unrelated to egg sac trans-
port, with such spigots routinely encountered
among those entelegynes that retain 2
8
am-
pullate spigots as adults, yet do not use am-
pullate silk for egg sac transport? Questions
such as these prompted us to begin examining,
as opportunities have arisen, spinnerets of
such non-lycosid entelegynes.
At present, our survey is very limited (Ta-
ble 3) and needs to be expanded, including
examining more lycosids and other lycosoids.
Thus, we lack satisfactory answers to the
above questions. As detailed below, from
what data we have and from observations re-
ported in the literature, it seems that 2
8
am-
pullate spigots are generally not larger than 1
8
236 THE JOURNAL OF ARACHNOLOGY
Figure 43.—Portion of left ALS containing the
MaA spigots (posterior at right, lateral at top),
showing MaA fibers that were being used for egg
sac attachment in Trochosa ruricola. The micro-
graph was taken after the egg sac was removed
from the spinnerets. As in Fig. 41, the shafts (*) of
both MaA spigots became detached from the bases
while processing the specimen for SEM. Scale bar
5
20
m
m.
ampullate spigots among entelegynes. But on
the other hand, larger 2
8
ampullate spigots are
neither restricted to lycosoids, nor are they
present in all lycosoids that use spinnerets,
solely or in part, to carry the egg sac. Among
the examined non-lycosoids in this study
(Figs. 28–37, Table 3), 2
8
ampullate spigots
tended to be smaller or about the same size as
1
8
ampullate spigots.
This description also applies to the MaA
spigots of several amaurobioids (sensu Gris-
wold et al. 1999). Davies (1998b:74) has not-
ed that in Tasmarubrius (Amaurobiidae) the
ALS of an adult female has two MaA spigots,
with the anterior one, i.e. the 1
8
MaA spigot,
larger. Wang (2000) provides micrographs of
ALS from several adult female amaurobiids,
including Rubrius and Callobius in which the
two MaA spigots appear similar in size. A mi-
crograph of a Coelotes right ALS in the same
paper (Wang 2000:fig. 4) indicates that the 2
8
MaA spigot is larger than the 1
8
MaA spigot,
but because the position of the MaA tartipore
in the Callobius figure indicates the right
ALS, rather than the left ALS as stated in the
caption, it may be that the figured Coelotes
spinneret is actually the left ALS, in which
case the 1
8
MaA spigot is larger. In a descrip-
tion of the amaurobioid subfamily Kababini-
nae, Davies & Lambkin (2000a) state that the
two MaA spigots on the female ALS are of
unequal size. From their fig. 5C of a right
ALS in Malarina, it appears that the 1
8
MaA
spigot is again larger. And in several amphi-
nectids, 1
8
MaA spigots are larger than 2
8
MaA spigots in adult females. Davies (1998a),
in describing a Quemusia species, states that
the anterior MaA spigot (i.e., the 1
8
) is ‘‘much
larger’’ than the posterior MaA spigot (2
8
),
and, in both a Magua species and a Buyina
species, reports that the anterior MaA spigot
is larger than the posterior one. 1
8
and 2
8
MaA
spigots of similar size can be seen on an ALS
of an adult female Liocranoides (Tengellidae)
in fig. 3 of Platnick (1999).
In contrast, several published micrographs
demonstrate that larger 2
8
ampullate spigots,
even if not prevalent, are not unique to lyco-
soids. In fig. 11 of Harvey (1995), an ALS
from the nicodamid Ambicodamus is shown
on which the 2
8
MaA spigot is conspicuously
wider than the 1
8
MaA spigot. Larger 2
8
am-
pullate spigots also occur in some lamponids
(Platnick 2000: Lamponina ALS, figs. 287 &
288; Lamponella ALS, figs. 354 & 355; fe-
male Centrothele PMS, figs. 408 & 409).
Among the pisaurids examined we saw ex-
amples of 2
8
ampullate spigots that were
smaller than, or about equal in size to, 1
8
am-
pullate spigots (Pisaurina, Figs. 26, 27), as
well as examples of 2
8
ampullate spigots that
were noticeably larger than their 1
8
counter-
parts (some, though not all, Dolomedes, Figs.
24, 25). If enlarged 2
8
ampullate spigots in
lycosids are part of an adaptation facilitating
egg sac transport using the spinnerets, then
these mixed observations may be a reflection
of the supplemental role, at most, that ampul-
late silk plays in pisaurid egg sac transport. If
ampullate silk is not used by pisaurids for egg
sac transport, then examples of larger 2
8
am-
pullate spigots in pisaurids might simply re-
flect phylogenetic relatedness between lycos-
237TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
ids and pisaurids (e.g., Dondale 1986;
Griswold 1993; Silva Davila in press).
