Table Of Content2005. The Journal of Arachnology 33:482^89
A FOSSIL HARVESTMAN (ARACHNIDA, OPILIONES)
FROM THE MISSISSIPPIAN OF
EAST KIRKTON, SCOTLAND
Jason A* Dunlop: Institut fur Systematische Zoologie, Museum fur Naturkunde der
Humboldt-Universitat zu Berlin, InvalidenstraBe 43, D^lOllS Berlin, Germany.
@
E-=mail: jason.dunlop museum.hu-berlin.de
Lyall I. Anderson: Department of Natural Sciences, National Museums of Scotland,
Chambers Street, Edinburgh, EHl IJF, EFnited Kingdom
ABSTRACT. A fossil harvestman (Arachnida, Opiliones) from the Mississippian (Visean: Brigantian)
of East Kirkton, Scotland is described as Brigantibunum Ustoni new genus and species. At ca. 340 Ma,
it represents the second oldest record of Opiliones. Although some details are lacking, this long-legged,
small-bodied and rather gracile harvestman is surprisingly modern-looking and appears to show the im-
pression of an annulate ovipositor. Its leg anatomy closely matches that of some living Eupnoi and it is
tentatively referred to this clade. Like the newly discovered Rhynie chert harvestmen, it reinforces the
idea that modern, crown-group Opiliones can be traced back to at least the mid-Paleozoic.
Keywords: Taxonomy, new species, Visean, Paleozoic, Eupnoi
Fossil harvestmen (Opiliones) are very rare. harvestmen having a very similar gross mor-
Although they are best known from Tertiary phology to living animals. The second oldest
ambers (e.g., Cokendolpher & Poinar 1998; harvestman is also rather modern looking and
Stargga 2002; Dunlop & Giribet 2003), there comes from the Mississippian (ca. 340 Ma) of
are also two Mesozoic records (Roger 1946; East Kirkton in Scotland (Wood et al. 1985).
Jell & Duncan 1986) neither of which have The East Kirkton harvestman (Figs. 1-3) is
been formally named and the first of which is preserved in a more typical fashion as a flat-
dubious. Until recently the earliestharvestmen tened impression and superficially resembles
were Pennsylvanian (ca. 300 Ma) fossils from living 'daddy long-legs’ forms. In general,
the Coal Measures of Commentry in France Mississippian arachnid fossils are far less
(Thevenin 1901) and Mazon Creek in the common than either Devonian or Pennsylva-
USA (Petrunkevitch 1913). Restudy of Pe- nian examples (see Dunlop & RoBler 2003 for
trunkevitch's (1913) putative extinct order a review), thus any record from this time pe-
Kustarachnida has shown that these Mazon riod is significant.
Creek arachnids are misidentified harvestmen The East Kirkton harvestman was briefly
(Beall 1986, 1997; Dunlop 2004a). Additional mentioned, with a figure, in the original sum-
Pennsylvanian-aged harvestmen have been mary paper dealing with the locality (Wood et
collected from Missouri, USA (Dunlop al. 1985), It has also been noted or listed in
2004b) and Poland (Maciek Kania, pers. some further publications (Smithson 1989;
comm., 2004). Selden 1993a, b; Clack 1998; Jerarn 2001;
The oldest recorded harvestmen are excep- Dunlop & RoBler 2003; Dunlop 2004a). Here
tionally preserved, three-dimensional fossils we formally describe and name this important
from the Early Devonian (ca. 400 Ma) Rhynie specimen and discuss its significance in the
cherts of Scotland (Dunlop et al. 2003, 2004). light ofother recent fossil harvestman discov-
Provisionally assigned to the Eupnoi clade as eries.
