Remarks on the new world pseudoscorpion genera Parawithius and Victorwithius, with a new genus bearing a remarkable sternal modification (Pseudoscorpiones, Withiidae)

2004·0.76 MB·

2004. The Journal of Arachnology 32:436–456
REMARKS ON THE NEW WORLD PSEUDOSCORPION
GENERA PARAWITHIUS AND VICTORWITHIUS, WITH
A NEW GENUS BEARING A REMARKABLE STERNAL
MODIFICATION (PSEUDOSCORPIONES, WITHIIDAE)
Mark S. Harvey: Department of Terrestrial Invertebrates, Western Australian
Museum, Francis St, Perth, Western Australia 6000, Australia. E-mail:
[email protected]
ABSTRACT. The withiid genus Parawithius Chamberlin is rediagnosed, based upon a detailed exam-
ination of the type species, P. nobilis (With) from Colombia, which is redescribed and illustrated. The
subgenus Parawithius (Victorwithius) Feio is returned to full genericlevel. Cystowithius,anewgenusof
Withiidae, is described from high elevation localities in central and South America for four species:
Cystowithiussmithersinewspecies(typespecies)fromEcuador,C.ecuadoricus(Beier)newcombination,
fromEcuadorandPeru,C.colombicusnewspeciesfromColombia,andC.chamberlininewspeciesfrom
Mexico and Guatemala. Males of Cystowithius are highly unusual due to the presence of sternal invagi-
nations, a feature that has not been previously observed in pseudoscorpions. PopulationsofCystowithius
smithersi occur within the leaf sheaths of the giant rosette plant Espeletia pycnophylla var. angelensis
(Compositae).
Keywords: Taxonomy, new species, new genus, Colombia, Ecuador, Peru, Mexico, Guatemala, Espe-
letia, high elevation, Cystowithius
The pseudoscorpion family Withiidae is volcano 15 km north of Tulca´n, in northern
distributed in most parts of the world with 34 Ecuador, Smithers et al. (2001) found many
genera and 153 species currently recognized pseudoscorpion specimens inhabiting the
(Harvey1991).Thefamilyisdividedintotwo spaces formed between the sheaths. Detailed
subfamilies, Withiinae Chamberlin and Para- examination of the specimens showed that
goniochernetinae Beier, but the relationships they represented a single species of Withiidae
both within and between these groups are with a peculiar modification to the male ster-
poorly known. The South American withiid nitesinwhichtheanterior-lateraledgeofster-
fauna consists of seven endemic genera (Har- nites V–VIII were invaginated to form pock-
vey 1991) which are currently placed in three ets within the inter-segmental membrane.
tribes: Cacodemonius Chamberlin in the Ca- Subsequent examination of museum collec-
codemoniini, Protowithius Beier 1955 in the tions revealed that while such a structure is
Protowithiini, and Balanowithius Beier 1959, rare among the Withiidae, further American
Dolichowithius Chamberlin 1931, Neowithius specimens were found with such invagina-
Beier 1932, Parawithius Chamberlin 1931 tions.Thefeaturewasfoundtobeabsentfrom
and Tropidowithius Beier 1955 in the Withi- otherwithiid generaand Ihereproposeanew
ini.Inaddition,thecosmopolitansynanthropic genus, here named Cystowithius, for these
species. In addition, the type species of Par-
species Withius piger (Simon 1879) has also
awithius, P. nobilis (With 1908), is rede-
been recorded from Chile. The composition
scribed based upon the male holotype and a
and relationships of these genera are only
second male specimen collected from a near-
poorlyknownandconsiderableproblemsexist
by locality, and the status of the species cur-
inthesystematicsofthewithiines,evenatthe
rently assigned to the subgenus Parawithius
generic level.
(Victorwithius) Feio 1944 is reassessed.
During a recent survey of the fauna asso-
ciatedwiththeleafsheathsofthegiantrosette METHODS
plantEspeletiapycnophyllavar.angelensison Thespecimensthatformedthebasisforthis
thewesternslopesofVolca´nChiles,anextinct study are lodged in the American Museum of
436
HARVEY—NEWWORLDWITHIIDAE 437
NaturalHistory,NewYork(AMNH),theNat- lateral tergalcrests;patchesofglandularsetae
ural History Museum, London (BMNH), the only present on sternites VII-IX of males;
CaliforniaAcademyofSciences,SanFrancis- males without sternal invaginations; and ter-
co (CAS), Museum Victoria, Melbourne gites without discrete the lateral fields char-
(NMV), Naturhistorisches Museum, Wien acteristic of Cacodemonius.
