Revision of F. R. C. Reed’s Ordovician trilobite types from Myanmar (Burma) and western Yunnan Province, China
Author
Fortey, Richard A.
Department of Earth Sciences, Museum of Natural History, Cromwell Road, London, SW 7 5 BD, UK.
Author
Wernette, Shelly J.
Department of Earth and Planetary Sciences, University of California, Riverside, CA 92521, USA. & Department of Geography and Environmental Studies, Texas State University, San Marcos, TX 78666, USA
Author
Hughes, Nigel C.
text
Zootaxa
2022
2022-07-08
5162
4
301
356
journal article
93239
10.11646/zootaxa.5162.4.1
7d343017-483e-43be-9439-d06e748fc54e
1175-5326
6810290
DD2279FA-E8F1-4951-A5CA-91082E875580
Subfamily
Birmanitinae
Kobayashi, 1960
Discussion.
Birmanites
Sheng, 1934
, with
type
species
Ogygites birmanicus
Reed, 1915
, was originally described as an asaphid, but was placed in a separate family
Birmanitidae
by
Kobayashi (1960
, p. 254) and in
Dikelokephalinidae
in the 1959
Treatise on Invertebrate Paleontology
(Harrington
et al. in
Moore, 1959
).
Lu (1975
, p. 319) regarded
Birmanites
as no more than an unusually flat asaphid, with a relatively small glabella, an opinion with which we concur; the taxon
Birmanitidae
has disappeared from the literature. It is, however, still available as a subfamily name within
Asaphidae
. Division of
Asaphidae
into subfamilies is itself contentious; a relatively conservative approach was used by Jaanusson (
in
Moore, 1959
), who employed seven subfamilies. The opposite approach was that of
Balashova (1976)
who elevated
Asaphidae
to subordinal level and divided the group into numerous families and subfamilies. Balashova nominated subfamilies centred on well-known Baltic endemic genera such as
Megistaspis
Jaanuson, 1956
,
Pseudoasaphus
Schmidt, 1901
and
Ptychopyge
Angelin, 1854
, to each of which she consigned a number of her own, finely divided genera. This is not an appropriate place to review this work of prolific taxonomic ‘splitting’, although the majority of Balashova’s taxa are confined to the Ordovician Baltic Shield, and her scheme has not been widely adopted. However,
Balashova’s (1976)
Subfamily
Nobiliasaphinae
(placed in Family
Pseudoasaphidae
Balashova, 1969
, by
Balashova [1976]
) includes the genera
Opsimasaphus
Kielan, 1960
, and
Nobiliasaphus
Přibyl & Vaněk, 1965
, both of which are closely similar to
Birmanites
, and must surely be included within the same subfamily, whatever taxonomy is eventually adopted. The name
Birmanitinae
Kobayashi, 1960
is the older name for this group even though it was not recognised by
Balashova (1976)
.
An objective appraisal of asaphid trilobites is overdue. An unpublished PhD thesis by Mark
Bell (2009)
under the supervision of RAF attempted a cladistic analysis of
Asaphidae
. One group that comprised a clade with moderate support included
Birmanites
,
and the name
Birmanitinae
is appropriate for it. The most important characters uniting the group are displayed by the hypostome, which has a characteristic rounded to oval middle body with prominent maculae at the rear, and a broad, U-shaped fork, and hypostome widest at, or in front of the maculae. The characteristic hypostome is illustrated by a specimen from the type series of
Birmanites birmanicus
figured herein (
Fig. 6.1
) and an illustrative reconstruction (
Fig. 7.b
), and by the Barrandian
lectotype
(selected
Kielan, 1960
, p. 76) of
Nobiliasaphus nobilis
(see
Horný & Bastl, 1970
, pl. 6 fig. 1;
Rabáno, 1989
, pl. 3, fig. 6; also,
Hughes, 1979
, fig. 11 for a similar hypostome, and
Hammann & Leone 1997
, pl. 6, fig. 3 and
Turvey, 2007
for further examples). Dorsal characters in the group are variable with regard to width of pleural areas, preglabellar field, dorsal effacement and the like, but a glabella with an inflated, often pyriform frontal lobe and with a pair of posteriorly inwardly sloping, and often deepened axial/glabellar furrows which constrict a narrow median glabellar lobe in front of the occipital ring, is also typical. Where known, the medial tubercle is immediately pre-occipital (
Birmanites
—see
Lu, 1975
, pl. 8, fig. 8), and the dorsal facial sutures run at, or at least very close to the anterior margin before they meet medially. A distinctive genus from central
Australia
,
Norasaphus
Fortey & Shergold, 1984
, which includes the only tuberculate asaphids, also belongs within the same group, showing similar hypostome construction and glabella furrows, even though it is much more convex and less flattened peripherally than
Birmanites
.
Turvey (2007)
added further Chinese taxa to this group.
Nobiliasaphus
is distinguished from
Birmanites
in having a curious structure of the pygidial axis, with chevron-like medial backward deflections of the ring furrows, the ring itself bisected transversely by fainter extra furrows and ridges of similar form.
This asaphid group is peri-Gondwanan, with scattered occurrences from
France
, Iberian Peninsula and
Wales
eastwards to southwest
China
. The genus
Pamirotetchites
Balashova, 1966
, from the Pamirs in central Asia is a junior synonym of
Nobiliasaphus
as
Balashova (1976
, p. 59) realised, and displays the same distinctive pygidial structure as
N. nobilis
(Barrande)
(see also
Opsimasaphus pseudodawanicus
Lu, 1975
,
in
Turvey, 2007
).
Norasaphus
is so far confined to
Australia
, while
Nobiliasaphus
does not extend east of the Pamirs.
Birmanites
is more widespread. Whether this group should be included within a more inclusive clade based on
Pseudoasaphus
(as claimed by
Balashova, 1976
) is uncertain. Many genera of
Asaphidae
are known to be endemic to Baltica, such as those placed in Megistaspidinae by
Balashova (1976)
, and
Pseudoasaphus
may be no exception.
Pseudoasaphus
spp.
have anterior branches of the facial sutures which cross the fontal area well away from the anterior cephalic margin before meeting in an acute point (e.g. Jaanusson, 1953, pl. 4, fig. 3), and hypostomes attributed to related genera are relatively long and narrow, with more extended ‘forks’ subtending an almost v-shaped outline (
Balashova, 1976
, pl. 3).