Eophalangium sheari Dunlop et al. 2004, this METHODS
exquisite, silicified material displays details of
internal structures such as genitalia and tra- The East Kirkton harvestman was borrowed
chea, all of which point towards these ancient on research loan from the Hunterian Museum,
482
DUNLOP & ANDERSON—MISSISSIPPIAN HARVESTMAN 483
Glasgow, United Kingdom (GLAHM A2854), comm. 2002). Rolfe et al. (1994) identifiedthe
where it is customarily on display. The spec- combined thickness of bed 81 and bed 82
imen was digitally photographed using a Lei- (with a lateral variation in thickness between
ca DC100 digital camera mounted on a Leica 80 and 140 mm) as the same unit which Gei-
MZFLIII microscope and drawn using a cam- kie (1861) named ‘bed bk This unit contains
era iucida attachment. Adobe Photoshop Lim- dense accumulations of ostracods on some
ited Edition 5.0 was used to manipulate the bedding surfaces as well as charcoalified plant
digital images. For comparative purposes the material. This is also the likely source bed of
type material ofNemastomoides longipes (Pe- the holotype of Westlothiana Uzziae. Durant
trunkevitch 1913) from the Peabody Museum, (1994) summarized the likely paleoeeviron-
M
Yale (YPM 171), depressus (Petrunkevitch ment in which the sediments exposed in the
1913) from the United States National Mu- East Kirkton Quarry were deposited in a shal-
seum (USNM 37974) and Kustarachne ten- low lake close to the flanks of an active vol-
uipes Scudder 1890 (USNM 37967) was ex- cano. Volcanic eruption products including
amined, along with Eophaiangium shear! ash and tuffs, were eroded and washed down
from Rhynie (held in the University of Mun- into the lake. Set against a backdrop of active
ster, Germany) and extant species preserved volcanism in the area in which it formed, it is
in the Museum fur Natu—rkunde Berlin. no surprise that ashy bands, pyroclastic frag-
Geological setting. The East Kirkton ments and chemical sediments influenced by
Limestone is a fossil Konservat-Lagerstatte hot spring activity dominate the sequence
located near Bathgate, West Lothian, Scot- hosting both terrestrial and aquatic plant and
land; about 27 km west of Edinburgh. The animal fossils. Widespread development of
limestone is the lowest of five which occur limestone formed from stromatolitic algae
within the Bathgate Hills Volcanic Formation. also point to an unusual physio-chemical en-
This in turn can be correlated with the lower vironment for the formation of this deposit.
part of the Brigantian Stage of the Visean Se- In the fossil’s plane of compression at least
ries of the Mississippian (= Lower Carbon- four elongate and articulated legs are pre-
iferous in European terminology). The Bri- served. A thin calcite vein cross-cuts the legs
gantian Stage spans the time interval 336.0- of the fossil (Figs. 1, 2). Close study of the
339.4 Ma. Further details can be found in harvestman body suggests that other legs may
Rolfe et al. (1994). The East Kirkton locality have broken off prior to preservation. How-
has also yielded scorpions (Jeram 1994, ever, it is obvious that such delicate structures,
2001), myriapods (Shear 1994) and a variety which easily break off in extant animals even
i of terrestrial tetrapods including the anthra- while still alive, indicate only a short period
I cosaurid Silvanerpeton miripedes and the fa- of transport into the preservational environ-
mous stem-ameiote Westlothiana Uzziae; ment. Perhaps this animal was rafted out onto
however early insects and arachnid groups the open lake on floating vegetation before
! like the extinct order Trigoeotarbida, which dropping into the water?
are usually fairly common in terrestrial de-
MORPHOLOGICAL INTERPRETATION
posits of this age, are so far absent. —
Preservation.—GLAHM A2854 is pre- Body. The body (Figs. 1-3) is small and
served as a compression fossil on the surface rounded. We suspect it is essentially a dorso-
ofathin, grey bed ofcalcareous tuff. The light lateral to ventro-lateral compression in which
red-brown coloration of the harvestman is the two sides of the body have become su-
analogous to that of the scorpion fossils perimposed. As is typical for harvestmen, the
etched from the East Kirkton limestone (Jer- prosoma and opisthosoma are broadly joined.
am 1994) and suggests that some constituent Features such as eyes ormouthparts are equiv-
ofthe original cuticle still remains, ratherthan ocal, even under higher magnification, but the
it being a wholly carbonized cuticle. This in body does come to a slight, angular point on
turn suggests that chitieoclastic bacteria were the dorsal side where an eye tubercle might
excluded from the preservatioeaJ environment be expected. Low-angle lighting reveals lines
during the formation of the deposit and the on the harvestman body which might corre-
arthropod fossils contained therein. The fossil spond to the original opisthosomal segmen-
originated from bed 82 (S.R Wood pers. tation and/or elements of the coxae which
484 THE JOURNAL OF ARACHNOLOGY
—
Figure 1. Brigantibunum listoni new genus and species, from the Mississippian (Visean: Brigantian)
ofEast Kirkton, Scotland. This modern-looking specimen is here tentatively assigned to the Eupnoi clade.