(NHMW), Museum National d’Histoire Na- Remarks.—The genus Parawithius com-
turelle, Paris (MNHN),PontificiaUniversidad prises two distinct groups which were treated
Cato´lica del Ecuador, Quito (PUCE), Bohart as subgenera by Beier (1959), Parawithius
Museum of Entomology, University of Cali- (Parawithius) and Parawithius (Victorwithius)
fornia, Davis (UCD), Western AustralianMu- Feio 1944, which differ in the position of tri-
seum, Perth (WAM) and Museum fu¨r Natur- chobothriaisbandit,whicharenearlyadjacent
kunde der Humboldt-Universita¨t, Berlin to each other in the nominate subgenus but
(ZMB). Terminology largely follows Cham- whicharesomewhatseparatedinallspeciesof
berlin (1931a) and Harvey (1992). In partic- Victorwithius.ThesubgenusParawithius(Par-
ular, it should be noted that the terminology awithius)currentlycontainsthetypespeciesP.
for the trichobothria used by Harvey (1992)
(P.) nobilis (With 1908) from Colombia, (P.
differs slightly from that used by other work-
(P.) nobilis nobilis), and Ecuador and Peru (P.
ers.TheratioTSisthedistancefromthebase
(P.) nobilis ecuadoricus Beier 1959), and P.
of tarsus IV to the tactile seta, divided by the
(P.) iunctus Beier 1932b and P. (P.) pseudo-
lengthoftheentiretarsus.Thetergalandster-
rufus Beier 1932b fromParaguay.Parawithius
nal chaetotaxies (including those of the glan-
(Victorwithius) consists of 12 species, P. (V).
dular setae) refer to the entire segment, and
coniger Mahnert 1979, P. (V). fiebrigi Beier
notmerelyhalf-segmentsasisoftenpresented
1932b, P. (V). gracilimanus Mahnert 1979, P.
in descriptions of some cheliferoid pseudo-
(V). incognitus Beier 1959, P. (V). mimulus
scorpion taxa. All measurements are in mm
(Beier 1954), P. (V). monoplacophorus Feio
and were made with an ocular micrometer on
1944, P. (V). proximus Ellingsen 1905, P. (V).
acompoundoradissectingmicroscope.Spec-
rufeolusBeier1959,P.(V).rufusBalzan1887,
imens were examined by either clearing in
P. (V). schlingeri Beier 1959, P. (V). similis
50% lactic acid or by permanently mounting
Beier 1959 and P. (V). venezuelanus Beier
inEuparalonmicroscopeslides.Thescanning
1932b.Theremovalofallofthosespeciespre-
electron micrographs were obtained in a Phil-
viously placed in Parawithius (Victorwithius)
ips XL30 scanning electron microscope after
to thefullgenus Victorwithius(seebelow)and
the specimens were prepared by dehydration
the transfer of P. nobilis ecuadoricus to the
in 1,1,1,3,3,3-Hexmethyldisilazane (HMDS),
new genus Cystowithius (see below) leaves
air-drying and mounting on SEM stubs with
only P. nobilis from Colombia, and P. iunctus
carbon tape.
and P. pseudorufus from Paraguay remaining
SYSTEMATICS inthegenusParawithius.Ihaveexaminedma-
terialofallthreeofthesespecies,includingthe
FAMILY WITHIIDAE CHAMBERLIN
type specimens of each and have confirmed
1931
that P. nobilis and P. pseudorufus are notcon-
SUBFAMILY WITHIINAE CHAMBERLIN
specificwithanyofthespeciesofCystowithius
1931
describedelsewhereinthispaper,andthatthey
Genus Parawithius Chamberlin 1931
lack the diagnostic features of Cystowithius.
Parawithius Chamberlin 1931b: 292; Beier 1932a: Beier (1932b) stated that the holotype of P.
212; Beier 1959: 216; Harvey 1991: 650. iunctus (ZMB 29667) was a male. In fact, the
Typespecies.—ChelifernobilisWith1908, specimen is a female, as indicated on the de-
by original designation. terminationlabelassociatedwiththespecimen.
Diagnosis.—ThegenusParawithiusishere It is currently not possible to easily separate
defined by thefollowing combinationofchar- isolated female specimens of Parawithius and
acters:tactilesetaoftarsusIVsub-distal(rath- Cystowithius,soitisnotcertaintowhichgenus
er than sub-medial); carapace broadest poste- thefemaleholotypeof P.iunctusbelongs.Itis
riorly; trichobothria est, isb and it nearly here retained in Parawithius until the Para-
adjacent to each other; male tergites without guayan withiid fauna becomes better known.
