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ALLOCHTHONOUS SPHENOTHALLUS (CNIDARIA) FROM A LACUSTRINE

LAGERSTÄTTE, CARBONIFEROUS OF NEW MEXICO, USA

Article · January 2011

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Allan J Lerner Spencer G. Lucas

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 Sullivan et al., eds., 2011, Fossil Record 3. New Mexico Museum of Natural History and Science, Bulletin 53.

86
ALLOCHTHONOUS SPHENOTHALLUS (CNIDARIA) FROM A
LACUSTRINE LAGERSTÄTTE, CARBONIFEROUS OF NEW MEXICO, USA

ALLAN J LERNER AND SPENCER G. LUCAS

New Mexico Museum of Natural History, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104

Abstract—New Mexico’s only record of Sphenothallus, a putative cnidarian of Paleozoic age, is from an unusual
Konservat-Lagerstätte deposit in the Tinajas Member of the Atrasado Formation (Carboniferous: Upper Pennsyl-
vanian Series, Kasimovian Stage) at the Tinajas Arroyo locality, Cerros de Amado, Socorro County. The Tinajas
Arroyo Sphenothallus record is a unique, allochthonous occurrence of this marine animal in a lacustrine environ-
ment that had intermittent connection to lagoonal/shallow marine waters. Sphenothallus adds to a recently aug-
mented and important record of Paleozoic cnidarians from New Mexico.

INTRODUCTION
Sphenothallus has undergone numerous changes in phylogenetic
placement since it was first described by Hall (1847), who interpreted it
as a plant resembling Calamites. Since then its identity or affinities have
been variably considered conulariid, scyphozoan, hydrozoan, tubiculous
annelid, or other worm (e.g., Ruedemann, 1898; Price, 1920; Mason and
Yochelson, 1985; Fauchald et al., 1986). Other authors have come to
regard Sphenothallus as incertae sedis at the family to phylum level (e.g.,
Wang et al., 2003; Li et al., 2004). The exact phylogenetic placement of
Sphenothallus at present remains unsettled, but we follow the assign-
ment of Van Iten et al. (1992), who interpreted Sphenothallus as a prob-
lematic fossil cnidarian and hypothesized homology of its test with the
theca of polypoid hydrozoans or scyphozoans.
Zhu et al. (2000), in a taxonomic review of Paleozoic phosphatic
tubular fossils, provided an emended generic diagnosis of Sphenothallus.
It was described as consisting of an apatitic, finely lamellar tube with a
basal attachment disk and two longitudinal thickenings at the end points
of the tube’s widest diameter. Sphenothallus is inferred to have been a
gregarious, sessile predator (Peng et al., 2005). It typically occurs in low-
energy, dysaerobic marine environments (Van Iten et al., 1992).
Sphenothallus has a stratigraphic range of Cambrian to Permian and has
published records from North America, South America, Europe, and
southern China (Zhu et al., 2000). It is one of the numerous animals to
have developed a biomineralized (calcium phosphate) skeleton during
the Cambrian Period (Li et al., 2004; Peng et al., 2005).
In this paper, NMMNH refers to the New Mexico Museum of
Natural History and Science, Albuquerque, New Mexico.
THE TINAJAS ARROYO LOCALITY
New Mexico’s only record of Sphenothallus comes from NMMNH
locality 4667, known as the Tinajas Arroyo locality, which is located in
the Cerros de Amado, near Tinajas Arroyo in Socorro County (Fig.1).
Lerner et al. (2009a) provided a detailed study of the geology, sedimen-
tology, and biota of the Tinajas Arroyo locality and discussed the paleo-
ecology of the site. Strata at NMMNH locality 4667 are assigned to the
Tinajas Member of the Atrasado Formation, which is inferred to be of
Carboniferous (late Kasimovian) age (equivalent to late Missourian age
of historic North American usage; Heckel, 2008) based on conodont
stratigraphy (Lerner et al., 2009a; Lucas et al., 2009).
A 5.2- to 7-m-thick laminar black shale bed (Fig. 1, unit 6) contain-
ing a prolific conchostracan assemblage occurs near the base of the mea-
sured section at NMMNH locality 4667. A single Sphenothallus speci-
men was collected from this unit, which also contains a diverse fossil
assemblage of crustaceans, insects, molluscs, fishes, and plants. Several
taxa within this assemblage, including Sphenothallus, were unknown
from the Carboniferous of New Mexico until reported from the Tinajas FIGURE 1. Location map and measured stratigraphic section at the Tinajas
Arroyo locality. Lerner et al. (2009a) considered the black shale to have Arroyo locality (from Lerner et al., 2009a).

