Phantom spiders: notes on dubious spider species from Europe

2015·13.7 MB·English

by  Rainer Breitling

5
Arachnologische Mitteilungen 50:65-80 Karlsruhe, November201
Phantom spiders: notes on dubious spider species from Europe
RainerBreitling, Martin Lemke,Tobias Bauer, Michael Hohner,ArnoGrabolle&Theo Blick
doi: 10.5431/aramit5010
Abstract.AsurprisinglylargenumberofEuropeanspiderspecieshaveneverbeen reliablyrediscoveredsincetheir
firstdescription many decades ago. Mostofthese are probably synonymouswith other species or unidentifiable,
due to insufficient descriptions or missing type material. Here we discuss about 50 ofthese cases, declare some
names as nomina dubia and establish the following new or re-confirmed synonymies:Agelenomengeella Strand,
1942=Allogelenagracilens(C.L.Koch, 1841)syn.conf.;Anyphoenaaccentuotoobscura(Sundevall, 1831)=Anyphae-
naaccentuoto(Walckenaer, 1802) syn.conf.;AnyphoenoaccentuotoobscuraLebert, 1877 =Anyphoenaaccentuoto
(Walckenaer, 1802)syn.nov.;AraneusdiadematusstellatusC. L.Koch, 1836=AraneusdiodemotusClerck, 1757syn.
nov.;Araneusdiodemotusislondicus(Strand, 1906)=AraneusdiodemotusClerck, 1757syn.nov.;Araneusquadrotus
minimusSimon, 1929=AraneusquadrotusClerck, 1757syn.nov.;Araneusquadrotussubviridis(Franganillo, 1913)=
AraneusquadrotusClerck, 1757syn.nov.;Centromerusunctus(L. Koch, 1870)=Leptorhoptrumrobustum (Westring,
1851) syn. nov.; Clubiona caliginosa Simon, 1932 = ClubionagermanicaThorell, 1871 syn. nov.; Coelotes atropos
anomalusHull, 1955=Coelotesatropos(Walckenaer, 1830)syn.nov.;CoelotesatropossilvestrisHull, 1955=Coelotes
atropos(Walckenaer, 1830) syn.nov.;CoelotesobesusSimon, 1875 =Pireneitegapyrenaea(Simon, 1870) syn.conf.;
CoelotessimoniStrand, 1907 = Coelotessolitarius (L. Koch, 1868) syn. nov.; Diplocephalussemiglobosus (Westring,
1861)nomenoblitum=Entelecaracongenera(O.P.-Cambridge, 1879)syn.nov.;Drassodesvoigti(Bösenberg, 1899)
= Scotophaeus blackwalli(Thorell, 1871) syn. conf.; ErigonedecensThorell, 1871 = Hylyphantesgraminicola (Sun-
devall, 1830) syn. nov.;Liocranoecastriatagracilior(Kulczynski, 1898) = Liocranoecastriata (Kulczynski, 1882) syn.
conf.;Phlegrarogenhoferi(Simon,1868)=Phlegracinereofasciata(Simon,1868)syn.nov.;Styloctetorstativus(Simon,
1881)=Styloctetorcompar(Westring, 1861)syn.nov.andcomb,nov.;Tapinocybabilacunata(L.Koch,1881)=Silom-
etopusincurvatus (O. P.-Cambridge, 1873) syn. nov.; Theridion variansmelanotum Strand, 1907 = Theridion varians
Hahn, 1833 syn. nov.; Thomisus trigonusGiebel, 1869 = Pistius truncatus (Pallas, 1772) syn. nov.; Titanoecapsam-
mophilaWunderlich, 1993 =Titanoecaspominima(Taczanowski, 1866) syn.nov.and comb,nov.;Xysticuspaniscus
L.Koch, 1875=Xysticuslineatus(Westring, 1851)syn.conf.
Keywords:Araneae,doubtful species,newsynonyms,nomendubium,speciesinquirendae
Zusammenfassung. Phantomspinnen: Bemerkungen zu zweifelhaften Spinnenarten aus Europa. Eine über-
raschendeAnzahl von europäischen Spinnenartenwurdeseit ihrerErstbeschreibung nie mehrzuverlässig wieder-
gefunden. In den meisten Fällen handelt es sich vermutlich um Synonyme anderer Arten oder die Arten bleiben
aufgrund von unzulänglichen Beschreibungen und verlorenem Typusmaterial unidentifizierbar. Hier besprechen
wiretwa 50dieserFälle,erklärenzahlreiche Namenzu nominadubia und identifiziereneine Reihevon neuen oder
bisherübersehenenSynonymien.
A
surprising number ofspider species listed as va- are probably nomina dubia, unidentifiable on the
lid on the European checklists and databases (e.g., basis ofthe original descriptions,but to conclusively
van Helsdingen 2014,World Spider Catalog 2015, determine the status ofthese species, a careful ex-
Nentwig et al. 2015) have never been reliablyredis- amination ofeach individual case is necessary (van
covered after their initial description. Most ofthese Helsdingen 2004).The status ofsome ofthese spe-
cies has been clarified as part oflarger revisionary
work or in isolated papers (e.g., Kronestedt 2000,
RainerBREITLING,FacultyofLifeSciences,UniversityofManchester, van Helsdingen 2008). An extended discussion of
ManchesterMl 7DN,UK;E-Mail:[email protected] dubious species described by Bösenberg was also
MartinLEMKE,Wakenitzmauer23,23552Lübeck,Germany;
E-Mail:[email protected] provided by Braun (1982), but many cases still re-
TobiasBAUER,Hattenbachweg12,70599Stuttgart,Germany; main to be examined.