438 THE JOURNALOF ARACHNOLOGY
Figures1–5.—Parawithiusnobilis(With),holotype(cid:63)unlessstatedotherwise:1.Rightpedipalp,dorsal;
2. Right chela, lateral; 3. Right tarsus IV, (cid:63) from 12 miles E. of Bogota´, Colombia;4. Carapace,dorsal;
5. Posterior sternites, ventral. Trichobothrial abbreviationsfollowChamberlin(1931)andHarvey(1992).
Parawithius nobilis (With 1908) Diagnosis.—Parawithius nobilis differs
Figs. 1–6 from P. iunctus and P. pseudorufus by the
Chelifer nobilis With 1908: 234–236, figs 4a–b. presence of pale spots on the carapaceal me-
Parawithius nobilis (With): Chamberlin 1931b: tazone, by the slightly stouter pedipalpal seg-
292;Beier1932a:213;Roewer1937:309;Beier ments (e.g. patella 3.24–3.30 times longer
1959: 216–217, fig. 25; Muchmore 1993: 99; than broad, compared with 3.4–3.6 timeslon-
Judson 1997: 30. ger than broad), and the morphology of the
Parawithius (Parawithius) nobilis (With): Harvey setae on the interior face of the pedipalpal
1991: 650.
segments, which are only slightly clavate in
Material examined.—Holotype male, Bo- P.nobilisandclearlyclavateinP.iunctusand
gota´, Cundinamarca, COLOMBIA [4(cid:56)36(cid:57)N, P. pseudorufus.
74(cid:56)05(cid:57)W], Keyserling collection (BMNH). Description.—Adult males: Color general-
Other material: Cundinamarca: COLOM- ly light red-brown; carapaceal metazone with
BIA 1 (cid:63), 12 miles E. of Bogota´ [ca. 4(cid:56)36(cid:57)N, largepairedpaleareas;tergitesI–IIIwithpale
73(cid:56)52(cid:57)W], 3 March 1955, E.I. Schlinger, E.S. areas on each tergal half, tergites IV–IX with
Ross (CAS). pale areas on each tergal half situated medi-
HARVEY—NEWWORLDWITHIIDAE 439
ally thus forming a distinct medialband. Ped- times longer than broad; tarsal tactile seta of
ipalp (Fig. 1): all segments granulate, except leg IV situated sub-distally, ca. 0.74 of tarsus
for chelal fingers, which are smooth; dorsal length; subterminal tarsal setae arcuate and
setae generally strongly foliate; trochanter acute; arolium slightly shorter than claws.
1.69–1.75, femur 4.22–4.25, patella 3.24– Dimensions (mm), males: Body length
3.30, chela (with pedicel) 3.60–3.71, chela 2.32–2.64. Pedipalps: trochanter0.422–0.435/
(without pedicel) 3.39–3.47, hand 1.73–1.78 0.248–0.250, femur 0.883–0.896/0.209–
timeslongerthanbroad,movablefinger0.99– 0.211,patella0.762–0.782/0.235–0.237,chela
1.04 times longer than hand. Fixed chelal fin- (with pedicel) 1.196–1.229/0.331–0.332, che-
gerwith8trichobothria,movablechelalfinger la (without pedicel) 1.125–1.150, hand length
with 4 trichobothria (Fig. 2): eb and esb sit- 0.567–0.591, movable finger length 0.588–
uated basally; est, it and isb grouped together 0.592. Chelicera 0.237/0.122, movable finger
sub-medially;istandibsituatedbasally;band length 0.180. Carapace 0.832–0.848/0.501–
sb situated near one another; st slightly closer 0.656 (width at medial area); eye diameter
to sb than to t. Venom apparatus present in 0.045–0.105. Leg I: femur 0.179/0.157, patel-
both chelal fingers, venom ducts long, termi-
la0.378/0.160,tibia0.352/0.105,tarsus0.340/
nating in nodus ramosus near est in fixed fin- 0.069. Leg IV: femur (cid:49) patella 0.662/0.224,
ger and near t in movable finger. Chelal teeth
tibia 0.511/0.128, tarsus 0.396/0.080, TS
strongly triangular, slightly retrorse; fixed fin-
0.294.
ger with 35 teeth; movable finger with 34
Remarks.—Parawithius nobilis has been
teeth;accessoryteethabsent.Chelicerawith5
infrequently recorded in theliterature,andthe
setae on hand, bs and sbs dentate; movable
only known specimens are the male holotype
finger with 1 submedial seta; galea short with
from Bogota´ (With 1908), a male specimen
4 small terminal rami; flagellum of 4 blades;
from ‘New Granada’ (now Colombia) (With
serrulaexteriorwith14–17blades;laminaex-
1908), a malefrom nearBogota´ (Beier1959),
terior present. Carapace (Fig. 4) 1.27–1.69
and six adultsfrom LagoZurucuchu,Ecuador
timeslongerthanbroad;lateralmarginseven-
and a female from Huamachuco, Peru (Beier
ly convex, but posteriorly widened; with 2
1959). The specimens from Lago Zurucuchu
non-corneate eyes, those of holotype appar-
and Huamachuco were regarded as a separate
ently smaller than those other male; with ca.
