New and already described fossil spiders (Araneae) of 20 families in mid and late Cretaceous Burmese Ambers, with notes on spider phylogeny, evolution and classification
Joerg Wunderlich, Patrick Müller
An emended classification and cladograms of high taxa of the Aeaneae
are provided. Ca. 48 families of spiders (Araneae) in Mid Cretaceous Burmese amber
are listed; the report of the families Atypidae, Idiopidae and Ochyroceratidae is doubtful.
The spiders of the Infraorder MYGALOMORPHA are treated, a provisional key to
its families is given. The 9 superfamilies of the diverse clade SYNSPERMIATA are
diagnosed, 20 families, 11 are known in Burmite, 5 are extinct. Synspermiata contains
the three subclades Caponiomorpha, Dysderomorpha and Pholcomorpha. Ca. three
quarters of haplogyne families in Burmite are members of the Synspermiata. I call the
Cretaceous the “age of Synspermiata and Palpimanoidea (= Archaeoidea)”. The colulate
and basically eight-eyed Synspermiata is actually not any more regarded as sister
group of the family Filistatidae but of the clade CRIBELLATAE in a new sense which
includes several diverse branches. Hence the name Basalhaplogynae WUNDERLICH
2019 is superfluous (name rejected). In my opinion the cribellum originated only once.
The taxon Microsegestriinae WUNDERLICH 2004 (under Segestriidae) in Lebanese
amber is now regarded as a questionable member of the superfamily Dysderoidea,
probably Oonopidae (quest. n. relat.). Segestrioidea is split off from the Dysderoidea
and contains three families. The family Ariadnidae WUNDERLICH 2004 (n. stat., elevated
from Ariadninae) is unknown in Burmite. Jordansegestria WUNDERLICH 2015 with its generotype J. detruneo in Jordanian amber is regarded as a synonym of Parvosegestria
WUNDERLICH 2015 of the new family Parvosegestriidae (n. syn. & n.
relat.), based on Parvosegestria WUNDERLICH 2015. The genus Denticulsegestria
WUNDERLICH 2015 is regarded as a synonym of Parvosegestria (n. syn.). The synonymy
of Myansegestria WUNDERLICH 2015 with Parvosegestria is not excluded (n.
quest. syn.). Magnosegestria tuber n. gen. n. sp. - a questionable member of the
Segestriidae - in Burmite is described. The holotype of Magnosegestria tuber is considered
to be the prey of a mygalomorph spider. Burmorsolidae WUNDERLICH 2015
(n. stat.) (from Burmorsolini) is transferred from the Plumorsolidae to the new superfamily
Burmorsoloidea (n. relat.). With some hesitation the extant family Trogloraptoridae
GRISWOLD et al. 2012 from Noth America is regarded as related to the Burmorsolidae
(quest. n. relat.). Loxodercinae WUNDERLICH 2017 has previously been transferred
from the family Eopsilodercidae WUNDERLICH 2008 to the Segestriidae:
Segestriinae but is now regarded as a synonym of the Burmorsolidae (n. syn.).
Burmorsolidae (under Burmorosolini) has erroneously been described by WUNDERLICH
(2015) as a member of the family Plumorsolidae WUNDERLICH 2008. Plumorsolidae
is known in Lebanese amber but is unknown in Burmese amber. This family is
now regarded as a plesion, probably of the branch Dipneumonomorpha: Oecobioidea?
(quest. n. relat.). Plumorsolidae in the previous sense of WUNDERLICH is not a monophyletic
taxon: Loxoderces WUNDERLICH 2017 and Pseudorsolus WUNDERLICH
2017 are regarded as junior synonyms of Burmorsolus WUNDERLICH 2015 (n. syn.)