Scale bar = 5 mm.
come up the side ofthe body in many living Shultz 2000) and may only be betrayed by
harvestmen. These lines do not form a clear folds or color patterns on the body surface.
pattern running the length of the body and The fossil also reveals fine tuberculation on
in similar-looking extant harvestmen seg- the body where cuticle has been preserved
mentation is generally poorly dehned (e.g.. and, while there is some degree of locali-
DUNLOP & ANDERSON—MISSISSIPPIAN HARVESTMAN 485
zation, the tubercles do not define identifi- lowed by a very short patella which is slightly
able segments.— thicker than the adjacent podomeres. It forms
OTipositor.- ^Oee intriguing feature is an a distinct and bulbous 'kneek The tibia is
apparently annulate structure overlaying the again elongate and slender, widening distally
left side of the body, which is only clearly to form a disjunct articulation with the next
visible under high magnification and low an- podomere,. the basitarsus. This basitarsus is
gle lighting (Fig. 3). It is associated with orig- also long and slender, although the transition
inal cuticle which tends to suggest it is not an to the telotarsus is indistinct. In many living
artefact, and at least towards the 'dorsaF end harvestmen the telotarsus is composed of
there are rows of tiny circular structures many short tarsomeres. These cannot be re-
across each aenulation which might be tuber- solved in the fossil, but the distal curvature of
cles or setal sockets. We tentatively interpret at least one of the legs (leg 1 in our scheme,
&
this structure as an ovipositor since its annu- see Figs. 1 2) implies that it, too, was
late morphology and proportions relative to formed from numerous articulating elements.
the body are suggestive of that in a modern Claws at the ends of the legs (the apotele) are
eupnoid harvestman (e.g., Shultz 2000). The equivocal.
East Kirktoe scorpions preserve respiratory SYSTEMATIC PALAEONTOLOGY
organs (Jeram 1994), which shows that there
is the potential at this locality to recover in- ?Eupeoi Hansen & Sprensee 1904
ternal features. Ifthis is an ovipositor, it is the —
second oldestrecord ofinternal genitalia, after Remarks, As noted by Selden (1993a),
the one recovered in Eophalangium sheari the East Kirkton fossil is clearly aharvestman,
from Rhynie (Dunlop et al. 2003, 2004). It but explicit diagnostic characters of higher
implies that the East Kirkton fossil is afemale taxa within Opiliones are not clearly pre-
which probably laid its eggs in the substrate. served. Nevertheless, its overall morphology
mm
In living harvestmen the ovipositor is appar- with a 4 globular body and long, essen-
ently extended through hemolymph pressure tially homogeneous legs is wholly inconsis-
(Shultz 2000). Perhaps the ovipositor in the tent with Cyphophthalmi, which are tiny (typ-
fossil was squeezed out of the body during ically 1“2 mm) with short, stubby legs. Nor
compression and came to lie across the opis- does it resemble the more robust Laniatores
thosoma? in which leg 4 is often enlarged and spiny
Legs.—-Four almost complete legs are pre- (although not, for example, in oecopodids)
served, all ofwhich are very long and gracile; and in which prominent, raptorial pedipalps
up to ca. twelve times the length of the body. would be expected. This leaves the Palpatores
A small fragment of either a fifth leg or, per- group, the monophyly ofwhich is currently in
haps, a pedipalp is also preserved. It is diffi- dispute (compare Shultz 1998 and Giribet et
cult to assign legs unequivocally to their se- al. 2002). The older Rhynie chert harvestmen
quence in the body, but the longest leg (which preserve convincing eupnoid characters (Dun-
is also the most gracile) is probably leg 2. lop et al. 2003, 2004a), thus both Eupnoi and
This leg is longest in most living (non-cy- its putative sister taxon lineage, Dyspnoi sen-
phophthalmid) harvestman and has a more su Shultz (1998) or (Dyspnoi + Laniatores)
tactile function. Indeed all 4 preserved legs sensu Giribet et al. (2002), can be predicted
express slightly different femur lengths (see from the Mississippiae.
