J. Paleont., 81(2), 2007, pp.

331–351
Copyright 䉷 2007, The Paleontological Society
0022-3360/07/0081-331$03.00

CENOZOIC TURRITELLIDAE (GASTROPODA) FROM SOUTHERN PERU

THOMAS J. DEVRIES
Burke Museum of Natural History and Culture, University of Washington, Seattle 98195;
mailing address: Box 13061, Burton, Washington 98013 USA

ABSTRACT—Cenozoic marine deposits of forearc basins in southern Peru contain a molluscan fauna that includes 15 species of turri-
telline gastropods. Twelve species were previously known; ten from northern Peru or Chile and two species solely from southern Peru.
Three new fossil species are described: Turritella riverae, Turritella cruzadoi, and Turritella salasi. Six species of turritellines with
crenulated primary spirals, a distinctive spiral ontogeny, and which are mostly endemic to Peru and Chile, are assigned to Incatella n.
gen., including I. cingulata (Sowerby, 1825), I. cingulatiformis (Möricke, 1896), I. chilensis (Sowerby, 1846), I. leptogramma (Philippi,
1887), I. hupei Nielsen, new name (⫽Turritella affinis Hupé, 1854), and I. trilirata (Philippi, 1887). Most Paleogene taxa range from
northern to southern Peru, while most Neogene taxa, including all species of Incatella, range from Peru to Chile. This biogeographic
asymmetry is attributed to a series of biotic events (e.g., extinctions, immigrations) impelled by global oceanographic changes acting
locally in a regime of coastal upwelling.

INTRODUCTION
T HE MODERN Peruvian Faunal Province contains just one spe-
cies of turritelline gastropod, Turritella cingulata Sowerby,
1825. The adjacent southern end of the Panamic Faunal Province,
which includes northernmost Peru, has five Recent species (Ala-
mo and Valdivieso, 1997; DeVries, 2003a) and was just as rich
in Turritella taxa throughout the Cenozoic (see Olsson, 1928,
1930, 1931, 1932). Studies by Sowerby (1846), Philippi (1887),
Tavera (1947, 1979), Herm (1969), and Nielsen (2003) have pro-
duced a list of about 10 Cenozoic species from Chile.
Fossils of Cenozoic Turritella Lamarck, 1799 from southern
Peru are less well known than those from northern Peru or Chile.
Only two paleontologists have contributed to the taxonomy of
turritellines from the Pisco, Sacaco, and Camaná basins (Fig. 1).
Lisson (1925) described two new species from Caravelı́, T. wood-
si Lisson, 1925 and T. portaroi Lisson, 1925, to which he as-
signed an Eocene age. Rivera (1957) described two new Eocene
turritellines from the Paracas area, T. paracasensis and T. lagun-
illasensis.
In this paper, I report several species of Cenozoic turritellines
from southern Peru formerly known only from more than 1,000
km away in Chile or northern Peru. Three new species are de-
scribed: Turritella riverae and Turritella cruzadoi, both from Up-
per Oligocene to Lower Miocene strata, and T. salasi, from Mid-
dle Miocene beds. Neotypes are designated for T. paracasensis
and T. lagunillasensis. Turritella woodsi is recognized as the se-
nior synonym for T. conquistadorana Hanna and Israelsky, 1925.
Incatella n. gen. is created for a largely endemic group of turri-
tellines present off Peru or Chile since the Late Oligocene.
GEOLOGY
The stratigraphy of the Pisco Basin has been described by Dun-
bar et al. (1990) and DeVries (1998) (Fig. 1). Within this forearc
basin, the Middle to Upper Eocene Paracas Formation overlies
crystalline bedrock. Unconformably overlying the Paracas se-
quence is the uppermost Eocene/Lower Oligocene Otuma For-
mation, which in turn is overlain by the uppermost Oligocene to
Middle Miocene Chilcatay Formation and Middle Miocene to Pli-
ocene Pisco Formation. Each Cenozoic formation is characterized
by a transgressive sandstone member representing nearshore pa-
leoenvironments and a finer-grained, tuffaceous, and diatoma-
ceous silty sandstone member representing outershelf paleoenvi-
ronments. Every formation can be found locally in contact with
crystalline basement rocks with associated deposits of bioclastic
gravel characteristic of intertidal and rocky shoreface paleoenvi-
ronments.
