Biostratigraphy of Devonian microvertebrates 
425 
cristatus and asymmetricus zones) horizons at 
Broken River. Cladolepis sp. scales have been 
reported from the early Eifelian Lauch Formation 
of Germany (Vieth-Schreiner 1983) in horizons of 
patulus Zone age (Weddige 1977); the late Emsian 
Santa Lucia Formation of Spain (Mader 1986); and 
the Middle Devonian of North America (Wells 
1944; Gross 1973) in horizons dated kockelianus 
Zone or earlier (BClapper and Johnson 1980). None 
of these occurrences is as young as the Broken 
River specimens. Scales possibly belonging to the 
genus are reported from the middle Givetian-early 
Frasnian Holy Cross Mountains of Poland 
(Liszkowski and Racki 1992), and the gigas Zone 
(early Frasnian) Mostyn Vale Formation of NSW 
(Turner 1993). 
Ohiolepis Wells, 1944 
Ohiolepis sp. Wells, 1944 
Figures 4D-G 
These specimens have previously been described 
and discussed (De Pomeroy 1994). They are 
included here simply as part of the comparison of 
ranges of the Broken River fauna with occurrences 
reported elsewhere. Scales assigned to the genus 
are present in horizons of Middle and Upper varcus 
zones at Broken River (Figure 2), which is later 
than other reports. However, the concept of the 
genus appears to have become confused over the 
years (see De Pomeroy 1994) and is in need of 
revision, so the apparent late appearance of these 
scales at Broken River is at present considered of 
little significance. 
Order Euselachii Hay, 1902 
Superfamily Ctenacanthoidea Zangerl, 1981 
Family Phoebodontidae Williams, 1985 
?Phoebodontid indet. 
Figures 4H, I 
Material 
Two broken cusps (QMF 31831 and one other). 
Localities 
SD128/134.1 m and SD164/124.1 m; Papilio 
Formation. 
Stratigraphic range 
Givetian (Middle varcus and hermanni-cristatus 
zones). 
Remarks 
The Broken River fragments are similar to cusps 
of Phoebodus australiensis and P. polihis from the 
late Famennian of Thailand discussed and figured 
by Long (1990 figures 2-4). These two species have 
three- and four-cusped teeth with, respectively, 
three to four and seven to ten coarse vertical striae 
on the lingual surface (Long 1990), compared with 
six striae on the Broken River specimens. The 
Broken River specimens also resemble cusps of P. 
bifurcalus, where the labial face of some cusps bears 
strong, subparallel ridges, as found at late Frasnian 
localities in the South Urals (Ginter and Ivanov 
1992: figures 4A-F, 5D-H). Turner (1982, figure 
6A) illustrates a 3-cusped tooth assigned to 
Phoebodus cf. politus, from the Famenian Burdekin 
Star Shelf in Queensland. These cusps have 
longitudinal, raised striae, similar to the ornament 
on the Broken River specimens. 
Description 
The conical fragments are less than 1 mm long, 
with a small central pulp cavity, six straight 
longitudinal ridges on one side, and a circular 
cross-section. Under high magnification (Figure 41), 
finer oblique striations are visible between the 
ridges; the other side of the cusp is smooth. Both 
fragments are apical ends of cusps, and are not 
curved. 
Discussion 
Phoebodont teeth are known from the Middle 
Devonian to Late Carboniferous (Zangerl 1981). 
Other reported Middle Devonian phoebodont teeth 
include the two species of Phoebodus erected by 
Wells (1944) from North American material, P. 
floxueri with three to five slightly spiralling 
striations on one side of the cusp, and P. ? bryanti 
with smooth cusps and ovoid cross-section, figured 
by Gross (1973: plate 34, figure 23, plate 35, figures 
7-8); those of the middle and late Givetian of the 
Holy Cross Mountains in Poland (Liszkowski and 
Racki 1992, figures 3, 4L-N); and a Givetian tooth 
from section SD146 at Broken River, reported by 
Turner (1993). 
Chondrichthyan incertae sedis 
Figures 4J-L 
Material 
Sixty-one teeth (QMF 31832,3 and 59 others). 
Localities 
SD15/115.6 m, SD127/43 m, SD128/68.6-132 m, 
SD130/25.5-109.3 m, SD164/40.2-80.8 m, SD210/ 
75-104 m, SD216/25-106 m, and SAGW/20 m; 
Papilio Formation, Spanner and Chinaman Creek 
limestones. 
Stratigraphic range 
Givetian (kockelianus to varcus zones). 