subspecies by Beier (1959) who named them
52 setae,including4nearanteriormarginand
Parawithius nobilis ecuadoricus Beier, but
7–8 near posterior margin; with 2 deep fur-
which are here treated as a distinct species of
rows, the posterior furrow slightly closer to
posterior carapaceal margin than to median Cystowithius (see below). The holotype of
furrow. Tergites with distinct medial suture, Chelifer nobilis lodged in the BMNH is in
sternitesbarelydivided.Chaetotaxyoftergites good condition, but a fine covering of minute
I-XII: 9–10: 9–10: 11: 12–15: 14–17: 15–16: crystals has obscured the specimen in some
15–17: 17–18: 17: 14: 6: 2; mostly uniseriate places,inparticularthecoxalregion,thuspre-
but some tergites with a few setae placed an- cluding the determination of the setal formula
teriorly; all setae foliate. Chaetotaxy of ster- of the coxae. The description presentedabove
nites I-XII: 12: (2)10(2): (2)11(2):14–16: 16: is based upon the male holotype and the male
16: 15–16: 11–12: 9–10: 9–11: 2; sternites from near Bogota´ lodged in CAS. Both spec-
VII-IX with patches of glandular setae, ar- imens are very similar in all morphological
ranged 26–41: ca. 63–77: 3–5 respectively; features including the size and shape of the
setae uniseriate and acuminate, except for pedipalpal segments and in the number of
smaller setae on sternite XI which are dentic- glandular setae on the male sternites. I have
ulate; glandular setae small and conical in not examined the specimen lodged in the
shape; (cid:63) without paired invaginations on an- BMNH from New Granada that was referred
terior margins of sternites. Coxal chaetotaxy to by With (1908).
12: 10: 10: 18; pedipalpal coxa with 2 apical
setae and very small sub-oral seta. Internal Genus Victorwithius Feio 1944
genitalia not observed. Legs: junction be- Victorwithius Feio 1944: 1–3.
tweenfemoraandpatellaeIandIIonlyslight- Cacodemoniellus Beier 1954: 326–327 (synony-
ly oblique; femur (cid:49) patella of leg IV 3.00 mized by Beier 1959: 216).
440 THE JOURNALOF ARACHNOLOGY
Parawithius (Victorwithius) Feio: Beier1959:216; Genus Cystowithius new genus
Harvey 1991: 651.
Typespecies.—Cystowithiussmithersinew
Type species.—Of Victorwithius: Victor- species.
withius monoplacophorus Feio 1944, by orig- Etymology.—The generic epithet refers to
inal designation; of Cacodemoniellus: Caco- the cyst-like invaginations that diagnose the
demoniellus mimulus Beier 1954, by original genus (kystis, Greek for bladder, sac or cell),
designation. combined with the generic stem Withius. The
Diagnosis.—SpeciesofVictorwithiusdiffer name was first used for this taxon by the late
from other South American withiids by the J.C. Chamberlin, who labelled the Mexican
following combination of characters: tactile specimens described below as ‘‘Cystowithius
seta of tarsus IV situated sub-distally; tricho- gen. nov.’’ The gender is masculine.
bothrium it situated mid-way between the tip Diagnosis.—Males of Cystowithius are un-
of the finger and isb; and tergites withoutdis- like those of any other pseudoscorpion in the
crete the lateral fields characteristic of Caco- possession of paired sac-like invaginationson
demonius. the anterior margins of sternites V-VIII (C.
Remarks.—The subgenus Victorwithius is smithersi and C. ecuadoricus) or VI-VII (C.
adistinctandeasilyrecognizabletaxonthatis colombicus) or VI-VIII (C. chamberlini)
here returned to full generic rank. It differs (Figs. 18, 19).
from Parawithius in the position of tricho- Description.—Most dorsal setae strongly
bothrium it which is situated mid-way be- denticulate; setae on sternites acicular. Fixed
chelal finger with 8 trichobothria, movable
tween the tip of the finger and isb in Victor-
chelal finger with 4 trichobothria; trichoboth-
withius but is situated close to isb in
ria est, it and isb grouped together sub-medi-
Parawithius. The distinction between Victor-
ally. Venom apparatus present in both chelal
withius and some other South American gen-
fingers, venom duct terminating in nodus ra-
era such as Balanowithius is difficult to con-
mosus near est in fixed finger and near t in
firm, and further research on the relationships
movable finger. Chelal teeth all closely
of these genera is necessary to establish their
spaced;accessoryteethabsent.Chelicerawith
interrelationships and hence the limits of the
5setaeonhand,bsandsbsdenticulate,others
genera. The following species are placed in
acuminate; movable finger with 1 subdistal
the genus Victorwithius:
seta; flagellum of 4 blades, the most distal
blade with several serrations on leading edge,
Victorwithiusconiger(Mahnert1979)NEWCOM- other blades smooth; lamina exterior present.