Given our observations on spinnerets from
lycosids and pisaurids, and considering that
data obtained thus far suggest the Trechaleidae
as sister group to the Lycosidae (Sierwald
1990b, 1993; Griswold 1993) or to Lycosidae
1
Pisauridae (Silva Davila in press), we
would have expected 2
8
ampullate spigots to
be larger than 1
8
ampullate spigots in trechal-
eids (at least with the MiA spigots since these
are used for egg sac attachment (Carico
1993)). But in Carico’s (1993) fig. 4 of a Hes-
ydrus PMS the 2
8
MiA spigot (to the right of
the 1
8
MiA spigot in this figure, above the
MiA tartipore) does not appear to be larger
than the 1
8
MiA spigot. It may, in fact, be
smaller. As mentioned above, the fibers
emerging from these spigots also do not differ
conspicuously in diameter. But both fibers are
very wide, relative to those that we have mea-
sured in Pardosa,Trochosa, and Dolomedes,
and both spigots are large relative to the acin-
iform and cylindrical spigots that surround
them. There is the possibility, therefore, that
trechaleids (Hesydrus at least) do have en-
larged 2
8
MiA spigots, but that it is not im-
mediately obvious because the 1
8
MiA spigots
are also enlarged. Since only MiA fibers are
used by trechaleids for egg sac attachment
(Carico 1993), it would be of value to exam-
ine the MaA spigots to see if they and the silk
fibers they produce are noticeably smaller
than their MiA counterparts. The larger di-
ameters of trechaleid MiA fibers, compared
with lycosid and pisaurid ampullate fibers,
lead us to speculate that this difference may
be related to a behavioral difference among
the three families. If an egg sac becomes de-
tached, lycosid and pisaurid females will re-
attach it, while trechaleid females will not
(Carico et al. 1985; Carico 1993). Nor, inci-
dentally, do Shinobius (Rhoicininae) females
reattach a detached egg sac (Kaihotsu 1988:
17). Do the larger-diameter trechaleid fibers
make detachment and, thus, the need for re-
attachment less likely than among lycosids
and pisaurids? The possibility was raised
above that the greater cost to a spider of pro-
ducing larger-diameter fibers may result in
more restricted use of such fibers. Thus, do
adult female trechaleids use MiA silk exclu-
sively or primarily for egg sac attachment?
Again, additional observations are clearly
needed.
The only scans of rhoicinine spinnerets we
have seen are those presented in the descrip-
tion of Heidrunea (Brescovit & Ho¨fer 1994).
In fig. 6c of Brescovit & Ho¨fer (1994), both
PMS of an adult female are shown. We ten-
tatively identify the tartipore, present on each
PMS and located just posterior to the three
most anterior spigots, as a MiA tartipore. The
two spigots immediately posterior to this tar-
tipore are presumably the MiA spigots, with
the 2
8
MiA spigot and MiA tartipore juxta-
posed. If these identifications are correct, then
the bases of the 1
8
MiA spigots are wider than
those of the 2
8
MiA spigots. The significance
of this observation to any correspondence be-
tween the size of 2
8
ampullate spigots and the
use of ampullate silk in egg sac transport, is
unknown at present since we do not know if
Heidrunea attach their egg sacs to their spin-
nerets and, if they do, if ampullate gland fibers
are used. Recall that Rhoicinus have been re-
ported to attach egg sacs to the PLS (Exline
1950, 1960), indicating that ampullate gland
silks are not involved.
As an aside, we note that amaurobiids of
the genera Amaurobius and Callobius are not
included in Table 2 even though Hajer’s
(1990) observations indicate that they con-
form to the description given in the table leg-
end. This is because others have reported only
a single MiA spigot on each PMS in adult
females of these two genera (Platnick et al.
1991:62, 64; Griswold et al. 1999; Wang
2000; the latter contains SEM scans of their
PMS), and our own observations on a single
specimen of an adult female Callobius ben-
netti (Blackwall 1846) coincide with these lat-
er reports. Thus, the 2
8
ampullate spigot sex-
ual dimorphism considered in Table 2 seems
to apply to the 2
8
MaA spigots only. Such is
the situation observed in the amaurobiid ge-
nus Tasmarubrius (Davies 1998b) and in the
amaurobioid subfamily Kababininae (Davies
1999; Davies & Lambkin 2000a, b). Even this
more limited sexual dimorphism is absent in
some genera currently included (some very
tentatively) in the Amaurobiidae (Platnick
2002) given that adult females of Storenoso-
ma,Otira,Midgee,Manjala,Malala (Davies
1999; Davies & Lambkin 2001), and Retiro
(Griswold et al. 1999) have only a single MaA
238 THE JOURNAL OF ARACHNOLOGY
Figures 44–47.—Attachment cones that affix ampullate fibers to the surface of the egg sac: 44, 45.