331
To the south of the Pisco Basin are Miocene and Pliocene ma-
rine sandstones of nearshore to shoreface origin around Sacaco
(Muizon and DeVries, 1985). A larger Cenozoic sedimentary ba-
sin surrounding the city of Camaná and a smaller area of outcrop
south of Caravelı́ have been discussed by DeVries (1998, 2001a),
Vega and Marocco (2004), and Sempere et al. (2004).
METHODS AND MATERIALS
Most Peruvian fossils in this study were found by the author.
Some were collected by J. Macharé (INGEMMET, Lima, Peru).
Comparative material was provided by the Paleontological Re-
search Institution (Ithaca, New York), the California Academy of
Sciences (San Francisco), S. Nielsen (Freie Universität Berlin,
Germany), and W. J. Zinsmeister (Purdue University, West La-
fayette, Indiana). Fossil turritellines from Chile were examined in
1993 at the Museo Nacional de Historia Natural and the Univ-
ersidad de Chile, both in Santiago, Chile. The latter site housed
the collections of J. Tavera, some of which are now stored at the
Museo Nacional de Historia Natural. Radiometric and biochron-
ostratigraphic ages are reported by Muizon and DeVries (1985),
Dunbar et al. (1990), and DeVries (1998).
Abbreviations for repositories of fossil specimens and localities
are as follows: CAS—California Academy of Sciences; F&C—
Collections of D. Frassinetti and V. Covacevich, Museo Nacional
de Historia Natural, Santiago, Chile; JM—José Macharé; MNHN-
LG—Muséum National d’Histoire Naturelle, Laboratoire de Géo-
logie, Paris; MUSM INV—Departamento de Paleontologı́a de
Vertebrados, Museo de Historia Natural, Universidad Nacional
Mayor de San Marcos, Lima, Peru; OSU—Orton Museum, Ohio
State University, Columbus, USA; PRI—Paleontological Re-
search Institution; SGO.PI.—Departamento de Paleontologı́a de
Invertebrados, Museo Nacional de Historia Natural, Santiago,
Chile; USNM—United States National Museum of Natural His-
tory, Washington, DC, USA; UWBM—Burke Museum of Natural
History and Culture, University of Washington, Seattle, USA; and
WJZ—W. J. Zinsmeister.
Museum numbers are followed by locality-sample numbers that
are listed in the Appendix. Measurements of length (L) and width
(W), in millimeters, were made with a digital caliper. Numbers
enclosed by parentheses indicate sizes for broken or deformed
specimens. Specimens were coated with ammonium chloride prior
to photography with a Nikon Coolpix 995 digital camera. Images
were prepared using Adobe Photoshop 6.0.
Morphological terminology follows that of Allmon (1996),
which is partly summative of earlier nomenclature (Merriam,
1941; Marwick, 1957a, 1957b, 1971). An ‘Allmon formula’ con-
sists of a string of hyphenated codes that refer to the trace of the
332 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
FIGURE 1—Location of the Pisco Basin in southern Peru and cities mentioned in the text. Sacaco and Camaná are centered in smaller sedimentary
basins. Stratigraphy is from DeVries (1998).
basal growth line (1 ⫽ convex, 2 ⫽ radial, 3 ⫽ concave, 4 ⫽
deeply concave, 5 ⫽ symmetrically sinuous, 6 ⫽ asymmetrically
sinuous), patterns of inflection on the lateral sinus of the growth
line (1 ⫽ anterior inflection, 2 ⫽ anterior and posterior inflections,
3 ⫽ posterior inflection, 4 ⫽ no inflections), depth of the lateral
sinus (S ⫽ shallow, M ⫽ moderate, D ⫽ deep), location of the
lateral sinus apex (A ⫽ posterior, B ⫽ medial, C ⫽ anterior), and
angle of the growth line (P ⫽ prosocline, OR ⫽ orthocline, OP
⫽ opisthocline) (Allmon, 1996). A ‘spacing formula’ consists of
a primary spiral’s designation (e.g., ‘C’) followed by a super-
scripted number. The number is the axial distance, in percent,
from the middle of anterior spiral ‘d,’ which separates the base
from the side of the whorl, to the posterior suture. A compilation
of spacing formulae for southern Peruvian turritellines is given in
Table 1.
SYSTEMATIC PALEONTOLOGY
Family TURRITELLIDAE Lovén, 1847
Genus TURRITELLA Lamarck, 1799
Type species (original designation).—Turbo terebra Linnaeus,
1758. Recent, Indo-Pacific.
TURRITELLA (TURRITELLA) INFRACARINATA Gryzbowski, 1899
Figure 2
Turritella suturalis NELSON, 1870, v. 2, p. 188 (not SOWERBY, 1846).