E-Mail:[email protected]
MichaelHOHNER,Juvenellstr.26,90419Nürnberg,Germany; The Working Group “Forum and Wiki” of the
AE-rMnaoilG:RaArBaOgLeLsE@m,hAomhnHeorr.nde13b,99425Weimar,Germany; Arachnologische Gesellschaft (Lemke et al. 2014)
E-Mail:[email protected] has recently started an online project documenting
TheoBLICK,Heidloh8,95503Hummeltal,Germany;E-Mail:info@theo- the information available on suspected “phantom
blick.de;SenckenbergResearch,StrictForestReserves,60325Frankfurt
amMain,Germany;E-Mail:[email protected] spiders”, with an initial focus on species from Cen-
submitted15.4.2015,accepted28.7.2015,online12.8.2015 tral Europe.
;
66 R.Breitling,M.Lemke, TBauer;M.Hohner;A.Grabolle&T.Blick
The curators of the following collections were types, stated (in Lebert 1877) that the species was
BMNH
contacted to trace possible type material: = new and occurred not only at the type locality in
British Museum ofNatural History (including large Switzerland,butalso in SouthTyrol (Italy).The type
parts ofthe L. Koch collection,Jan Beccaloni),MfN locality at an altitude of 1280 m indicates that this
= Museum für Naturkunde, Berlin (Dahl collection, maybe a montane or alpine species, such as Pellenes
Jason Dunlop), NMB = Naturhistorisches Museum lapponicus(Sundevall, 1833),which shows suggestive
Basel (Schenkel collection, Ambros Hänggi), NRS similarities in the genitalia, but this remains specu-
= Naturhistoriska riksmuseet Stockholm (parts of lative. Neither the illustration of the palpus, which
the Thorell collection, including Westring material, is apparently shown in expanded state, nor the very
NSMW
Torbjörn Kronestedt), = Naturhistorische schematicfigureoftheepigyne,northeextensivede-
Sammlungen Museum Wiesbaden (Zimmermann scription seem sufficient to allow a confident identi-
OUM
collection, Fritz Geller-Grimm), = Oxford fication ofthis species.The type material is probably
University Museum (Pickard-Cambridge collection, lost (see Introduction).
ZoeSimmons),SMF= SenckenbergMuseumFrank-
furt (Braun andWunderlich collections,PeterJäger), Agelena mengeella Strand, 1942 =Allagelenagracilens
ZMH = Zoologisches Museum Hamburg (parts of (C. L. Koch, 1841) syn. conf. - syn. nov. in Bonnet
the Bösenbergcollection,includingBertkaumaterial, (1955) (Agelenidae)
ZMW
Kai Schütte), = Zoological Museum Warsaw This specieswasfirstdescribedbyMenge (1871:285,
(Kulczynski and Taczanowski collections, Dominika pi. 52, f. 165) as Agalena brunea, matching a species
Mierzwa-Szymkowiak),ZSH = Zoologische Samm- similar to Allagelena gracilens to egg sacs similar to
lung der Martin-Luther-Universität, Halle (Giebel those ofAgroeca brunnea (Blackwall, 1833). Strand
collection, Karla SchneiderScjoachim Händel). (1942) noticedthe errorandproposedthe newname
Some of the most important collections in A. mengeella however, he did not examine the type
this context are known to be lost, including tho- material andwas uncertain about the actual identity
se of Menge (formerly in the Provincial Museum of the species, although he realized that it is most
Gdansk, but probably lost at the end ofWorld War likelythatMenges specimensbelonged to eitherAl-
II; Kraus 2009), Lebert (formerlyperhaps in Zurich lagelena gracilens or Agelena labyrinthica. These two
or Wroclaw, where it could not be found; Hänggi species were commonly confused at the time, but
pers. comm.) and Bösenberg (formerly in the Kö- the illustrated pedipalp and epigyne both support an
nigliches Naturalienkabinett in Stuttgart, but des- identification with the former, and Menge himself
troyed during a bombing raid on September 12th, had alreadypointed outthe similarity-his misiden-
1944; Renner 1988).Inthispaper,we summarize the tification was apparently only based on the wrongly
results for a selection ofspecies for which the type assigned egg sacs.Against Strand (1942), and in ag-
specimenshavebeenlocated andexamined,orwhere reementwith Bonnet (1955),we therefore conclude
they are in all probability lost. We also synonymise that even in the absence ofthe type material the sy-
a number ofsubspecies with their nominate form, if nonymy ofthe two species can be established with
theywere originally described as sympatric (or even confidence.This is alsoin agreementwithPröszynski
&
syntopic) variations and thus cannot be considered Star^ga (1971), who also synonymizedA. brunea
as subspecies in the modern sense. More detailed with A. gracilens, following the use ofthe name by
supporting information, including all original de- several earlier Polish authors.