BINATION
Carapace with 2 non-corneate eyes; with 2
Victorwithius fiebrigi (Beier 1932b) NEW COM-
distinct furrows, posterior furrow closer to
BINATION
posterior margin of carapace than to anterior
Victorwithius gracilimanus (Mahnert 1979) NEW
furrow. Tergites I–X with medial suture, ster-
COMBINATION
nites IV–X with faint medial suture. Males
Victorwithius incognitus(Beier1959)NEWCOM-
BINATION with paired sac-like invaginations on anterior
Victorwithius mimulus (Beier 1954) NEW COM- margins of sternites V–VIII or VI–VII or VI–
BINATION VIII;maleswithpatchesofglandularsetaeon
Victorwithius monoplacophorus Feio 1944 sternites VII–X, females with 2 glandular se-
Victorwithius proximus (Ellingsen 1905) NEW tae per segment on segments VII–IX (C.
COMBINATION smithersi)orVIII–IX(C.ecuadoricus,C.col-
Victorwithius rufeolus (Beier 1959) NEW COM-
ombicus and C. chamberlini); glandular setae
BINATION
small and conical in shape. Male genitalia
Victorwithius rufus (Balzan 1887) NEW COMBI-
with lateral apodeme long and triangular. Fe-
NATION
male genitalia with single oval median crib-
Victorwithius schlingeri (Beier 1959) NEW COM-
riform plate and 2 smaller lateral cribriform
BINATION
plates; with poorly defined membranous sper-
Victorwithius similis (Beier 1959) NEW COMBI-
NATION mathecal sac irregularly covered with pores;
Victorwithius venezuelanus (Beier 1932b) NEW and with 2 lateral hooked apodemes. Pleural
COMBINATION membrane longitudinally striate. Posterior
HARVEY—NEWWORLDWITHIIDAE 441
maxillary lyrifissure present and sub-distally nert 1988 (Mahnert 1988), Rexwithiusgirardi
situated. Spiracular helix present. Legs: junc- Heurtault 1993 (Heurtault 1993), Trichotow-
tionbetweenfemoraandpatellaeIandIIonly ithius abyssinicus Beier 1944 (Dashdamirov
slightly oblique; subterminal tarsal setae ar- 1992) and possibly in Victorwithiusmonopla-
cuate and acute; arolium slightly shorter than cophorus Feio 1944 (Feio 1944). An alterna-
claws; claws slender and simple. tivemorphologyinwhichthelateralapodeme
Remarks.—The presence of sternal invag- and ejaculatory canal is not lengthened pos-
inations in Cystowithius is unique within the teriorly occurs in some species of Withius in-
Pseudoscorpiones. This feature has been ob- cluding the type species W. piger (Simon
servedinallmaleshereassignedtothegenus, 1878) (Heurtault 1971) and in severalAfrican
but was found to be absent in other withiids species (Mahnert 1988), as well as in Aisthe-
(Table 1). The structure appears to be auta- towithius rossi Beier 1967 (Mahnert 1988),
pomorphic for the four species described be- Girardwithius pumilus Heurtault 1993 (Heur-
low. While the feature is easily seen in slide- tault 1993) and Ectromachernes lamottei Va-
mounted specimens (either as permanent chon 1952 (Vachon 1952).Althoughitispos-
mounts or as temporary mountsinglycerolor sible that those species in which the lateral
lactic acid) using a compound microscope, it apodeme is triangular may represent a mono-
is still visible in material examined in ethanol phyleticgroup,itshouldbenotedthatthevast
using a dissecting microscope, as the antero- majority of withiids have yet to have their
lateral margins of the sternite are slightly male genitalia described in detail, thus pre-
curved and somewhat darkerthantheremain- cludinganycomprehensivestatementsregard-
ing sternal cuticle. It was not observed using ing the utility of this feature.
a scanning electron microscope as the slight AlthoughthespeciesdescribedhereinCys-
contraction of the specimen during the dehy- towithius clearly form a monophyletic group,
dration process hides from view the invagi- the disposition of the tactile seta of leg IV
nation (Fig. 22). varies considerably in its position. The tactile
The conformation of the male genitalia of seta is sub-distal in C. smithersi (TS (cid:53) 0.75–
Cystowithius spp. resembles that of someoth- 0.77), C. ecuadoricus (TS (cid:53) 0.74–0.75) and
er withiids as they possess an elongate, tri- C. colombicus (TS (cid:53) 0.77–0.79) but is only
angular lateral apodeme that bears an extend- slightly distad of medial in C. chamberlini
ed ejaculatory canal (Fig. 21). This feature (TS (cid:53) 0.59–0.61). This variation may dimin-
also occurs in several species of Withius, in- ish the usefulness of this character in distin-
cludingW.hispanus(L.Koch1873),W.faun- guishing between different genera of Withi-
us (Simon 1879), W. neglectus (Simon 1878) idae.