been primarily deposited in a fresh-to-brackish water lake with anoxic-
dysoxic bottom waters favoring organic preservation. A lacustrine origin
of the black shale is supported by the abundance of conchostracans;
presence of darwinuloid (freshwater) ostracods, land plants, and insect
wings; lack of marine indicators; and the lithology and geometry of the
deposit. Lerner et al. (2009a) concluded that the lake was formed in a
coastal setting and had an intermittent connection to lagoonal/shallow
marine waters, as indicated by the presence of taxa (e.g., Sphenothallus
and some fishes) within the assemblage that normally inhabited saline
waters. Indeed, the lake deposit is directly overlain by marine shale with
Dunbarella capped by marine limestone (Fig.1 ), and this suggests that
the Tinajas locality represents a coastal lake that was transgressed over
by the advancing marine shoreline (Lerner et al., 2009a). No other occur-
rence of a similar paleoenvironment is known from the Carboniferous of
New Mexico. Similar Lagerstätten deposits and biota have been de-
scribed from the Carboniferous (Upper Pennsylvanian Series) to Per-
mian (Cisuralian Series) of Kansas (e.g., Dunbar, 1924; Babcock et al.,
2000), but Sphenothallus is unknown from these strata.
SYSTEMATIC PALEONTOLOGY
Phylum Cnidaria Hatschek, 1888
Class, order, and family uncertain
Genus Sphenothallus Hall, 1847
Sphenothallus sp.
Fig. 2
Illustrated specimen: NMMNH P-37916 (part and counter-
part).
Description: Specimen P-37916 consists of a crushed tube pre-
served along the bedding plane, as part (Fig. 2A-B) and counterpart (Fig.
2C), on small slabs of gray laminar shale. The tube is brownish gray in
color and has in places a thin covering of what appears to be carbon. The
tube is fragmentary, lacking both a basal and an apertural end. As best
seen on the part slab, the tube is 33 mm long and from 7 to 9 mm in
diameter. The tube margins are mostly straight along their length and then
gently curved towards one end. A short length of tube shows longitudinal
thickening, which is best seen on the counterpart exposed along the
inwardly bent margin of the curved end. Beginning mid-length on the
opposite margin is a gently sinuous length of tube, possibly a displaced
longitudinally thickened area that lies on the tube surface is present. One
end of this section is nearly in contact with a somewhat regular longitu-
dinal fold that extends submedially along the tube. The counterpart con-
tains additional tube material towards the curved end that is not seen on
the part, although it is poorly preserved and fragmentary. Holdfasts are
present along the entire length of the tube surface and number as many as
200 individuals. Some of these holdfasts occur in isolation, whereas
others are grouped into clusters (Fig. 2D). A somewhat linear arrange-
ment is also seen. Adjoining specimens frequently touch at their borders,
although overlap between them also occurs. The holdfasts are variably
preserved, but most show a raised outer collar that is generally circular to
elliptical in shape. The average holdfast diameter approaches 1 mm,
although forms as large as 1.6 mm also occur. Some well-preserved hold-
fasts have an additional inner central collar with a central depression.
None of the holdfasts retain their own tubes, and isolated basal mem-
branes are not preserved. Conchostracan valves assigned to Lioestheria
carinacurvata Martens and Lucas, 2005, and ostracods provisionally
referred to Carbonita sp., are preserved in association on the part and
counterpart slabs.
Remarks: The specimen is assigned to the genus Sphenothallus
based on the presence of a lateral thickening along the tube margin and the
characteristic morphology of the holdfasts. The fragmentary condition
of the tube precludes making a specific assignment, so it is left in open
nomenclature. Sphenothallus tube diameters generally range from less
than 1 to 17 mm (Bodenbender et al., 1989). The Tinajas Arroyo speci-
men, with a tube diameter of 7 to 9 mm, is thus near the middle of this
range. Although tapering is a commonly reported feature of Sphenothallus
87
tubes, the variation in tube diameter of the Tinajas Arroyo specimen
likely represents taphonomic distortion rather than accurately reflecting
its morphology during life. The Tinajas Arroyo specimen has a relatively