scriptions and figures, is available on the associated
Wiki page (http://wiki.spinnen-forum.de/index. AgelenamengeiLebert,1877= nomendubium(Age-
php?title=Phantomarten). lenidae)
Leberts description ofafemale isveryextensive,but
Speciesaccountsinalphabeticalorder does not allow an unambiguous identification (Le-
Aelurillus simoni (Lebert, 1877) = nomen dubium bert 1877: 211, pi. 6, f. 42). The most likely candi-
(Salticidae) date would seem to be Agelena labyrinthica, which
The original description was based on three adult matches the description and illustration very well;
females and a male (Lebert 1877: 310, pi. 6, f. 45- however, Lebert reports A. labyrinthica from many
47; as Aelurops simoni). L. Koch, who had seen the locations, and insists that this specimen belongs to a
Shorttitle:PhontomspidersfromEurope 67
different species, although the only diagnostic diffe- ofwhite spots,the smaller size and the psammophi-
rence explicitly mentioned seems to be a slightvari- lous (not titanophilous) habitat.The large numberof
ation in bodyproportions. Possiblythe species could specimens examined byTaczanowski makes it unli-
even be a memberofTegenarias.lat. kelythatthespecimenwasanunusuallysmallordark
formofanotherspecies.Eventhoughthedescription
Agyneta resima (L. Koch, 1881) = nomen dubium onlymentionsthesizeofafemalespecimen(perhaps
(Linyphiidae) because it was particularlylarge), there is no indica-
ThisspecieswasdescribedinthegenusErigone(Koch tion that only females were found, and the striking
1881: 50, pi. 2, £ 4), which at that time included a whitespotsofmales (andmostsubadultmales) ofre-
largepartofthe smallLinyphiidae,butthe similarity lated species would not have escapedTaczanowskis
withAgynetarurestris(C.L.Koch,1836) discussedin attention. Braun (1969) had mentioned Tpsammo-
the original descriptionjustifies the transfer toAgy- phila as a“melanistic and nanistic form”ofTquadri-
neta. A more precise identification seems, however, guttatafrom the Mainzer Sand (SMF20769/15119,
impossible,based on the veryvague illustrations and examined by TB). Other records are known from
textualdescription.Thetypematerialseemstobelost southern Sweden (Öland,Östergötland, Stockholm)
(not in BMNH).T. resima is one ofseveral dubious and Finland (Aboland), from the dunes north-east
speciesincludedbyRoewer(1928) inhis keyofGer- of Berlin (Pimpinellenberg), from sandy meadows
man spiders.He even addedadditionaldetails on the in the South ofthe Czech Republic (Hodonin area),
habitat and phenology ofthe species, although the and neighbouring regions of Slovakia (Laksarska
source ofthis information is unclear.The popularity Nova Ves), from South Hungary (Kiskunsag Natio-
and easyaccessibilityofRoewer’sworkis mostlikely nal Park), from the Perm Region in the easternmost
responsible for reports ofthe species from Cieszyn part ofthe European part ofRussia, and in Poland
in South Poland (Ksi^zköwna 1936), as well as for from Biebrza National Park, 200 km north-east of
the notorious “Balkan rediscoveries”ofmany ofthe Warsaw (Kupryjanowicz 1997a, Jakobitz 8c Broen
species discussed here (see Braun 1982 for details). 2001, Gajdos 8cMajzlan 2005, Esyunin 2006, Galle
8c Feher 2006, Kronestedt 2010, Hula et al. 2014).
Amaurobius spominimus Taczanowski, 1866 = 77- Thus, although no recent records of Tpsammophila
tanoecapsammophila Wunderlich, 1993 = Titanoeca are known from the Warsaw area, the locus typicus
spominima (Taczanowski, 1866) syn.nov.and comb, ofA. spominimus is located in the epicentre of the
nov.(Titanoecidae) known distribution and consists ofvery typical ha-
This species was described byTaczanowski without bitat. As T. psammophila was only described quite
figures andwith averyshort description ofless than recently and has beenveryrarelyreported,the name
3 lines (Taczanowski 1866: 4), and the type materi- is not protected by prevailing usage, and the older
al appears to be lost (not in ZMW). It would thus synonym takes priorityas Titanoecaspominima.
seem an obvious candidate forbeing a nomen dubi-
um. However, the short description, which is based Anyphaena accentuata obscura (Sundevall, 1831) =
on specimens collected in the dunes of Prag^ and Anyphaena accentuata (Walckenaer, 1802) syn. conf.
D^browaclose toWarsaw,mentions a numberofdi- - syn. nov. in Sundevall (1833) (Anyphaenidae)
stinguishing characters that allow a confident iden- AnyphaenaaccentuataobscuraLebert, 1877 =Anypha-
tification: “Prosoma reddish-brown; opisthosoma enaaccentuata (Walckenaer, 1802) syn.nov.
short,rounded,hairy,black;legs reddish-blackhairy; Anyphaenasabina Bertkau, 1880 (misidentification)
length: female 4 mm.About 10 specimens collected =AnyphaenafurvaMiller, 1967
in sand under a lawn ofreindeer lichen Cladonia)”. Anyphaena accentuata obscura Bertkau (in Förster 8c
(
Of all cribellate species in the area, only Titanoeca Bertkau 1883) (misidentification,notA.a. o. Lebert,
psammophila shares these characters Titanoeca spe- 1877) =AnyphaenafurvaMiller, 1967
(
cies were often placed inAmaurobius at the time of Anyphaena obscura Bösenberg, 1902 (misidentifica-
Taczanowskis work). Tpsammophila was long con- tion, not A. a. o. Lebert, 1877) = Anyphaenafurva
fusedwith Titanoecaquadriguttata(Hahn, 1833),but Miller, 1967
is distinguished from this and other Central Euro- The name obscura was first used by Sundevall for a
pean Titanoeca species by the combination ofa lack specimen similar to Tegenaria domestica (Sundevall
68 R.Breitling,M.Lemke, T.Bauer,M.Hohner,A.Grabolle&TBlick
1831: 21,subAgelenaobscura),but alreadytwoyears Araneusdiadematusstellatus C. L. Koch, 1836 =Ara-
later he realized that this specimen was an old fe- neusdiadematusClerck, 1757 syn.nov.(Araneidae)
male ofAnyphaena accentuata in which the charac- Araneusdiadematusislandicus(Strand,1906) =Arane-
,
teristic markings ofthe opisthosoma had been ob- usdiadematusClerck, 1757 syn.nov.
literated (Sundevall 1833: 265,269).The name was Araneus quadratus minimus (Getaz, 1889) = nomen
later used independently by Lebert (1877: 242) for nudum
A
a dark variety (“Spielart”) of accentuata, without Araneus quadratus minimus Simon, 1929 = Araneus
.
reference to Sundevall andwithout the intention to quadratusClerck, 1757 syn.nov.
establish a subspecies in the modern sense. Bert- Araneus quadratus subviridis (Franganillo, 1913) =
&
kau (in Förster Bertkau 1883: 210) uses Leberts AraneusquadratusClerck, 1757 syn.nov.