(Heurtault 1971), Metawithius yurii (Redikor- SpeciesofCystowithiusaregenerallyfound
zev1938)(Harvey1988),M.murrayi(Pocock at high altitudes with three species occurring
1900) (pers. obs.), M. philippinus Beier 1937 above 3,000 m in the northern Andes in Co-
(pers. obs.), M. spiniventer Redikorzev 1938 lombia,EcuadorandPeru,andabove2,000m
(pers. obs.), Pycnowithius cavernicola Mah- in Mexico (Fig. 7).
Key to species of Cystowithius
1. Chelal hand smooth; setae on chelal hand only barely denticulate; tactile seta of tarsus IV
situated sub-medially (TS (cid:53) 0.59–0.61); males with sternal invaginations on sternites VI-
VIII ................................................. C. chamberlini new species
Chelal hand evenly granulate; setae on chelal hand distinctly denticulate; tactile seta of
tarsus IV situated sub-distally (TS (cid:53) 0.74–0.79); males with sternal invaginations on ster-
nites V-VIII ............................................................... 2
2(1). Larger species, e.g. chela (with pedicel) greater than 1.35 mm (Fig. 8) ..........
...................................................... C. smithersi new species
Smaller species, e.g. chela (with pedicel) less than 1.20 mm (Fig. 8) ............... 3
3(2). Setae on tergite XI long and only slightly clavate (Fig. 27) ....... C. ecuadoricus (Beier)
Setae on tergite XI short and strongly clavate (Fig. 32) ....... C. colombicus new species
442 THE JOURNALOF ARACHNOLOGY
Table1.—Withiidaeinwhichthemalesternalinvaginationsareabsent.Asteriskdenotesthetypespecies
of the genus.
Specimens
Taxon examined Museum Locality Remarks
Balanowithius egregius holotype (cid:63), para- CAS Pichilingue, Ecua- Personal observa-
Beier 1959* type (cid:47) dor tion
Cacodemonius satanas Dr M. Judson (in
(With 1908) litt.)
Cacodemonius seg- Dr M. Judson (in
mentidentatus (Bal- litt.)
zan 1887)
Dolichowithius argen- holotype (cid:63) CAS Salta, Argentina Personal observa-
tinus Beier 1959 tion
Dolichowithius exten- 1 (cid:63), 1 (cid:47) syn- BMNH La Moka, Vene- Personal observa-
sus Beier 1932b types zuela tion
Dolichowithius granu- 1 paratype (cid:63) AMNH Guyana Personal observa-
losus Hoff, 1945 tion
Ectromachernesmira- holotype (cid:63) BMNH Jemme Forest, Personal observa-
bilis Beier 1944* Ethiopia tion
Hyperwithiusdawydof- 4 (cid:63) syntypes NHMW Cao Nguyeˆn Laˆm Personal observa-
fi Beier 1951 Vieˆn (Plateau tion
von Langbian),
Laˆm Dong, Vi-
etnam
Hyperwithiustonkinen- 1 (cid:63), 1 (cid:47), 2 proto- NHMW Lau Chau, Lau Personal observa-
sis Beier 1951 nymphs syn- Chau, Vietnam tion
types
Juxtacheliferfructuo- paratypes (cid:63), (cid:47) AMNH Santa Fe, New Personal observa-
sus Hoff 1956* and tritonymphs Mexico, U.S.A. tion
Metawithius murrayi 1 (cid:63), 1 (cid:47) syntypes BMNH North West Point, Personal observa-
(Pocock 1900) Christmas Is- tion
land, Australia
Metawithius philippi- several (cid:63), (cid:47) syn- ZMB Philippines Personal observa-
nus Beier 1937 types tion
Metawithius spiniven- several (cid:63), (cid:47) NHMW Cao Nguyeˆn Laˆm Personal observa-
ter Redikorzev 1938 specimens Vieˆn (Plateau tion
von Langbian),
Laˆm Dong, Vi-
etnam
Metawithius (Micro- 1 (cid:63) NMV Sertung Island, In- Personal observa-
withius) yurii (Redi- donesia tion
korzev 1938)
Metawithius (Micro- 2 (cid:63), 1 (cid:47) syn- NHMW Telom Valley, Personal observa-
withius) tweediei types Cameron High- tion
Beier 1955 lands, Pahang,
Malaysia
Nesowithius seychelle- 2 (cid:63) syntypes BMNH Praslin, Seychelles Personal observa-
sensis Beier 1940* tion
New genus, similar to several (cid:63), (cid:47) and WAM Manngarre Rain- Personal observa-
Metawithius Cham- tritonymphs forest, Kakadu tion
berlin 1931 National Park,
Northern Terri-
tory, Australia
Parawithius nobilis holotype (cid:63) BMNH Bogota´, Colombia Personal observa-
(With 1908)* tion
Parawithius nobilis 1 (cid:63) CAS near Bogota´, Co- Personal observa-
(With 1908)* lombia tion
HARVEY—NEWWORLDWITHIIDAE 443
Table 1.—Continued.