Turritella infracarinata GRYZBOWSKI, 1899, p. 643, pl. 20, fig. 5; SPIEK-
ER, 1922, p. 79, pl. 3, figs. 9, 10; WOODS, 1922, p. 109, pl. 18, figs.
2, 3; STEINMANN, 1929, p. 200, fig. 247; OLSSON, 1932, p. 196–197,
pl. 22, fig. 8.
Turritella infracarinata var. zorritensis SPIEKER, 1922, p. 80, pl. 3, fig.
11.
Turritella inca GRYZBOWSKI, 1899, p. 24, pl. 20, fig. 1.
Turritella inca var. trita NELSON. SPIEKER, 1922, p. 73, pl. 3, fig. 4.
Turritella rotundata GRYZBOWSKI, 1899, p. 643, pl. 20, fig. 6.
Turritella nelsoni SPIEKER, 1922, p. 74, pl. 3, figs. 5, 6.
Turritella nelsoni var. truillissatia SPIEKER, 1922, p. 78, pl. 3, fig. 8.
Diagnosis.—Shell frustate, with evenly spaced spirals of equal
strength. Primary spirals situated anteriorly. Lateral growth line
strongly prosocline.
Description.—Shell length to 110 mm, elongate, pleural angle
10⬚–11⬚. Whorls frustate. Sutures moderately impressed. Proto-
conch two to three whorls, turbinate. Early teleoconch whorls
with equally spaced spirals, situated anteriorly, spiral formula
c3B1s4a2. Later whorls with addition of secondary spirals posterior
to spiral ‘A.’ Adult whorls with seven strong spirals, formula
dC3B1S4A2R5R5R5, with additional secondary spirals often present.
Spacing formula C15B33A56 (n ⫽ 9). Lateral growth line strongly
prosocline with no points of inflection. Apex of shallow lateral
sinus situated medially or anteriorly. Basal growth line radial.
Allmon formula 2-4-S-B/C-P.
Material examined.—UWBM 97759, DV 571-5, L (38.2), W
13.0; UWBM 97760, DV 420-1, L (41.9), W 14.6; UWBM
97761, DV 579-2, L (47.8), W (19.1); UWBM 97762, DV
1653-1, L (26.5), W 8.0; UWBM 97763, DV 420-1, L (13.8), W
(4.5); MUSM INV 96, DV 420-1, L (44.8), W 12.6; MUSM INV
97, DV 1021-4, L (104.9), W (25.5). DeVries collection: DV
420-1, lot of 35; DV 419-4, lot of 69. CAS 328.01, CAS 329.02,
CAS 337.01, CAS 61706.01, northern Peru. Type locality is Zor-
ritos, Peru.
Occurrence.—Middle to lower Upper Miocene, northern and
southern Peru.
Discussion.—Turritella infracarinata is assigned to Turritella
(Turritella) Lamarck, 1799, based on its spiral ontogeny and
strongly prosocline growth-line pattern. Olsson’s (1932) obser-
vations on the variability of this species in northern Peru apply
equally to populations in southern Peru.
Specimens of Turritella infracarinata are common in Middle
to lower Upper Miocene sandstones of the Zorritos and Montera
formations of the Talara and Sechura basins, respectively (Olsson,
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 333

TABLE 1—Sculptural formulae for species of southern Peruvian turritellines. Allmon formulae are defined in text. Letters for early spiral formulae and spacing
formulae refer to spirals from anterior to posterior. Lowercase letters indicate weak spirals; upper case letters indicate strong spirals. Subscripted numbers
describe the order of appearance of spirals. Spirals share the same subscripted number when the order of appearance cannot be determined. Superscripted
numbers refer to axial distance of primary spiral, in percent, from middle of anterior spiral ‘d’ to posterior suture of whorl. ‘Sample size’ refers to data
used to calculate an average spacing formula.