name for the males ofa dark species ofAnyphaena The name Araneus quadratus minimus was first used
found in Bonn, Germany. This species is, however, by Getaz (1889: 60; sub Epeira quadrata, var. minima)
clearly distinct from A. accentuata as can be seen in a list ofspiders from Pays-d’Enhaut (canton Vaud,
,
A
from the illustration ofthe male pedipalp provided Switzerland),butwithoutanydescription. descripti-
by Bösenberg (1902: 258, pi. 24, f. 373) based on onwas onlyprovidedfortyyears laterbySimon (1929:
Bertkaus material (nowlost; Braun 1982). It seems 683),who must therefore be regarded as the valid au-
verylikelythatthesespecimens actuallybelongedto thor ofthis taxon.A. q. minimus was the only named
A.furva, a rare species ofAnyphaena. Although the varietyofA. quadratusSimonmaintainedinhisArach-
tibial apophysis as illustrated by Bösenberg is cer- nides de France, reporting it as a local montane form,
tainlyexaggerated,itis sufficientlysimilarto thatof found on dwarf shrubs of alpine meadows; it is thus
A.furva which is broader and more massive (“brei- nota subspecies in the modern sense. Similar melanis-
,
terund plumper”) than thatofA. accentuata (Miller ticspecimens aretypicalforboreoalpinepopulationsof
1967). Also, the lack ofventral spines at the base Araneus diadematus as well (e.g., var. islandicus Strand,
of the pedipalpal femur, the more uniform dorsal 1906,andvar.stellatusC.L. Koch, 1836,bothofwhich
hairs on the pedipalpal tibia,and the more cylindri- wouldnotbeconsideredsubspeciesinthemodernsen-
cal (rather than anteriorly broadened) shape ofthe se,andhave tobe treated as synonyms ofthe nominate
tibia are clearlyvisible in comparison to the figures form).The synonymyforvar. stellatuswas alreadypro-
ofA. accentuata on the same plate and match the posed by Thorell (1870) and Lessert (1910), but not
diagnostic features ofA.furva (Miller 1967). Mo- acceptedbyalllater authors (e.g., Simon 1929).
reover,A.furva is regularly found as almost black Another montane form ofAraneus quadratuswas re-
specimens (Bauchhenss 2009). No other European portedbyFranganillo from Spain (Franganillo 1913:
species of Anyphaena matches the description of 127),where he found female specimens in their sil-
Bertkaus specimens. Bertkau (1880: 253) had ori- ken retreats “in gorse and low shrubs on the slopes
ginallyreported his specimens asA. sabina but had of the mountains” in the surroundings of Gijon or
,
changed his opinion after a male had been exami- La Guardia.From the description itis clear that this
ned by Simon, and the shape ofthe pedipalpal ti- greenishformofthe species (“withfourstrikinglyvi-
bia certainly excludes this identification.A.furva is sible spots”)wasneverintendedas asubspeciesinthe
foundinxerothermichabitats andwouldbe another modern sense,anditwas neverusedas suchbyFran-
example ofa distinctlythermophilic elementrepor- ganillo,whodescribesitas avarietyonly.^. quadratus
ted by Bertkau for the Bonn area. Other thermo- has been reported as being able to activelychange its
philous species, often with Ponto-Mediterranean colour (Bunn 1957), and the features ofthe epigyne
affinities, found by Bertkau around Bonn include, (“scapus large and bent upwards, as in Epeira trifoli-
e.g., Cetonana laticeps Sagana rutilans, Euryopis um Hentz”,referring to a lateralview ofthe epigyne
,
quinqueguttata Heriaeus graminicola (sub Heriaeus illustratedin Emerton 1884) also seem to fallwithin
,
hirtus in Braun 1960),Pellenes nigrociliatus Philaeus the normalvariation ofA. quadratus.
,
chrysops and Saitis barbipes (Bertkau 1880, Bösen-
,
berg 1903, Braun 1960).Therefore, his discovery of Araniella silesiaca (Fickert, 1876) = nomen dubium
A.furva which extends the known areaofthis rare- (Araneidae)
,
lyreported species byseveralhundred kilometres to The status ofthis species, which had been first de-
thewest,is not all that surprising. scribed as Epeiras. based on a female specimen from
Shorttitle:PhantomspidersfromEurope 69
the environs ofWroclaw (Fickert 1876: 70),was ex- 1843). Koch’s mistake was first noticed by Thorell
amined in detail by Blanke (1982), who concluded (1871), who redescribed the species under the new
thatthespeciescannotbeunambiguouslyrecognized name Clubionagermanica. Simon, however, felt that
based on the original description. While many au- themales andfemalesillustratedbyKochdidnotbe-
thors have identified the species as Araniella alpi- longtothe same species andintroducedanothernew
ca (L. Koch, 1869), the existence of highly similar name for the latter. The justification for this move
forms,including the sister speciesA. inconspicua (Si- is unsatisfactory: the (admittedly crude) illustrati-
mon, 1874),precludes aconfidentassignmentto this on ofthe epigyne provided by Koch does certainly
species.The originaltypes areverylikelyto be lost. show sufficient similarity with that of C. germanica
,
andnothingin Koch’s description argues againstthis
Bathyphantes enslini Strand, 1910 = nomen dubium identification.The collection O. Pickard-Cambridge
(Linyphiidae) inthe OxfordUniversityMuseum ofNatural Histo-
This specieswas describedbyStrand (1910: 48) from ry contains specimens ofC. holosericea from Nurem-
a juvenile and poorly preserved specimen found in berg labelled as types (Bottle 281.9); these are most
a cave in Franconia, Germany. The types are most likelythe materialunderlyingThorell’s descriptionof
likelylost (not inMfN),and even iftheywere redis- C. germanica. The type of C. caliginosa however, is
,
covered, a confident identification would be close to the illustration ofthe epigyne publishedbyKoch.
impossible.
CoelotesatroposanomalusHull,1955 = Coelotesatropos
Centromerus ludovici Bösenberg, 1899 = nomen du- (Walckenaer, 1830) syn.nov.(Agelenidae)
bium(Linyphiidae) CoelotesatropossilvestrisHull, 1955 = Coelotesatropos
The type material of this species described by Bö- (Walckenaer, 1830) syn.nov.
senberg (1899: 115, pi. 1, f. 2) was destroyed du- The two “varieties” described by Hull were always
ring World War II (Renner 1988), like many of found together with the typical forms; they are not
Bösenberg’s types. Wunderlich (1973) and Braun subspecies as currentlyunderstood,butratherindivi-
(1982) consider the species as a member ofwhat is dualvariantsofasingle,highlyvariablespecies (types
nowthegenusAgyneta,but agree thatamoreprecise notin BMNH).
identification is impossible.
