The first Permian Diaphanopterodea (Insecta, Megasecopteromorpha) from China
Author
Yang, Nan
https://orcid.org/0009-0000-2687-3916
College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, China
Author
Cui, Yingying
https://orcid.org/0009-0008-2544-2821
College of Life Sciences, South China Normal University, Guangzhou, China
Author
Xu, Ziqiang
0000-0002-2682-2550
College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, China
Author
Xu, Yanqi
Geological Survey of Jiangsu Province, Jiangsu, China
Author
Ren, Dong
0000-0001-8660-0901
College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, China
Author
Béthoux, Olivier
0000-0002-3178-8967
CR 2 P (Centre de Recherche en Paléontologie – Paris), MNHN – CNRS – Sorbonne Université, Paris, France
text
Fossil Record
2024
2024-08-15
27
2
247
258
journal article
10.3897/fr.27.e128892
E93DEB3D-DFFF-4BE5-8B06-E79DB3F271F7
Family
Parelmoidae
Rohdendorf, 1962
Type genus.
Parelmoa
Carpenter, 1947
.
Included genera.
Diapha
Kukalová-Peck, 1974
;
Elmodiapha
Kukalová-Peck, 1974
;
Paradiapha
Kukalová-Peck, 1974
;
Permelmoa
Prokop & Nel, 2011
;
Permodiapha
Kukalová-Peck, 1974
;
Permuralia
Sinichenkova & Kukalová-Peck, 1997
;
Protodiapha
Kukalová-Peck, 1974
;
Pseudelmoa
Carpenter, 1947
;
Stenodiapha
Kukalová-Peck, 1974
;
Sinoelmoa
gen. nov.
Commented diagnosis.
ScP long, ending beyond the first fork of
RP
(plesiomorphy within
Diaphanopterodea
); near wing base, shortly after its origin, CuA fused for some distance with, or running closely along,
R
+ M (apomorphy; as currently documented, shared with all
Diaphanopterodea
except
Sinodiaphidae
, in which CuA is connected with M by a short cross-vein, and
Diaphanopteridae
, in which the connection of CuA and M is very brief); first cross-vein in the CuA – CuP area very short and oblique, with CuA displaying a clear inflexion at the point of connection with this cross-vein [also present in
Diaphanopteridae
,
Carrizodiaphanoptera
and, to some extent,
Elmoidae
(Fig.
3
); possibly an apomorphy of the entire
Diaphanopterodea
– except for
Sinodiaphidae
, in which this state is primarily absent –, and with presumed secondary losses in various families, such as
Martynoviidae
];
MP
branched (plesiomorphy within
Diaphanopterodea
); CuP with 1–3 distal branches (plesiomorphy within
Diaphanopterodea
); developed anal area (putative apomorphy for the family).
Remarks.
The combination of (i) an overall rich venation, (ii) a long fusion of CuA with
R
+ M (or, CuA running very close to
R
+ M for some distance), and (iii) a cua-cup cross-vein very short, is generally used to identify members of the
Parelmoidae
. The Pennsylvanian
Diaphanopteridae
differ from this family only by lacking character state (ii) (see
Béthoux and Nel 2003
). Given the polarity of several character states listed as diagnostic of the family, it is not excluded that this taxonomic concept might represent a paraphyletic entity, to include other
Diaphanopterodea
families, such as
Elmoidae
.
Carpenter (1992)
considered the six genera and 10 species reported from the Obora locality (Permian, Cisuralian, Sakmarian), and originally assigned to the
Elmoidae
or
Parelmoidae (
Kukalová-Peck 1974
)
, as of uncertain Parelmoidael affinities. Addressing aspects of species delimitation, and the systematics of these species, is made difficult by post-depositional deformations this material endured. Nevertheless, wing venation character states they display tend to indicate a placement to the family
Parelmoidae
as delimited above, in particular the well-developed anal area. Also, despite deformation, this comparatively large sample allows appreciating variation in character states variability (in other words, how the extent of variability varies within families), likely to differ among (and within) the closely related families
Diaphanopteridae
,
Parelmoidae
and
Elmoidae
(see below and
Kukalová-Peck and Sinichenkova 1992
).
Notably, a distal fork of CuP is common in the Obora material, with some specimens displaying an early fork, and even a 3 - branched CuP. The relation between
RP
and MA is also very variable across the corresponding species, ranging from a complete lack of fusion to a long one. In contrast, species of the genus
Parelmoa
Carpenter, 1947
(Fig.
2 A – D
) show more stable venational features (CuP simple; MA and
RP
connected by a short cross-vein). Among other
Parelmoidae
, the monotypic genera
Pseudelmoa
Carpenter 1947
(Fig.
2 E
) and
Permuralia
Sinichenkova & Kukalová-Peck, 1997
(Fig.
2 F
) remain similar to
Parelmoa
spp.
in most of their venational features. Also,
Elmoa trisecta
Tillyard, 1937
(
Elmoidae
; Fig.
3
) shows rather stable venational features, with a consistent occurrence of a (i) distally forked CuP and (ii) a MA distinct from
RP
, with a cross-vein connecting the two veins shortly after the origin of the latter. The Obora material is therefore unusual in several respects. Instead of attempting to finely resolve relationships between the corresponding taxa, whose variability in many aspects cannot be properly appreciated, we believe it is more sensible to use the
Parelmoidae
as a broad taxonomic concept, possibly paraphyletic (i. e., a grade), to include the Obora material.