Systematics) and this might indicate that the Both the Eupnoi and Dyspnoi clades, which
fossil is essentially a lateral view preserving make up the traditional Palpatores group, in-
legs 1--4 on one side of the body; with the clude long-legged taxa. The extremely long
GLAHM
corresponding legs from the other side either and gracile legs in A2854, which are
A
missing or still within the matrix. tentative up to about twelve times body length, tend to
numbering scheme is proposed based on this favor affinities with phalaegiid or scleroso-
assumption (Fig. 2). matid harvestmen (Eupnoi), for example
Discrete podomeres can be recognized, and members of common eupnoid genera like
the basic leg anatomy is a precise match for Leiobunum C.L. Koch 1839, Opilio Herbst
living eupnoid harvestmen (cf. Shultz 2000), 1798 and Phalangium Linnaeus 1758. Among
The femur is elongate and slender. It is fol- the Recent Dyspnoi, common genera such as
486 THE JOURNAL OF ARACHNOLOGY
—
Figure 2. Interpretative drawing of the specimen shown in Fig. 1. Abbreviations: bs = basitarsus, fe
= femur, pp = possibile pedipalp, pt = patella, ti = tibia, tt = telotarsus. Legs tentatively numberedfrom
1 to 4, with leg 2 longest. Scale bar = 5 mm.
I
Nemastoma C.L. Koch 1836 and Dicranolas- phthalmi (rejected for the reasons outlined
ma S0rensen 1873 typically have legs which above) and Eupnoi (Shultz 1998, 2000; Giri-
are somewhat shorter in relation to body bet et al. 2002). However, explicit autapo-
length. Members of the dyspnoid genus Mi- morphies of Eupnoi cannot be resolved un-
tostoma Roewer 1951 are closer to GLAHM equivocally in the fossil. The lakeside
A2854 in terms of leg lengths. Data from the paleoenvironment is unlikely to have trapped
fairly widespread M. chrysomelas (Hermann either high mountain and/or cave animals. The
1804) in Martens (1978, p. 143) suggests that long and gracile legs in the fossil are more
leg length (13.7 mm) is, at best, about eight characteristic for certain phalangiid and scle-
times body length (1.7 mm), although in a rosomatid eupnoids, as opposed to the usual
highly-specialized Alpine cave species likeM. range in non-specialist dyspnoids. On these
anopthalmum (Page 1946) leg length (22.8 grounds we tentatively assign the East Kirkton
mm) can be over fourteen times body length fossil to Eupnoi.
(1.6 mm); data from Martens (1978, p. 149). Brigantibunum new genus
Clearly leg length is not an ideal characterand —
as noted by Martens, these parameters can Type and only species. Brigantibunum
vary even within a species and between males listoni new gen—us and species.
and females. Etymology. From the Brigantian age of
The putative ovipositor also hints at a eup- the fossil and the suffix “bunum” used in
noid. If our interpretation is correct, this an- modern genera of small bodied, long-legged
nulate morphology only occurs in Cypho- harvestmen such as Leiobunum.
DUNLOP & ANDERSON—MISSISSIPPIAN HARVESTMAN 487
—
Figure 3. Detail ofthe body region under low angle lighting and immersion in alcohol. Abbreviations:
ov? = possible annulate ovipositor, tb = tuberculation of cuticle. Leg sequence as in Fig. 2. Scale bar =
mm.
1
—
Diagnosis. Extremely gracile Paleozoic Kirkton fossil belongs to Eophalangium Dun-
harvestman with long, slender legs up to lop et al., 2004.
twelve times the length of the small, ovate Among the Pennsylvanian opilionids, our
body. Femora at least two and a half to three fossil is clearly not congeneric with the pu-
times the len—gth of the body. tative Commentry troguloid Eotrogulusfayoli
Remarks. The Eophalangium sheari ma- Thevenin 1901, which is a robust animal with
terial from the Devonian of Rhynie, Aber- an elongate body and comparatively short
deenshire, Scotland includes a long-legged legs. This leaves three species ofKustarachne
i specimen. Given the very different modes of Scudder 1890, two ofwhich are rather incom-
preservation, direct comparisons with the East plete and doubtful, and three species of Ne-
Kirkton fossil are problematic. The three-di- matostomoides Thevenin 1901; of which N.
mensional Rhynie material yields many char- depressus from Mazon Creek is a misidenti-
acters not testable in the East Kirkton speci- fied phalangiotarbid (Beall 1997; pers. obs.).