Coelotes obesus Simon, 1875 = Pireneitega pyrenaea
Centromerus unctus (L. Koch, 1870) = Eeptorhoptrum (Simon, 1870) syn.conf.-syn.nov.in Simon (1937)
robustum (Westring, 1851) syn.nov.(Linyphiidae) (Agelenidae)
The original descriptionby Koch (1870: 24,sub Eri- This synonymy was already recognized by Simon
gone uncta) already doubted the validity ofthis spe- (1937: 1034), but overlooked in subsequent cata-
cies and pointed out the close similarity to Erigone logues.
huthwaitii (O. R-Cambridge, 1861) (= L. robustum).
The mentioned diagnostic characters do not allow a Coelotes simoni Strand, 1907 = Coelotes solitarius (L.
discriminationfromthis species,and Koch mentions Koch, 1868) syn.nov.(Agelenidae)
that any observed differences could easilybe explai- C. simoni was suggested as a new name for a speci-
nedbythe commonlyobservedexpansionofthepal- men ofC. solitarius illustrated by Simon (1898: 173,
pal organs. Even though the type material seems to f. B),which Strand (1907: 392) considered misiden-
belost,thesynonymyseemsjustified,consideringthe tified, without examination ofthe original material
highly distinct male genitalia ofL. robustum and the and without any further explanation. There is no
factthatitisthesolememberofitsmonotypicgenus. indication that Strand’s decision wasjustified, given
that Simon was well acquainted with C. solitarius
,
ClubionacaliginosaSimon, 1932 = Clubionagermani- as shown by numerous records ofthe species in the
caThorell, 1871 syn.nov.(Clubionidae) Arachnides de France.
The name C. caliginosa was introduced by Simon
(1932: 965) for the female of a species originally Diplocephalus semiglobosus (Westring, 1861) nomen
considered by Koch (1867: 311) as Clubiona holo- oblitum = Entelecara congenera (O. P.-Cambridge,
sericea De Geer (= Clubionaphragmitis C. L. Koch, 1879) syn.nov.(Linyphiidae)
70 R.Breitling,M.Lemke, T.Bauer,M.Hohner,A.Grobolle&TBlick
The type ofthis species,whichwas described as very Drassodes myogaster (Bertkau, 1880) = nomen du-
similar to Entelecara acuminata (Wider, 1834) and bium(Gnaphosidae)
Diplocephalus picinus (Blackwall, 1841) (Westring This species, described as Drassus m. based on a sin-
1861: 235; sub Erigone semiglobosa), is preserved in gle female from Bonn (collected at exactly the same
Thorell’s collectionintheNRS.Itwas originallykept location asAnyphaenaaccentuata obscura),was repea-
in dried state on a pin, and transferred into alcohol tedlysynonymizedwithDrassodeslapidosus(Walcke-
by Ake Holm, who also studied the specimen and naer, 1802) (e.g., by Reimoser 1937, Grimm 1985).
discovered the synonymywithE. congenera as recor- This is, however, dubious, considering not only the
,
ded on a label he added to the vial (Kronestedt pers. presence ofthe morphologically all but indistingu-
comm.). D. semiglobosus is the senior synonym, but ishable sister species D. cupreusin the same area,but
it has not been used as a valid name after 1899; to also the fact that Bertkau did describe D. lapidosus
our knowledge, it only occurs in general catalogues, in the same paper, and saw closer similarities ofD.
which should be considered mere nomenclators ac- myogaster with D. pubescens D. luteomicans (sub D.
,
cording to article 23.9.6 of the International Code portator), D. rubidusand D. villosus.As the type ma-
ofZoological Nomenclature. In contrast, the junior terial is apparently lost, no reliable identification of
synonymE. congenerahasbeeningeneraluse forthe the species is possible.
last 100years,beingusedbyfar more than 25 works
by more than 10 authors in the last 50 years (for ex- Drassodes voigti (Bösenberg, 1899) = Scotophaeus
ample,Miller 1971, Klomp &,Teerink 1973, Locket blackwalli (Thorell, 1871) syn. conf. - syn. nov. in
et al. 1974,Punda 1975,Albert 1979,Bauchhenss et Grimm (1985) (Gnaphosidae)
al. 1987,Roberts 1987,Baehr 1988,Blick&,Scheid- The species, described as Drassus voigtii was initi-
& ,
ler 1991, Hauge Hansen 1991, Heimer &cNent- ally synonymized with Drassodes villosus (Thorell,
wig 1991,Braun 1992, Schultz 1992,Millidge 1993, 1856) (Reimoser 1937). Only the discovery ofa pu-
Albrecht 1995, Finch 1997, Hermann 1998, Svaton tative female syntype (paratypoid) in Bösenberg’s
&
Pridavka2000,Tutelaers2000,Harveyet al.2002, collection in the Zoological Museum Hamburg by
Ratschker et al. 2005, Van Keer &, Van Keer 2005, Grimm (1985) revealed that the species is synony-
&
De Köninck 2006, Otto Floren 2007, Russell- mous with Scotophaeus blackwalli. In retrospect, this
Smith 2011, Wunderlich 2011, Staudt et al. 2012, matches Bösenberg’s illustrationofthe epigyne quite
Kostanjsek &,Kuntner 2015).Therefore,we propose well (Fig. 1), and the type locality in the inner city
that Diplocephalus semiglobosus (Westring, 1861) is ofBonn also agrees with the synanthropic habits of
considered as nomen oblitum, and that Entelecara S. blackwalli in Central Europe (Grimm 1985; Ro-
congenera(O.P.-Cambridge, 1879) isvalidas nomen berts 1998).Incidentally,this case illustrates that the
protectum accordingto article23.9 ofthe Internati- declaration of taxa as nomina dubia will always be
onal Code ofZoologicalNomenclature. tentative; an initial careful revision of Bösenberg’s
Fig. 1: Comparison
oftheoriginal illus-
trationofDrassodes
voigti (Bösenberg,
1899: pi. 1, f. 5)
and the epigyne of
Scotophaeus blaclc-
wa///(Thorell,1871),
showing an excel-
lent agreement in
overall shape and
proportions (photo
byArnoGrabolle)
Shorttitle:PhantomspidersfromEurope 71
collectionbyBraun(1982) failedtodiscoverthetype The holotype ofErigone compar, a single dried male
and concluded that D. voigti was a nomen dubium and pedipalp in the collection ofthe Naturhistoris-
(“dubioseArt”). ka riksmuseet, Stockholm,was examined in 1942 by
Ake Holm, who added a corresponding identifica-
ErigonedecensThorell, 1871 = Hylyphantesgraminico- tion labelto the specimen (Kronestedtpers.comm.).