Specimens
Taxon examined Museum Locality Remarks
Parawithius pseudoru- holotype (cid:63) ZMB 29668 Paraguay Personal observa-
fus Beier 1932b tion
Scotowithius helenae Dr M. Judson (in
Beier 1977* litt.)
Stenowithius angulatus 1 (cid:63) CAS (JC- no data Personal observa-
(Ellingsen 1906) 906.01001) tion
Stenowithius bayoni 7 adult syntypes BMNH Kampala, Uganda Personal observa-
(Ellingsen 1910) of Stenowithius 1932.11.12.14– tion
ugandanus 20
Beier 1932b
Stenowithius duffeyi 11 adult syntypes BMNH Boatswain-birdIs- Personal observa-
Beier 1961 1964.8.17.2–12 land, Ascension tion
Island
Thaumatowithius tibia- 1 (cid:63), 2 (cid:47) syn- BMNH Le Pouce, Mauri- Personal observa-
lis Beier 1940* types tius tion
Trichotowithius abys- 1 (cid:63), 3 (cid:47) syn- BMNH Jemme Forest, Personal observa-
sinicus Beier 1944* types Ethiopia tion
Victorwithius coniger Dr M. Judson (in
(Mahnert 1979) litt.)
Victorwithius rufeolus 2 (cid:63) and (cid:47) syn- CAS Pichilingue, Ecua- Personal observa-
(Beier 1959) types dor tion
Victorwithius schlin- holotype (cid:63) CAS Tingo Mar´ıa, Personal observa-
geri (Beier 1959) Monson Valley, tion
Peru
Victorwithius similis 1 (cid:63) and 1 (cid:47) syn- CAS 10 miles N. of Personal observa-
(Beier 1959) types Trancas, Argen- tion
tina
Victorwithius venezue- 1 (cid:63) and 2 (cid:47) syn- BMNH La Moka, Vene- Personal observa-
lanus (Beier 1932b) types zuela tion and Dr M.
Judson (in litt.)
Withius hispanus (L. 2 (cid:63) CAS (JC- Sardinia Personal observa-
Koch 1873) 511.04004–5) tion
Withius piger (Simon numerous speci- WAM, CAS Australia,Hawaii, Personal observa-
1878)* mens Europe,North tion
America,South
America,India,
etc.
Cystowithius smithersi new species tritonymph, same data as holotype (BMNH);
Figs. 7–23 1 male, 1 female, 1 tritonymph, same data as
holotype (PUCE); 1 male, 1 female, 1 trito-
Material examined.—Holotype male,Vol- nymph, same data as holotype (MHNG); 1
ca´n Chiles, ca. 15 km N. of Tulca´n, Carchi male, 1 female, 1 tritonymph, same data as
Province, ECUADOR, 0(cid:56)48(cid:57)35.0(cid:48)N, holotype (MNHN).
77(cid:56)57(cid:57)15.0(cid:48)W, altitude 3,600 m, August1997, Etymology.—The specific epithet honors
in leaf sheath of Espeletia pycnophylla, P. Peter Smithers, University of Plymouth, who
Smithers, A. Bond, M. Burne (WAM provided me with the opportunity to examine
T54598). Paratypes: ECUADOR: Carchi the specimens described here, and who first
Province: 3 males, 3 females (1 with brood- published upon the unusual biology of this
sac), 5 tritonymphs, same data as holotype species (Smithers et al. 2001).
(WAM T54599–54606); 1 male, 1 female, 1 Diagnosis.—Cystowithius smithersi differs
444 THE JOURNALOF ARACHNOLOGY
Figures 6–7.—Maps showing known distributions of species mentioned in the text: 6. Parawithius
nobilis; 7. Cystowithius species.
from the other three species of the genus by brown; carapaceal metazone with paired pale
its longerandmoreslenderpedipalps,e.g.the spots; tergites with paler areas, usually sur-
chela (with pedicel) of C. smithersi is greater rounding setal areoles. Pedipalp (Fig. 16): all
than 1.35 mm and the pedipalpal femur is segments granulate, except for chelal fingers,
greater than 0.90 mm. In addition, it differs which are smooth; dorsal setae generally
from C. chamberlini in the evenly granulate strongly foliate; femur 4.93–5.43 ((cid:63)), 4.70–
chelal hand and the presence of male sternal 5.28 ((cid:47)), patella 4.04–4.45 ((cid:63)), 3.83–4.19
invaginations on sternites V–VIII. ((cid:47)),chela(withpedicel)4.24–4.59((cid:63)),3.88–
Description.—Adults: Color dark red- 4.18 ((cid:47)), chela (without pedicel) 4.04–4.35
Figure 8.—Graph depicting pedipalpal chela (with pedicel) length versus pedipalpal femur length in
Cystowithius species.