Early Spiral
Turritelline Species Allmon Formula Formula Spacing Formula Sample Size (n)
infracarinata 2-4-S-B/C-P C3B1s4a2 C15B33A56 9
cingulata 1/2-1/2-M-A-OR/OP C2B1a3 C24B47A80 8
cingulatiformis 1/2-1-M-A-OR/OP c2B1a3 C25B47C76 12
chilensis 2-1-M-A-OR/OP c1B1a1 C25B46C76 1
leptogramma 2-1-M-A-OP C1B1A1 C22B48C76 3
hupei new name 2-1/2-M/D-A-OR/OP c2B1a3 C20B49A87 23
trilirata 2-1-M-A-OR/OP c2B1a3 C23B47A78 10
paracasensis 3/2-1-M-A-P/OR dC2B1a3 C35B58A78 8
cochleiformis 3-1-M-A-P/OR dC1B1 C35B61 4
cruzadoi n. sp. 3-2-D-A/B-OR/OP c2B1a3 C19B44A77 9
lagunillasensis 3/4-1-M/D-A-OR C1B2A1 C17B37A73 17
woodsi 5/4-1/2-D-B-P/OR C1B1A1r1 C19B37A68 29, 9, 15 for C, B, A
conquistadorana1 5/4-1/2-D-B-P/OR ? C17B42A74 6
‘trongolensis’ 5/4-4-D-B-P/OR C1B1s1A1r1 C18B38A74 4
salasi n. sp. ?-4-D-A-P C1B1A1 C13B45A69 1
chira (s. Peru) 2-1-M-A-P/OR c2B1a3 C28B57A71 6
chira (n. Peru) 2-1-M-A-OR — C29B58A70 1
riverae n. sp. 2-1/2-S-B-P/OR C1B1a2 C29B57A73 10
1
PRI and CAS samples from northern Peru (⫽Turritella woodsi).

1932). Specimens from the Pisco Basin include those from shore-
face and inner shelf sandstones of the lower part (Middle Mio-
cene) of the Pisco Formation (DeVries, 1998). Specimens were
also found farther south in a lower Upper Miocene shoreface bio-
clastic sandstone at Alto Grande (DV 571-5), several meters
above an ash bed dated radiometrically at about 9 Ma (Muizon
and DeVries, 1985).
Genus INCATELLA new genus
Type species.—Incatella cingulata (Sowerby, 1825). Recent,
Ecuador to southern Chile.
Other species.—I. cingulatiformis (Möricke, 1896), I. chilensis
(Sowerby, 1846), I. leptogramma (Philippi, 1887), I. hupei Niel-
sen, new name (⫽Turritella affinis Hupé, 1854), I. trilirata (Phi-
lippi, 1887).
Diagnosis.—Spiral formula c2B1a3 on early whorls. Adult pri-
mary spirals thick, ropelike, crenulated or beaded. Posterior and
anterior inflections on latter only; apex of lateral sinus in growth
line posteriorly situated. Basal growth line radial to slightly con-
vex.
Description.—Shell small- to medium-sized. Pleural angle 12⬚–
24⬚. Whorl profile flat-sided, subquadrate or convex. Primary spi-
rals thick, ropelike, strongly crenulated or beaded by intersections
with growth lines. Earliest spiral formula c2B1a3, becoming
dC2B1A3 on succeeding whorls. Spacing of spirals variable; in-
terspace C/B narrower than B/A in Peruvian populations, formula
about C23B47A78; interspace C/B wider than B/A in some Chilean
populations. Adult whorls with variable addition of secondary spi-
rals and tertiary threads. Lateral growth lines orthocline to op-
isthocline, inflected anteriorly and sometimes posteriorly. Lateral
sinus moderately deep, apex situated posteriorly, just posterior to
spiral ‘B.’ Base convex, with five to seven spirals anterior to
spiral ‘d.’ Basal growth line radial to slightly convex adapertur-
ally. Allmon formula 1/2-1/2-M/D-A-OR/OP.
Etymology.—Prefix ‘Inca,’ a ruler of the pre-Columbian Inca
Empire.
Occurrence.—Uppermost Oligocene (Peru), Miocene (Peru,
Chile, Argentina), Pliocene to Recent, Peru and Chile.
Discussion.—Incatella includes six species from Peru: I. cin-
gulata, Pleistocene to Recent; I. cingulatiformis, Late Miocene to
Pliocene; I. chilensis, Late Miocene to Pliocene; I. leptogramma,
Early to Late Miocene; I. hupei, Early to Middle Miocene; and I.
trilirata, latest Oligocene to Early Miocene. All six species are
found in Chile (Philippi, 1887; Tavera, 1979). These few species
and perhaps others from Argentina (e.g., Turritella patagonica
Sowerby, 1846; see Ortmann, 1902; Ihering, 1907; del Rı́o, 2004)
are distinguished from other turritellines in the Americas and else-
where by a combination of their early spiral sculptural formula;
strong ropelike primary spiral cords, usually three in number;
spacing formula; and Allmon formula.