la (Sundevall, 1830) syn.nov.(Linyphiidae) He identified the specimen as belonging to Stylocte-
This species was described in a footnote to the dis- torstativus. Westring’s name is the senior synonym,
cussion ofErigonedentifera (= H.graminicola) based and as ithas been used repeatedlysince 1899,due to
,
on a male specimen thatThorell could “scarcely dis- a mistaken synonymization ofG. comparand G. late-
tinguish from E. dentifera (Thorell 1871: 128).The bricola(sensu Simon) byHull (1932),the oldername
type specimen is preserved in Thorell’s collection in cannot be considered as a nomen oblitum according
the Naturhistoriska riksmuseet, Stockholm; its exa- to article23.9 ofthe International Code ofZoologi-
mination by Ake Holm revealed that it is a gynan- cal Nomenclature. Even though S. stativus has been
dromorphous specimen ofH. graminicolawith fully verywidelyusedinthelast 100years,anditsreplace-
formed epigyne and typical male pedipalps (Krones- ment by the senior synonym will be inconvenient, it
tedtpers. comm.). does not seemjustifiedto appealto the International
Commission on Zoological Nomenclature reques-
Euophrysstriolata(C.L.Koch, 1846) = nomendubi- tingarulingundertheplenarypowerto suppress the
um(Salticidae) older synonym.Thus,we suggestthat Styloctetorcom-
This species,firstdescribedbasedon apoorlypreser- par(Westring, 1861) should be considered the valid
ved female from near "Carlsbad in Böhmen”, which name forthe species.
is now Karlovy Vary in the Czech Republic (Koch
1846: 47-48, f. 1306), is similar to Euophrysfron- Eepthyphantes beckeri Wunderlich, 1973 = nomen
talis and E. terrestris. This is one ofthe few species dubium(Linyphiidae)
described by C. L. Koch that Simon (1864) lists as This species, in contrast to most ofthe species dis-
"speciesinvisa”inhisrevisionofEuropean Salticidae, cussed here, has been found again after the original
so presumablythe type was alreadylostbythen.The description. It turned out that all specimens found
description and figure do not allow an unambiguous were parasitized females, and it is likely that they
identification,beyondthefactthatthisis almostcer- are malformed representatives ofa common species
tainly the synonym of a common species (Bonnet (Harvey et al. 2002) belonging in the genus Tenu-
1955). iphantes. The original description emphasizes the
similarity in habitus to 77 mengei, but Wunderlich
Gonatiumfuscum Bösenberg, 1902 = nomendubium (2008) statedthatboth T.mengeiand T.flavipeswere
(Linyphiidae) commonatthetypelocalityandconsideredL.beckeri
GonatiumgilbumBösenberg, 1902 = nomendubium a nomen dubium.
Gonatium pallidum Bösenberg, 1902 = nomen
dubium Eepthyphantes thienemanni Schenkel, 1925 = nomen
The type material ofthese species was destroyed in dubium(Linyphiidae)
World War II (Renner 1988). The descriptions do The female type specimen is preserved in Schenkels
not allow an unambiguous identification. Despite a collection in the Naturhistorisches Museum Basel;
number oftentative identifications in the literature, however, its epigyne is missing.The description em-
an unambiguous identification is impossible in all phasises the noticeablysmall and pale appearance of
cases (Braun 1982). All reported specimens from the epigyne as the main diagnostic character. This
Eastern and Southern Europe that could be exa- indicates that the specimenwas probably a subadult
mined turned out to belong to well-known species femaleofawidespread andcommon species ofLept-
(Braun 1982). hyphantess.lat.,especiallyas anotherfourfemalespe-
cimenswerelaterfoundinFebruary,MarchandJune
Gongylidiellum compar (Westring, 1861) = Styloctetor inbogsinNorthwestGermanybyPeus (1928;mate-
stativus(Simon,1881) = Styloctetorcompar(Westring, rial determinedby Schenkel,but apparentlylost,not
1861) syn.nov. and comb.nov.(Linyphiidae) in NMB).
,
72 R.Breitling,M.Lemke, J.Bauer,M.Hohner,A.Grabolle&TBlick
Liocranoecastriatagracilior(Kulczynski, 1898) = Lio- species certainlybelongs to Oedothorax but also sug-
,
cranoeca striata (Kulczynski, 1882) syn. conf. - syn. gests a possible synonymy with Dismodicus elevatus
,
nov. in Simon (1932) (Liocranidae) based on misidentified specimens from Romania
This taxon, described asAgroecagracilior which pro- (his reasoning in this case is not quite clear: there is
,
bably represents individual intraspecific variability, no reason to assume that the Romanian specimens
according to the original description is identical in had been compared to authentic material).The type
genitalic structure to the nominate form. It was al- material was probably lost together with the other
ready synonymized by Simon (1932), and this deci- Gonatium types ofBösenberg (Renner 1988).