large number of Sphenothallus holdfasts along the length of the tube. The
association of individuals or holdfasts attached to other sphenothallids
has been recorded from the Cambrian of China (Zhu et al., 2000), the
Middle Ordovician of New York (Hall, 1847; Ruedemann, 1898), the
Devonian of Brazil (Clarke, 1913), the Upper Devonian of Ohio
(Feldmann et al., 1986; Neal and Hannibal, 2000), and the Carboniferous
(Upper Mississippian) of Montana (Van Iten et al., 1992). The Tinajas
Arroyo specimen thus adds to this record.
Sphenothallus holdfasts have been described in detail from several
North American occurrences (e.g. Ruedemann, 1898; Bodenbender et al.,
1989; Van Iten et al., 1992; Neal and Hannibal, 2000). The Tinajas Ar-
royo specimens most resemble those described by Bodenbender et al.
(1989) from the (Upper Ordovician) Dillsboro Formation of Indiana by
displaying a central collar with an inner circular depression. This feature
was interpreted by Neal and Hannibal (2000) as the point of continuity
between the basal part of the tube and the holdfast, which was left
behind when the tube separated from the holdfast.
Sphenothallus appears to have been a euryoxic and eurytopic
organism that tolerated a variety of marine conditions (Malinky et al.,
2004). The Tinajas Arroyo Sphenothallus record is unique in coming
from a fresh to brackish lacustrine/lagoonal environment, but we con-
sider it an allochthonous element within the fauna. It lacks a basal hold-
fast, which prevents knowing whether it was attached to an object when
transported or detached then carried by a current to its burial site. Trans-
portation of this marine animal from its habitat into a lacustrine deposi-
tional setting may have played an important role in its preservation out
of its normal context, as microbial decomposition of marine organisms
can be inhibited in freshwater and brackish water settings (Babcock,
1998; Babcock et al., 2001). Rapid burial, possibly associated with coastal
storm surge conditions, also may have contributed to preservation of
this Sphenothallus tube in this unusual Konservat-Lagerstätte deposit
(see for comparison Babcock et al., 2000).
DISCUSSION
Sphenothallus adds to a recently augmented and important fossil
record of Paleozoic cnidarians from New Mexico. Lerner and Lucas
(2005) first reported Sphenothallus and later discussed it in more detail
within the broader context of the Tinajas Arroyo biota and
paleoenvironment (Lerner et al., 2009a). This was followed soon after-
ward by a report of porpitid hydrozoan body impressions from the
Robledo Mountains Formation (Permian: Cisuralian Series; or
Wolfcampian of historic North American usage; Ogg et al., 2008) of
south-central New Mexico (Lerner et al., 2009b). This was the first
record of porpitids from the Permian. Continued collecting from the
Robledo Mountains Formation near the newly established Prehistoric
Trackways National Monument then revealed a previously unknown
cnidarian-dominated trace and body fossil assemblage that primarily
consists of porpitid body impressions, actinarian (sea anemone) bur-
rows assigned to Bergaueria, and the limulid trace Selenichnites (Lerner
and Lucas, 2010). Several of the Bergaueria specimens, the first record
of the ichnogenus from New Mexico, are highly unusual in that they are
preserved with tentacular resting impressions. These recent discoveries,
including Sphenothallus, indicate that New Mexico contains a scientifi-
cally important late Paleozoic cnidarian fossil record that warrants fur-
ther collecting and study.
ACKNOWLEDGMENTS
The authors are indebted to Loren Babcock and Joerg Schneider
for having greatly improved the text with their thoughtful reviews. We
thank Joseph Hannibal for having initially identified the Sphenothallus
specimen. Larry Rinehart provided helpful technical assistance and dis-
cussion. The New Mexico Friends of Paleontology provided field col-
lecting assistance at the Tinajas Arroyo locality.
88
FIGURE 2. Sphenothallus sp., NMMNH P-37916. A-B, Two views (differing contrasts) of part; multiple holdfasts are seen along the length of the tube
surface. C, Counterpart; the inset photo (arrow) better shows the lateral thickening (t). D, Detail of holdfasts. Scale bars = 2 mm.
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