The combination of spiral ontogeny, growth-line form, and
whorl profile exhibited by species assigned to Incatella is shared
in part by Colpospira (Platycolpus) Donald, 1900, a genus of very
small- to medium-sized modern turritellines from Australia (Don-
ald, 1900; Allmon, 1996), but the genera differ consistently in the
position of the apex of the lateral sinus and to a lesser extent in
the inclination of the growth lines.
There are no Eocene species in northern Peru figured by Olsson
(1928, 1930, 1931) that are plausible ancestors to the Incatella
clade of turritellines, nor have any been identified in Eocene de-
posits of Chile. The existence of crenulated tricordate turritellines
in Oligocene deposits of Argentina (Ortmann, 1902; Ihering,
1907; del Rı́o, 2004) suggests a South Atlantic origin for Inca-
tella.
INCATELLA CINGULATA (Sowerby, 1825)
Figure 3.1–3.3
Turritella cingulata SOWERBY, 1825, p. 56, appendix p. xiii; REEVE,
1849, v. 5, pl. 6, fig. 23; MÖRICKE, 1896, p. 556, pl. 11, fig. 5; DALL,
1909, p. 231; CARCELLES AND WILLIAMSON, 1951, p. 273; HERM,
1969, p. 132–133, pl. 14, figs. 14, 15, not figs. 12, 13; KEEN, 1971, p.
392, fig. 436; MARINCOVICH, 1973, p. 30, fig. 61; DEVRIES, 1986, p.
521, pl. 28, fig. 7; ALAMO AND VALDIVIESO, 1997, p. 20, fig. 50;
FORCELLI, 2000, p. 72, fig. 150.
Diagnosis.—Shell stout, pleural angle 18⬚–24⬚. Sculpture of
three equally strong primary spirals. Secondary spirals variably
developed; tertiary spirals poorly developed or absent.
Description.—Shell length 40–50 mm, stout, pleural angle 18⬚–
24⬚. Whorls usually flat-sided, sometimes weakly subquadrate or
slightly convex. Sutures shallow, slightly impressed to appressed.
Protoconch unknown. Early whorls with three equally spaced spi-
rals, spiral formula C2B1a3; interspaces as wide as spirals. Later
334 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
FIGURE 2—Turritella infracarinata Gryzbowski, 1899. All from the Mid-
dle Miocene portion of the Pisco Formation. 1, MUSM INV 96, ⫻1.9;
2, UWBM 97762, ⫻2.8; 3, UWBM 97761, ⫻1.8; 4, UWBM 97760,
⫻1.8; 5, UWBM 97760, basal view, ⫻2.0; 6, UWBM 97763, ⫻5.3.
whorls with primary spirals equally strong (formula C2B1A3),
spacing formula C24B47A80 (n ⫽ 8). Primary spirals strongly cren-
ulated or beaded. Secondary spirals inserted on later whorls, spiral
formulae include C2B1s4A3r5 and C2t4B1s4A3. Tertiary threads ab-
sent or weakly developed. Lateral growth line orthocline to op-
isthocline with posterior and anterior inflections or latter only;
apex of lateral sinus situated posteriorly. Convex base with five
to seven spirals; basal growth line radial to convex. Spirals brown;
background cream-colored. Allmon formula 1/2-1/2-M-A-OR/OP.
Material examined.—OSU 37542, DV 333-21, L 34.5, W 16.4;
UWBM 97710, DV 769-1, L (40.0), W 15.8; UWBM 97711,
82JM 017, L 39.3, W 14.8; MUSM INV 76, DV 769-1, L 29.4,
W 12.6. DeVries collection, DV 769-1, lot of 8; DV 381-5, lot
of 9; DV 465a-1, lot of 1; WJZ 346, Chile, lot of 2; WJZ 343,
Chile, lot of 5. Type locality is Valparaiso, Chile.
Occurrence.—Lower Pleistocene, northern Peru. Middle Pleis-
tocene, northern Peru to central Chile. Upper Pleistocene to Re-
cent, Ecuador to Chiloé, southern Chile.
Discussion.—Specimens of Incatella cingulata from Peru are
distinguished by a wide pleural spiral angle, shallow sutures, and
whorls that are flat-sided. These features are always present in
specimens from Lower and Middle Pleistocene deposits of north-
ern Peru (DeVries, 1986), Middle and Upper Pleistocene marine
terraces of southern Peru, and modern Peruvian beaches, and are
usually present in specimens from Chile. Some modern Chilean
specimens, however, differ by having narrow pleural angles (10⬚–
15⬚) and deeper sutures (see Herm, 1969, pl. 14, figs. 14, 15).