sion was followed by most subsequent authors (e.g.,
Reimoser 1937, Bonnet 1955, Sterghiu 1985). The Oedothoraxpallidus (Bösenberg, 1902) = nomen du-
type material seems to be lost (notin ZMW). bium(Linyphiidae)
This specieswas originallydescribed inKulczynskiel-
Micrargus incomtus (O. R-Cambridge, 1872) = no- lum F. O. Pickard-Cambridge, 1895.Wunderlich (in
mendubium(Linyphiidae) Braun 1982) considered the species as belonging to
The type material ofthis species seems to be missing Oedothorax probably O. retusus but Braun disagreed
,
in the collection O. Pickard-Cambridge in the Ox- and suggested a possible identity with Gongylidium
fordUniversityMuseum ofNatural History.The de- rufipes (Linnaeus, 1758) instead. The type material
scription,as Erigoneincomta,compares the species to probablywas losttogetherwith most otherlinyphiid
Agyneta saxatilis (Blackwall, 1844) and Maso sunde- types ofBösenberg (Renner 1988),makingareliable
valli (Westring, 1851),but the form ofthe pedipalp, identification impossible.
especiallythelongspiralembolus exclude acloseraf-
finitywith these species and instead support aplace- Oedothoraxsubniger(Bösenberg, 1902) = nomendu-
ment in MicrargusDahl, 1886. However, the typical bium(Linyphiidae)
prosomalgrooves ofthe males ofthis genus are mis- Suggested identifications for this species, described
sing in the figures and description. In the absence of as Kulczynskiellum subnigrum have been Gonatium
,
type material,the species remains unidentifiable. rubens (Blackwall, 1833) and Hylyphantesgraminico-
la (Sundevall, 1830) (Braun 1982). However, as the
Microneta iracunda (O. P.-Cambridge, 1879) = no- type material probably was lost together with most
mendubium(Linyphiidae) otherlinyphiid types ofBösenberg (Renner 1988), a
This species, described as Neriene iracunda, after a reliable identification is impossible.
single male collected by Eugen Count Keyserling
in “Lüvland” (present-day Latvia and Estonia), is Oedothorax tener (Bösenberg, 1902) = nomen dubi-
described as being allied to Agyneta subtilis (O. P.- um(Linyphiidae)
Cambridge, 1863), A. conigera (O. P.-Cambridge, Another ofBösenberg’s species, originallyKulczyns-
1863) and A. innotabilis (O. P.-Cambridge, 1863). kiellum tenerum for which the type material is lost
,
Pickard-Cambridge was obviously quite familiar (Renner 1988) and an identification based on the
with this group, and it is very likely that the species incomplete descriptionis impossible.Even the gene-
should be placed in Agyneta. However, as the type ric assignment is uncertain and Wunderlich (1973)
material could not be found in the Pickard-Cam- suggested apossible placementin Tapinocyba Simon,
bridge collectionin Oxford,noreliable identification 1884.Nevertheless,the species,whichwas illustrated
at the species levelis possible. by Roewer (1928), has been reported from the Bal-
kans (Drensky 1929, 1936).
Oedothorax insignis (Bösenberg, 1902) = nomen du-
bium(Linyphiidae) Pardosa bernensis (Lebert, 1877) = nomen dubium
Originally described in Gonatium, the species was (Lycosidae)
&
transferred to Oedothorax (Bertkau, in Förster As the type ofthe speciesLycosabernensisis probabi-
Bertkau, 1883) byWunderlich (1974), based on the lity lost, no unambiguous identification ofthis spe-
similarity of the epigyne to that of species such as cies is possible. The two most likely candidates are
Oedothorax retusus and Oedothorax apicatus. Braun Acantholycosapedestris (Simon, 1876) and Pyrenecosa
mm
(1982) confirms that according to the epigyne the rupicola (Dufour, 1821), based on the size (12
Shorttitle:PhantomspidersfromEurope 73
total length), the type locality in the Bernese limes- was malformedduetoparasitism,orthetypewas in-
tone Alps, and the “strange”(“sonderbar”) habitus of deed a rare hybrid with intermediate characters (see
the specimen: dark black, long-legged and almost Martin 2013 for a discussion of possible causes of
without pattern. Both A. pedestris and P rupicola genital malformations in Pardosa). Pardosa species
are large, almost uniformly black species from the are the most commonly known hosts ofmermithid
&
limestone Alps. The type locality (Kandersteg, can- worms (Penney Bennett2006),anditseems likely
ton Bern) is at the centre ofthe known distributi- thatparasite-inducedmalformations arethebasisfor
on ofthe alpine population ofP. rupicola and too far otherphantom species in this genus aswell.
&
west forA.pedestris (Buchar Thaler 1993), but an
identification based solely on a zoogeographical ar- Philodromus depriesteri Braun, 1965 = nomen dubi-
gument seems tenuous.The otherAcantholycosa spe- um(Philodromidae)
cies are more distinctly coloured,while the similarly This species, a member of the Philodromus aureolus
builtPardosanigra (C.L. Koch, 1834) is describedin group,was first described from twowidely separated
detailbyLebert and reported from numerous locali- localities (Krimml, Austria, and Geisenheim, Ger-
ties, thus can probably be excluded as an alternative many), separated by 600 km including the German
synonym. Alps. Nonetheless, despite its presumably extensive
range and much increased collecting activities in
Pardosa intermedia (Bösenberg, 1903) = nomen du- the last decades, the species has never been found
bium(Lycosidae) again since its description 50years ago.The reason is
Like manyother types ofBösenberg,the material of probably that the two female types preserved in the
this species,descirbed as Lycosa intermedia,was dest- Senckenberg Museum Frankfurt are malformed (or
royedinWorldWarII (Renner 1988).Intheoriginal subadult) specimens with incompletely developed
description,Bösenbergremarkedthatthe specieshas epigynes (Jäger pers. comm.). Vulval structures very
an intermediate position between Pardosa agrestis similar to those ofP depriesteri were observed in a
,
Pardosa albatula Pardosa monticola and Pardosapa- female Philodromusofthe aureolusgroup that turned
,
lustris. Given the general difficulties of identifying out to be infectedbya parasiticworm (Mermithidae
females of the monticola group, it is impossible to or Nematomorpha; Fig. 2).This specimen was coll-
identify the species solely based on the description ected bybeating the field layer ofawet meadow,to-
and figures. In particular, an identification with P getherwithtypicalspecimens ofPhilodromuscollinus.