men. Furthermore, the (male) Rhynie fossil The East Kirkton fossil appears longer-legged
associated with the long legs is incomplete than all these Pennsylvanian forms, although
and the full extent of both the legs and body the Mazon Creek specimens are in nodules
remains equivocal. Further discoveries may and the full extent of the legs is, of course,
change this interpretation, but we can find no not preserved. In detail, the length of the fe-
explicit autapomorphies or even reliable ratios mur offers a potential diagnostic characterand
of body proportions to argue that the East the femora in the East Kirkton fossil are pro-
488 THE JOURNAL OF ARACHNOLOGY
portionately longer (ca. 3 times body length) mm, leg 3: 6 mm, leg 4: 9 mm. Basitarsi
than the femora in Nemastomoides and Kus- lengths as follows. Leg 1: 8 mm, leg 2: un- !j
tarachne (ca. 1-2 times body length). Overall, clear, leg 3: 8 mm, leg 4: 9 mm. Telotarsi
GLAHM A2854 is a unique find and the only incomplete, but long and slender. i
|
record of a Mississippian harvestman. It most DISCUSSION
closely resembles the Commentry species N.
elaveris Thevenin 1901, but based on its ex- Brigantibunum listoni fits into a developing
treme leg length and gracile appearance we pattern (e.g., Dunlop et al. 2003, 2004; Dun- i
assign it to a new genus diagnosed on the lop 2004a, b) in which harvestmen appear to
body-femur ratio. have evolved relatively early into recogniz-
able crown-group forms (i.e. animals assign-
Brigantibunum listoni new species able to clades with Recent representatives)
Figs. 1, 2. and exhibit a high degree of stasis, with little
?Earliest known harvestman (Arachnida, Opili- fundamental change over hundreds ofmillions
ones): Wood et al. 1985: 355-356, fig. 1. of years. Harvestmen with the same basic
Opilionid or harvestman: Smithson 1989: 676-678; shape as the East Kirkton fossil, remain com-
Selden 1993a: 392-393; Selden 1993b: 305-306; mon and abundant today; particularly in the
Clack 1998: 66-69; Jeram 2001: 374, tabs. 16.1, northern hemisphere. To put this into context,
16.2; Dunlop & RoBler 2003: 389; Dunlop & Gi- the oldest crown-group spiders (Selden 1996)
ribet 2003: 371; Dunlop 2004a: 24; 2004b: 67.
are mesotheles (the most basal living spider
—
Type. Holotype, from the East Kirkton clade) and are first recorded from the end of
Quarry, near Bathgate, (27 km west of Edin- the Pennsylvanian. This is some 100 million
burgh), West Lothian, Scotland (Grid refer- years after the oldest crown-group harvestmen
ence NS 991690), collected by Mr. Stan P. from Rhynie, which can be assigned to the
Wood, derived from Unit 82 ofthe East Kirk- eupnoids; a somewhat derived clade. It is also
ton Limestone, West Lothian Oil-Shale For- worth mentioning that in recent arachnid phy-
,
mation, Strathclyde Group, Upper Visean logenies (e.g. Giribet et al. 2002) harvestmen
(Brigantian), Mississipian (= Lower Carbon- seem to resolve in a fairly basal position
iferous in European stratigraphy) (GLAHM (compared to spiders) as part of the so-called '
A2854). Dromopoda clade along with scorpions, pseu-
—
Etymology. ForJeffListon (University of doscoipions and solifuges.
Glasgow & Hunterian Museum). ACKNOWLEDGMENTS
—
Diagnosis. —As for the genus. We thank Neil Clark and Jeff Liston !
mmDescription. Body small, rounded, ca. 4 (GLAHM), Tim White (YPM) and MarkFlor-
in diameter. Red-brown cuticle includes ence (USNM) for access to material in their
fine tuberculation and possible segment/coxal care, Stan Wood for information on the type
boundaries. Elongate, annulate structure
(?ovipositor), length 2.5 mm, curves across locality and Maciek Kania for information on
new Polish material. Gonzalo Giribet, Bill
body on left side. Small, incomplete limb el-
ement (?pedipalp), length 2 mm, projects from Shear and the editors provided helpful com- i
ments on the manuscript during review.
body on right side. Four legs preserved, all
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