Specimens of Incatella cingulatiformis are longer and propor-
tionally narrower than those of I. cingulata and have deeper su-
tures with steeply inclined sutural platforms.
INCATELLA CINGULATIFORMIS (Möricke, 1896)
Figure 3.4–3.7, 3.20
Turritella cingulatiformis MÖRICKE, 1896, p. 556, pl. 11, fig. 4; HERM
AND PASKOFF, 1967, p. 23; HERM, 1969, p. 132, pl. 14, figs. 10, 11.
Turritella cingulata SOWERBY. HERM, 1969, pl. 14, figs. 12, 13.
Diagnosis.—Shell length to 70 mm. Whorls strongly subquad-
rate; sutures deep. Secondary spirals and tertiary threads often
well developed.
Description.—Shell length to 70 mm, pleural angle 15⬚–17⬚.
Whorls strongly subquadrate. Sutures deeply impressed; sutural
platform steeply inclined. Protoconch unknown. Whorls with
three primary spirals, strongly crenulated or beaded. Spiral for-
mula on early whorls c2B1a3, followed by C2B1A3 and, with ad-
dition of secondary spirals on adult whorls, C2t5B1s4A3 or
C2t5B1s4s5A3. Tertiary threads often present. Spacing formula
C25B47C76 (n ⫽ 12). Lateral growth line orthocline to opisthocline
with anterior and posterior inflections or latter only. Lateral sinus
with apex situated posteriorly. Convex base with five to seven
spirals; basal growth line radial. Spirals brown. Allmon formula
1/2-1-M-A-OR/OP.
Material examined.—UWBM 97706, DV 381-1, L 30.0, W
10.5; UWBM 97707, DV 571-1, L (42.4), W 17.4; UWBM
97708, DV 523-2, L (44.0), W (16.8); UWBM 97709, DV
1453-1, L (44.5), W 17.0; MUSM INV 77, DV 523-2, L (43.6),
W 12.0; MUSM INV 78, DV 381-1, L (32.4), W 9.9; MUSM
INV 79, DV 1235-4, L (22.9), W 13.0; MUSM INV 80, DV
1235-4, L (32.1), W (16.1); SGO.PI.1083, Quebrada Blanca, Cal-
dera, L (41.4); SGO.PI.1084, Quebrada Blanca, Caldera. DeVries
collection: DV 381-1, lot of 1; DV 1235-4, lot of 1; DV 523-2,
lot of 10; WJZ 343, Chile, lot of 6. Type locality is Coquimbo,
Chile.
Occurrence.—Upper Miocene, southern Peru. Pliocene to Mid-
dle Pleistocene, southern Peru to central Chile.
Discussion.—Incatella cingulatiformis was first described from
Pliocene strata in Chile (Möricke, 1896), where its elongate, sub-
quadrate form persisted into the Pleistocene (Herm, 1969). In
northern Peru, the species is unknown, and in southern Peru, spec-
imens are found in beds no younger than the Middle Pleistocene,
at which time they were replaced by those of the stouter I. cin-
gulata. Teusch et al. (2002) cite specimens from upper Pleistocene
terraces in southern Chile, but did not describe criteria for distin-
guishing specimens of I. cingulatiformis from I. cingulata. The
species is longer-lived than indicated by its record in Chile, with
large specimens found in lower Upper Miocene beds in both the
Pisco and Sacaco basins of southern Peru.
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU
INCATELLA CHILENSIS (SOWERBY, 1846)
Figure 3.8
Turritella chilensis SOWERBY, 1846, p. 257, pl. 4, fig. 51; D’ORBIGNY,
1852, p. 33; PHILIPPI, 1887, p. 72, pl. 9, fig. 6; TAVERA ET AL., 1985,
pl. 2, figs. 8, 9; FRASSINETTI, 2000, p. 134, pl. 1, figs. 1–3.
Diagnosis.—Whorls keeled, medial primary spiral ‘B’ stron-
gest. Secondary spirals absent; tertiary threads numerous.
Description.—Shell length to 60 mm, elongate, pleural angle
12⬚–14⬚. Whorls keeled. Sutures weakly to moderately impressed.