palustris proposed by Simon (in Bösenberg 1903) It therefore seems verylikely that the unusual geni-
and Bonnet (1958) seems unlikely, given the high- talia are the result ofa parasite-induced malformati-
ly characteristic epigyne ofthat species and the fact on, comparable to the case ofLepthyphantes beckeri.
that Kulczyriski examinedthe type and considered it Nematode infections have beenrepeatedlydescribed
a new species. Possibly the epigyne ofthe specimen in spiders (Meyer 2014 and references therein), and
Fig.2: Parasitized Philodromusfemale (left),collected intheAllgäu region, Bavaria,Germany, in August2012. Itsepigyne (centre,
dorsalview)lacksreceptaculaandafullydevelopedmedianseptum,justasseeninP.depriesteri,andduringthegenitalpreparation
aparasiticworm(right)about10cminlengthwasdetected.(PhotographscourtesyofStefan Rehfeldt)
74 R.Breitling,M.Lemke, TBauer,M.Hohner,A.Grabolle&T.Blick
it has been speculated before that they might be the ety of Philodromus aureolus, and this seems indeed
causeofgenitalmalformations (Martin2013).Braun the most likely identification, based on the figures
(1965) alreadyrecognized the possibilitythat his ty- in both the original description and the later re-de-
pes were aberrant specimens, but nonetheless deci- scription by Bösenberg (1902). Bertkau (1880) had
ded to describe them as a new species, even though already considered Philodromus micans as the male
the highly unusual genital morphology (lacking re- of P aureolus. However, given that Muster 6cTha-
ceptacula and median septum, which are otherwise ler (2004) tentatively assign the male illustrated by
uniformly present and distinct in all species of the Bösenberg to the closely related Philodromus buchari
group) would require a radically divergent pedipalp Kubcovä, 2004, it seems currently impossible to un-
structure and mating behaviour, which is inconsis- ambiguouslyidentifyMenge’s species.
tentwith thegeneralconservative trendswithinPhi-
lodromus. Such a dramatic divergence would be par- Phlegrarogenhoferi(Simon,1868) =Phlegraeinereofa-
ticularlyunexpected as Braunidentifies aclear“sister sciata (Simon, 1868) syn.nov.(Salticidae)
species”,PhilodromuscollinusC.L.Koch,1835,which This species was described based on a single male
is highly similar in all non-genitalic characters. The collected by Octavius Pickard-Cambridge in Ba-
reason for considering P. depriesteri as a nomen du- den (close to Vienna,Austria) during a trip through
bium, instead ofa synonym ofP. collinus lies in the Europe and probably passed on to Eugene Si-
,
overall similarityoffemales in the aureolusgroup. mon during his subsequent visit to Paris (Pickard-
Cambridge 1918). According to Kulczynski (1898)
Philodromus dispar obscurus Lebert, 1877 = nomen the species is very close (and possibly identical) to
dubium(Philodromidae) P.fuscipes Kulczynski, 1891, currently considered a
This form was described by Lebert (1877: 271) as a junior synonym of P cinereofasciata (Simon, 1868).
melanisticvarietyofPdispar,foundintheUrserental, More recently, the species was discussed by Stefania
Switzerland, at an altitude of1500 m. Later authors H^ciak in her unpublished PhD thesis (H^ciak, ca.
eitherignoredthisvarietyorconsidereditasynonym 1983),based on material from “Galicia Vallombrosa”
ofthe nominate form (Lessert 1910).The descripti- (probablyin Spain orpossiblyItaly) in the collection
onofPd. obscurusis,however,impossibletoreconcile ofthe Museum National d’Histoire Naturelle,Paris.
with P dispar, the male is described as having enti- However, it is unlikely that this material is part of
rely dark brown legs with black margins (“mit ganz the type series ofP rogenhoferi, and it quite possibly
dunkelbraunen, schwarz berandeten Beinen”) and belongs to a different species. Despite Proszynski’s
white-grey spots and a grey transverse band on the claim (2014) that “Simon’s 1937: 12661267 re-
blackopisthosoma.Thefemaleis evenmore different mark on occurrence [in Austria and once in Landes,
(“weicht besonders ab”), and is described as entirely France] must be wrong”, it is all but inconceivable
dark,andlargerandmoremassivethanthe nominate that Simon, whose excellent memory was legen-
form. The alpine location suggests that Lebert may dary (Savory 1961), would have confused material
actuallybe describingdarkspecimens ofPhilodromus obtained from Pickard-Cambridge at the “summit
vagulus(Simon, 1875),ahigh-altitude species thatis meeting”ofthe GoldenAge ofarachnology.This in-
reasonably similar in general habitus to Pdisparand terpretation is confirmed by the presence ofa speci-
has a similarly elongated epigyne, but is darker, lar- men from Baden in Pickard-Cambridge’s collection
ger, and without distinct sexual dimorphism. How- in Oxford (vial 1744.7); it is not clear ifthis is the
ever, the description is so vague and the number of type described by Simon,but the specimen certainly
remaining discrepancies so large that, in the absence belongs to the same series. Geographical arguments
oftype material, it seems prudent to consider P dis- support the identification ofP. rogenhoferiwith Pci-
parobscurusas a nomen dubium. nereofasciata. The latter is the only species ofPhlegra
sufficientlysimilarto the species describedbySimon
Philodromus micans Menge, 1875 = notnen dubium occurring in the wider vicinity of the type locality.
(Philodromidae) Perhaps even more importantly, the first published
As for most ofthe species described by Menge, the record for Austria we are aware ofcame from a xe-
type material of P micans is probably lost (Kraus rothermic hillside in easywalking distance ofBaden
2009). The form was originally described as a vari- (just 10 km to the north ofBaden city centre; Franz