From a Gladicosa gulosa egg sac, with the boxed area in Fig. 44 shown at higher magnification in Fig.
45; 46, 47. From a Trochosa ruricola egg sac, with the boxed area in Fig. 46 shown at higher magnification
in Fig. 47. On the G. gulosa egg sac, it appeared that all the ampullate fibers were attached by this one
cone, while on the T. ruricola egg sac, one 1
8
/2
8
pair of ampullate fibers was attached by a separate, more
poorly formed or damaged cone. In Figs. 45, 47 note the fusion among many of the fibers that constitute
the cones. Scale bars (44, 46)
5
100
m
m; (45, 47)
5
5
m
m.
239TOWNLEY & TILLINGHAST—EGG SAC ATTACHMENT IN LYCOSIDS
Figures 48–49.—Surface of a Pardosa moesta egg sac, at the site where the eight ampullate fibers
coming from the ALS and PMS are attached. 48. Each 2
8
ampullate fiber (the four thickest, most obvious
fibers in the micrograph) is attached by a separate attachment cone, one of which is not well-defined or
is damaged. Only two of the 1
8
ampullate fibers are affixed in the same cone as their 2
8
counterpart. An
arrow points to the site where one 1
8
ampullate fiber is affixed by its own very small cone. 49. The six
more closely spaced ampullate fibers from Fig. 48 are shown at higher magnification so that the 1
8
fiber
accompanying each 2
8
fiber can be seen more clearly. Scale bars (48)
5
25
m
m; (49)
5
10
m
m.
240 THE JOURNAL OF ARACHNOLOGY
spigot on each ALS, accompanied by a MaA
nubbin.
Ampullate fiber attachment to the egg
sac.—We have not made a specific attempt to
determine the glandular origin of the principal
fibers that form the cone-like structures by
which ampullate fibers are affixed to the sur-
face of the egg sac. Casual observations from
SEM micrographs suggest the piriform glands
as the most likely candidates. Among females
with egg sacs, the only fibers we have ob-
served emerging from spigots are ampullate
and piriform fibers. Also, fusion among fibers
seen on cones is reminiscent of the fusion that
has been described among piriform fibers
from webs of C. citricola (Peters 1993). It
must be acknowledged, however, that fusion
has also been observed among other fiber
types, including aciniform-A fibers from at
least some uloborids (Peters & Kovoor 1989)
and cylindrical fibers from A. aurantia
(Stubbs 1991; Stubbs et al. 1992; Foradori et
al. 2002). A role in attaching ampullate fibers
to the egg sac surface is consistent with the
piriform glands’ well known function of pro-
ducing attachment discs that secure ampullate
fibers to various substrates (e.g., Apstein
1889; Warburton 1890; Richter et al. 1971).
From descriptions of trechaleid egg sacs in
Sierwald (1990a:8–9; 1993:62), Brescovit et
al. (2000:14), and especially Carico (1993:
230, 236), and from figs. 5–6 in the latter pa-
per, it appears that a single attachment cone
secures the four MiA fibers to the surface of
the egg sac in at least several genera within
this family. A single cone may also affix the
eight ampullate fibers to the egg sac in some
lycosids (e.g., Gladicosa), but others (Pardo-
sa) typically produce several closely spaced
cones that serve to attach these ampullate fi-
bers.
ACKNOWLEDGMENTS
Partial financial support for this work was
contributed by Joel Tillinghast. Matthew For-
adori (Zoology Department, UNH) and Mes-
bah Creitz collected some of the spiders used
in this study, Margaret Tillinghast made trans-
lations from the French, Tomoko Kakazu
made a translation from the Japanese, and Jo-
seph Danahy and Douglas Prince (UNH Com-
puting and Information Services) helped ready
the figures for publication. Nancy Cherim
(UNH EM Facility) was extremely accom-
modating in making much time on the SEM
available to us. Figure 1 was produced at the
suggestion of Dr. Jonathan Coddington (Na-
tional Museum of Natural History, Smithson-
ian Institution) with additional design sugges-
tions made by Dr. Gail Stratton (University of
Mississippi) and Charlene Newton. Drs. Strat-
ton and Coddington and an anonymous re-
viewer made a number of recommendations
that greatly improved the text. Dr. Petra Sier-
wald (Field Museum of Chicago) identified
and determined the sex of juvenile P. mira
examined in this study and Dr. Charles Don-
dale (Eastern Cereal & Oilseed Research Cen-
tre, Agriculture Canada) identified the exam-
ined Coras individuals. Diana Silva Davila
kindly sent us a copy of her in press manu-
script. To all these individuals we are indebted
and very grateful.
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Manuscript received 14 April 2002, revised 26 Oc-
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