Protoconch unknown. Earliest preserved teleoconch whorls with
spiral formula c1B1a1; adult whorl formula d2c1B1a1. Medial pri-
mary spiral thick, sometimes forming keel; all primary spirals
strongly beaded. Secondary spirals absent; tertiary threads nu-
merous. Spacing formula C25B46C76 (n ⫽ 1). Lateral growth line
orthocline to opisthocline, weakly inflected anteriorly. Lateral si-
nus moderately deep, apex situated posteriorly. Base with three
to five primary spirals; basal growth line radial. Allmon formula
2-1-M-A-OR/OP.
Material examined.—The type locality of Incatella chilensis is
Isla de la Mocha, south-central Chile. P. Jeffery (Natural History
Museum, London, personal commun., 1995) noted that labels
with both Isla Guafo and Isla de la Mocha are attached to a block
with syntypes. The block also bears a number associated with a
larger block of sedimentary rock that comes from Mocha, leading
Jeffery to conclude that the syntypes are from Isla de la Mocha.
UWBM 97703, DV 599-2, L 48.2, L 13.5; UWBM 97704, DV
599-2, L (36.6), W 13.5; UWBM 97705, DV 599-2, L (31.1), W
10.5; F&C 280884.1-1, Isla Guafo, L (45.7), W 12.2; F&C
280884.1-2, Guafo, L (26.6), W 8.4; F&C 259, Guafo, L (38.2),
W (13.4); F&C 170693-8-1, Isla de la Mocha, L (32.0), W (11.8);
F&C 170693-8-2, Mocha, L (25.5), W (8.1); SGO.PI. 610.
Occurrence.—Upper Miocene to Lower Pliocene, southern Peru
to southern Chile.
Discussion.—Specimens of Incatella chilensis differ from those
of I. cingulata, I. cingulatiformis, and I. leptogramma most no-
tably in the persistent prominence of primary spiral ‘B’ and less
consistently in the absence of secondary spirals.
Sowerby was of two minds whether to refer specimens with a
pronounced medial primary spiral to Turritella cingulata or a new
species, but was swayed by d’Orbigny to the latter course (Sow-
erby, 1846). Oddly, T. chilensis is not mentioned by Hupé (1854)
or Möricke (1896). References to Paleogene occurrences of I.
chilensis in Argentina (e.g., Stilwell, 2003) are mistaken (Stilwell,
personal commun., 2005).
Mollusks on Isla de la Mocha, south-central Chile, found by
Frassinetti and Covacevich (personal commun., 1993) with In-
catella chilensis suggest a Late Miocene age, whereas species
associated with I. chilensis from Isla Guafo, southern Chile, sug-
gest a Pliocene age (Frassinetti, 1997). The few poorly preserved
specimens of I. chilensis from southern Peru come from the Mon-
temar area of the Sacaco Basin (DV 599-2), where other mollusks
and radiometrically dated strata suggest an age of Late Miocene
to Early Pliocene (Muizon and DeVries, 1985).
INCATELLA LEPTOGRAMMA (Philippi, 1887)
Figure 3.9–3.11
Turritella leptogramma PHILIPPI, 1887, p. 70, pl. 9, fig. 30.
‘Turritella leptogramma’ PHILIPPI, 1887. NIELSEN, 2003, p. 47, pl. 7, figs.
1, 2.
Turritella breantiana D’ORBIGNY. PHILIPPI, 1887, p. 73, pl. 9, fig. 1b.
Not Turritella breantiana D’ORBIGNY, 1847, part 4 (Géologie), pl. 5, figs.
37, 38 (replacement name for Turritella monilifera FORBES, 1846 [not
DESHAYES, 1825]); D’ORBIGNY, 1850, p. 218. Late Cretaceous, India.
Turritella breantiana var. nana TAVERA, 1979, p. 93, pl. 18, fig. 61a,
61b.
335
Diagnosis.—Whorls flat-sided to slightly convex, suture weakly
impressed to appressed. Sculpture of three weak primary spirals
and numerous secondary spirals and tertiary threads.
Description.—Length to 50 mm, elongate, pleural angle 9⬚–12⬚.
Whorls flat-sided to slightly convex, sutures weakly impressed
to appressed. Protoconch and earliest teleoconch whorls
unknown. Three equally strong primary spirals on earlier whorls,
spiral formula C1B1A1. Additional secondary spirals on adult
whorls, typical formulae include C1t4t4B1s3s2A1 (Chile) or
duC1t4t4B1s2s2A1r3r3r3r3 (Peru). Additional tertiary threads some-
times present. Spacing formula C22B48C76 (n ⫽ 3). Lateral growth
line opisthocline, inflected anteriorly, apex of lateral sinus situated
posteriorly. Base with several primary spirals; basal growth line
radial. Allmon formula 2-1-M-A-OP.
