Superfamily
Tenthredinoidea Latreille, 1802
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Family
Xyelotomidae Rasnitsyn, 1968
Type genus:
Xyelotoma Rasnitsyn, 1968
(type species
X. nigricornis Rasnitsyn, 1968
, Upper Jurassic, Callovian–Kimmeridgian or Oxfordian–Kimmeridgian ( Zherikhin and Gratshev 1993; Mostovski and Martínez−Delclòs 2000), Karatau, Chimkent region, Southern Kazakhstan).
Included genera.—
Xyelocerus Rasnitsyn, 1968
(type species
Xyelocerus admirandus Rasnitsyn, 1968
, Upper Jurassic, Callovian–Kimmeridgian or Oxfordian–Kimmeridgian, Karatau, Chimkent region, Southern Kazakhstan).
Pseudoxyela Rasnitsyn, 1968
(type species
P. heteroclita Rasnitsyn, 1968
, Upper Jurassic, Callovian–Kimmeridgian or Oxfordian–Kimmeridgian, Karatau, Chimkent region, Southern Kazakhstan).
Undatoma Rasnitsyn, 1977
(type species
U. dahurica Rasnitsyn, 1977
, Upper Jurassic/Lower Cretaceous, Unda river, Chita Province, Transbaikalia (Far Eastern Russia); other species:?
U. undurgensis Rasnitsyn, 1990
,?
U. taksha Rasnitsyn, 1990
, both from the Lower Cretaceous of Transbaikalia,
U. rudwickensis Rasnitsyn and Jarzembowski, 1998
, Lower Cretaceous, Lower Barremian, West Sussex, England,
U. stigmatica Rasnitsyn and Jarzembowski, 1998
, Lower Cretaceous, Upper Berriasian, Dorset, England,
U. bicolor Rasnitsyn and Jarzembowski, 1998
, Lower Cretaceous, Upper Hauterivian, Surrey, England).
Liaotoma Ren et al., 1995
(type species
L. linearis Ren et al., 1995
, Late Jurassic or Lower Cretaceous?, Liaoning Province, China). Ren et al. (1995: 194, text−fig. 3−69) only compared this taxon with
Dahurotoma Rasnitsyn, 1990
and “
Protenthredo
” Hong, 1982. In fact,
L. linearis
shows no significant difference with
U. rudgwickensis
, except, maybe, in the shape of the pterostigma, which is apparently narrower in
L. linearis
than in
Undatoma
. A revision of the type specimen of
L. linearis
is necessary in order to determine whether Liadotoma is a junior synonym of
Undatoma
.
Dahurotoma Rasnitsyn, 1990
(type species
D. robusta Rasnitsyn, 1990
, Lower Cretaceous, Transbaikalia).
Leridatoma Rasnitsyn and Ansorge, 2000
(type species
L. pulcherrima Rasnitsyn and Ansorge, 2000
, Lower Cretaceous, Spain).
Davidsmithia Pagliano and Scaramozzino, 1990
, one species
D. suni ( Hong, 1982)
, Upper Jurassic/Lower Cretaceous, Guyong Basin, Neimenggol Region, China.
D. suni
was originally described in the genus
Protenthredo Hong, 1982
, as the type genus of the fossil family
Protenthredinidae Hong, 1982
. As the generic name
Protenthredo
was preoccupied by
Protenthredo Pongracz, 1928
(in
Tenthredinidae
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), Pagliano and Scaramozzino (1990: 5, line 10) proposed a generic name
Davidsmithia
for this taxon, but the same authors proposed the genus name
Thomasia
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in the same page, line 45. Abe and Smith (1991) noted the problem but did not solve it. We propose as the valid name the first one:
Davidsmithia
. Rasnitsyn and Ansorge (2000: 59) erroneously indicated that Pagliano and Scaramozzino proposed the new name
Xaxexis
to replace
Protenthredo Hong, 1982
, but in fact these last authors (1990: 6) proposed
Xaxexis
to replace
Euryxyela Hong, 1984
. Abe and Smith (1991) put
Protenthredo Hong, 1982
in
Xyelotomidae
, without further explanation, and did not synonymize the
Protenthredinidae
with the
Xyelotomidae
. A revision of the type specimen is necessary before any definite conclusion about the position of this enigmatic genus.
Vitimilarva Rasnitsyn, 1969
(type species
V. paradoxa Rasnitsyn, 1969
, Early Cretaceous, Baissa, Vitim river, Transbaikalia). This species is based on a larva that Rasnitsyn (1969) put in “Tenthredinidea incertae sedis ” but that was transferred latter in to the
Xyelotomidae
( Rasnitsyn 1990; Abe and Smith 1991).
Kuengilarva Rasnitsyn, 1990
(type species
K. inexpectata Rasnitsyn, 1990
, Lower Cretaceous, Transbaikalia). This taxon is also based on a larva that Rasnitsyn (1990) attributed to the
Xyelidae
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or
Xyelotomidae
.
Remarks on the family composition.— Rasnitsyn (1988: 120, fig. 1; 1996: 14; 2002: fig. 331) clearly considered the
Xyelotomidae
as a paraphyletic, “ancestral” group of the other
Tenthredinoidea
, with no recorded autapomorphies. At least, the three genera
Undatoma
,
Liaotoma
,
Leridatoma
, and
Davidsmithia
have their forewing vein Sc “not developed, at least not as a longitudinal trunk” ( Rasnitsyn 1977 —English translation in 1978, 1990; Hong 1982: 93, fig. 19; Ren et al. 1995: 194, text−fig. 3−69; Rasnitsyn et al. 1998). This character state is currently supposed to be the unique apomorphy in the wing venation of the
Tenthredinoidea
minus
Xyelotomidae
( Rasnitsyn 1988, 2001; Ronquist et al. 1999; Schulmeister, 2003). Thus, it would support the possible exclusions of
Undatoma
,
Liaotoma
,
Leridatoma
, and
Davidsmithia
from the
Xyelotomidae
. The longitudinal part of Sc is absent or very weak and Sc1 and Sc2 are absent in
Xyelocerus admirandus
(after Rasnitsyn 1968: 226, text−fig. 35, 1996: fig. 5). On the contrary, the genus
Pseudoxyela
has “retained” Sc and R separated and parallel, and the forewing Sc is basally free and distally fused with R in
Dahurotoma
and probably
Xyelotoma
.
The structure of the prosternum (character 34, state 1 of Ronquist et al. 1999) is a potential apomorphy of the
Xyelotomidae
but it is present in several other hymenopteran groups (e.g.,
Megalyridae
and
Stephanidae
). Furthermore, it is unknown in several species currently attributed to the
Xyelotomidae
. At least in
Undatoma
and
Thomasia
, the body characters are too badly known and cannot be used in a phylogenetic analysis. Ren et al. (1995) did not describe the body structures of
Liaotoma
. This suggests that the
Xyelotomidae
could be an artificial group of taxa with no direct common ancestor.
Only a detailed phylogenetic revision of the
Xyelotomidae
and the
Tenthredinoidea
will help to solve the problems in the definition of the
Xyelotomidae
and the positions of the genera
Undatoma
,
Liaotoma
,
Leridatoma
, and
Davidsmithia
.
Lastly, it is not possible to have a correct proof of the attribution of
Vitimilarva
and
Kuengilarva
to the strictly fossil family
Xyelotomidae
, because the hymenopteran larvae are very different from the adults.