HARVEY—NEWWORLDWITHIIDAE 445
anterior margin and 10 near posterior margin;
with 2 deep furrows, the posterior furrow
slightly closer to posterior carapaceal margin
than to median furrow. Tergites with distinct
medial suture, sternites barely divided. Chae-
totaxyoftergitesI–XII:holotype(cid:63),10:9:10:
13: 15: 15: 17: 18: 17: 13: 9 (including 2 tac-
tile setae): 2; paratype (cid:47), 12: 10: 10: 17: 17:
16: 18: 18: 19: 16: 8: 2; mostly uniseriate but
some tergites with a few setae placed anteri-
orly; all setae foliate. Chaetotaxy of sternites
I–XII: holotype (cid:63), 13: (1)12(1): (2)12(2):16:
19:18[gls]:16[gls]:11[gls]:9:12:2;paratype
(cid:47), 15: (1)17(1): (2)12(2): 17: 20: 17[gls]:
17[gls]: 17[gls]: 14: 10: 2; sternites VII–IX
withpatchesofglandularsetae[gls],arranged
(cid:63), ca. 75: 72: 60 respectively; (cid:47), 2: 2: 2, re-
spectively;setaeuniseriateandacuminate,ex-
cept for lateral setae on sternite XI which are
finely denticulate; glandular setae small and
conical in shape; (cid:63) with paired invaginations
on anterior margins of sternites V–VIII (Figs.
18, 19). Coxal chaetotaxy: paratype (cid:63), 10: 9:
10: 18; paratype (cid:47), 14: 13:12:22;pedipalpal
coxa with 2 apical setae and very small sub-
oralseta.Internalgenitaliaofmalewithlateral
Figure 9.—Cystowithius smithersi new species, apodeme long and triangular (Fig. 21); of fe-
holotype male.
male with single oval median cribriformplate
and 2 smaller lateral cribriform plates, and
((cid:63)), 3.71–4.00 ((cid:47)), hand 1.98–2.07 ((cid:63)), with poorly defined membranous spermathe-
1.74–1.95 ((cid:47)) times longer than broad, mov- cal sac irregularly covered with pores; with 2
able finger 1.04–1.15 ((cid:63)), 1.00–1.22 ((cid:47)) lateralhookedapodemes(Fig.20).Legs(Figs.
times longer than hand. Fixed chelal finger 13, 14): junction between femoraandpatellae
with 8 trichobothria, movable chelal finger I and IIonly slightly oblique;femur(cid:49)patella
with 4 trichobothria (Fig. 10): eb and esb sit- oflegIV3.80((cid:63)),4.36((cid:47))timeslongerthan
uated basally; est, it and isb grouped together broad;tarsaltactilesetaoflegIVsituatedsub-
sub-medially;istandibsituatedbasally;band distally, ca. 0.77 ((cid:63)), 0.75 ((cid:47)) of tarsus
sb situated near one another; st mid-way be- length; subterminal tarsal setae arcuate and
tween sb and t. Venom apparatus present in acute; arolium slightly shorter than claws.
both chelal fingers, venom ducts long, termi- Dimensions (mm), males (females): Body
nating in nodus ramosus near est in fixed fin- length 2.40–2.75 (2.44–3.71). Pedipalps: tro-
ger and near t in movable finger. Chelal teeth chanter 0.484–0.493/0.222–0.232 (0.464–
triangular, slightly retrorse; fixed finger with 0.515/0.224–0.246), femur 0.982–1.072/
42 ((cid:63)), 43 ((cid:47)) teeth; movable finger with 49 0.186–0.200 (0.925–1.104/0.193–0.215),
((cid:63)),52((cid:47))teeth;accessoryteethabsent.Che- patella 0.864–0.908/0.198–0.224 (0.789–
licera: with 5 setae on hand, all acuminate; 0.960/0.206–0.229), chela (with pedicel)
movable finger with 1 subdistal seta; galea 1.392–1.440/0.307–0.340 (1.378–1.586/
with 3 small terminal rami; flagellum of 4 0.333–0.390), chela (without pedicel) 1.318–
blades (Fig. 12); serrula exterior with 19 ((cid:63), 1.376(1.306–1.517),handlength0.634–0.672
(cid:47)) blades; lamina exterior present. Carapace (0.592–0.718), movable finger length 0.670–
(Fig. 15) 0.97–1.09 ((cid:63)), 1.13–1.32 ((cid:47)) times 0.736 (0.656–0.776). Chelicera 0.250/0.128
longerthanbroad;lateralmarginsevenlycon- (0.290/0.128), movable finger length 0.147
vex, but posteriorly widened; with 2 non-cor- (0.197). Carapace 0.846–0.909/0.816–0.938
neate eyes; with ca.44setae,including4near (0.845–0.944/0.640–0.832); eye diameter