of the family Burmorsolidae. – The relationships of the family Copaldictynidae
WUNDERLICH 2004 n. stat. of the new taxon Agelenomorpha in subrecent COPAL
FROM MADAGASCAR is revised and regarded as related to the extant families Titanoecidae
and Nicodamidae of the Nicodamoidea (n. relat.). - The following FURTHER NEW
TAXA are described (Burmorsoloidea and Segestrioidea: See above): Ctenizidae: Parvocteniza
parvula n. gen. n. sp.; Nemesiidae: Burmesia sordida n. gen. n. sp., Myannemesia
glaber n. gen. n. sp.; Theraphosidae: Protertheraphosinae n. subfam. based
on Protertheraphosa spinosa n. gen. n. sp.; Oonopidae: Burmorchestina circular n.
sp.; Burmorsolidae: Burmorsolus: globosus n. sp., longembolus n. sp. and longibulbus
n. sp.; Eopsilodercidae: Propterpsiloderces crassitibia n. sp., P. cymbioseta n. sp., P.
duplex n. sp.; Psilodercidae: Priscaleclercera furcate n. sp., P. hamo n. sp., P. liber
n. sp.; the family Aliendiguetidae n. fam., a plesion probably close to the Ochyroceratoidea
and Plectreuroidea, based on Aliendiguetia praecursor n. gen. n. sp.; Praepholcidae
n. stat., from Eopsilodercidae: Praepholcinae: Hamoderces opilionoides n.
gen. n. sp.; Tetrablemmidae: Bicornoculus granulans n. sp., Cymbioblemma fusca n.
sp., C. hamoembolus n. sp., Electroblemma bifurcate n. sp., E. caula n. sp., E. pinnae
n. sp., Eogamasomorpha rostratis n. sp., Unicornutiblemma n. gen., U. brevicornis n.
gen., U. gracilicornis n. sp., U. longicornis n. sp.; Hersiliidae: ?Burmesiola kachinensis
n. sp.; Archaeidae: ?Burmesarchea bilongapophyses n. sp.; Pholcochyroceridae:
Spinicreber vacuus n. sp.; Praearaneidae: Praearaneus araneoides n. sp.; Zarqaraneidae:
Palazarqaraneus hamulus n. gen. n. sp., Paurospina fastigata n. sp. ?Baalzebub
mesozoicum PENNEY 2014 from the Late Cretaceous OF FRANCE (under Theridiosomatidae)
is transferred to the family Zarqaraneidae (n. relat.) and regarded as the
member of an undescribed genus. – A note on Burmese Tilin amber is added.
Most important results of my studies (most are based on fossil spiders, too):
(1) The orb web originated twice: First in the Deinopoidea (remains of a Cretaceous
cribellate orb web exists) and - probably distinctly - later in the ecribellate Araneoidea
(no sure proof of an orb weaving araneoid taxon exists in the Cretaceous in constrast
to members of irregular web dwellers like Theridiidae and Zarqaraneidae); (2) the irregular (space) capture webs within the superfamily Araneoidea did not originate
from an orb web. They are not derived but are ancient web types and an extinct
araneoid species gave rise to the orb web;
(3) the cribellum originated only once. The Dipneumonomorpha retained basically a
DIVIDED cribellum - divided in the Family Filistatidae, divided, entire or lost in the remaining
Dipneumonomorpha; it is secondarily entire in the Hypochilomorpha (see
fig. A). (4) Losses (e. g.): Tarsal and metatarsal trichobothria were lost numerous times during
spider evolution (like the cribellum, book lungs, feathery hairs, leg bristles and the anterior
(5) the RTA-clade may be close to the Deinopoidea (see fig. A).
(6) the predecessor of the Synspermiata did PROBABLY not possess a cribellum – in
this clade the anterior median spinnerets were directly transformed to a colulus (see
figs. A, C).
(7) The Cretaceous - and apparently already the Jurassic - was the era of haplogyne
spiders, mainly of the Synspermiata (fig. C) and the - really haplogyne? – Palpimanoidea
(= Archaeoidea) at least in higher strata of the vegetation.
(8) Not a single sure proof of a Cretaceous member of the Retrolateral Tibial Apophysis
(RTA)-clade in Burmite exists. Members of this clade – e. g. Jumping spiders (Salticidae)
and Wolf spiders (Lycosidae) – are the most frequent and the most diverse spiders
today besides members of the superfamily Araneoidea.