Material examined.—UWBM 97699, DV 523-2, L 40.0, W
12.0; UWBM 97700, DV 532a-3, L (17), W (10); UWBM 97701,
DV 523-4, L (24.7), W 10.8; UWBM 97702, Lo Abarca, L (8.5),
W 3.8; MUSM INV 73, DV 1653-1, L (19.4), W 9.9; F&C
150693.1, F&C 150693.2, F&C 150693.3, F&C 150693.5, F&C
170693.1, F&C 170693.2, F&C 170693.5, F&C 170693.8, Isla de
la Mocha; F&C 271080-1, Cerro Los Pololos; SGO.PI. 600, Al-
garrobo?; SGO.PI. 618, Navidad. Tavera collection: Punta Perro,
lot of 35; Algarrobo, lot of 1 (T. leptogramma).
Occurrence.—Lower Miocene, central Chile, Argentina. Middle
to Upper Miocene, southern Peru to central Chile.
Discussion.—Turritellines from Chile with poorly differentiated,
ropelike, primary and secondary spirals have long been classified
as ‘Turritella breantiana d’Orbigny’ (Philippi, 1887; Tavera,
1979). This name, however, was intended for a Cretaceous species
from India, Turritella monilifera Forbes, 1846, whose name was
preoccupied by T. monilifera Deshayes, 1824, an Eocene species
from the Paris Basin (d’Orbigny, 1850; Stoliczka, 1868). (The
Recent Chinese turritelline, T. monilifera Adams and Reeve,
1850, has never figured in the synonymy of the Chilean ‘T. brean-
tiana.’)
Confusion over the provenance of ‘Turritella breantiana’ spec-
imens stems from d’Orbigny’s (1847) inclusion of a figure of
‘Turritella breantiana,’ without text, on plate 5, ‘Coquilles fos-
siles des terrains crétacés,’ a plate that elsewhere (Grange, 1848,
p. 217–218) is entitled ‘Coquilles fossiles des terrains crétacés du
Chili.’ Not all of the figured specimens on plate 5, however, are
from Chile. Grange’s (1848) misleading plate caption and a su-
perficial similarity between the strongly tricordate Indian and Chi-
lean turritellines probably led Philippi (1887) to accept Grange’s
(1848) geographic attribution of ‘T. breantiana’ to Chile. Philippi
(1887) did doubt d’Orbigny’s (1847) assignment of the species to
the Cretaceous, however, concluding instead that ‘T. breantiana’
was a Tertiary species.
The Indian taxon differs in significant respects from the Chilean
species, having a smooth, strongly convex base with sinuous basal
growth lines, a weakly developed ‘d’ spiral, a tendency towards
obsolescence of spiral sculpture on the ultimate and even penul-
timate whorl, and a very thin outer lip (Stoliczka, 1868).
The next available name for ‘Turritella breantiana’ of Chile is
T. leptogramma Philippi, 1887. One syntype, SGO.PI 610, said
by Philippi (1887) to be from Navidad, is best referred to Inca-
tella chilensis (DeVries, 1993, personal data; Nielsen, 2003). An-
other syntype, SGO.PI 600, purportedly from Algarrobo, is com-
parable to Chilean material assigned by Chilean paleontologists
to ‘T. breantiana’ (DeVries, 1993, personal data; Nielsen, 2003).
Nielsen (2003) did note that the turritellines and rock matrix of
SGO.PI 600 were unlike samples he had collected from Algar-
robo, leaving the type locality and Philippi’s (1887) Cretaceous
to Tertiary range of I. leptogramma in doubt.
The primary spirals of Incatella leptogramma are weaker than
in any other Incatella species. Specimens of I. leptogramma have
336 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 337

weakly impressed sutures like those of I. cingulata, but are nar-
rower and have a greater number of secondary and tertiary spirals.
Specimens of Incatella leptogramma are found in Middle and
Upper Miocene strata in southern Peru. Specimens from Chile
intermediate between I. chilensis and I. leptogramma are found
on Isla de la Mocha in Upper Miocene deposits with specimens
of I. chilensis. A Late Miocene age is indicated for a mainland
occurrence of I. leptogramma at Lo Abarca, Chile (Fig. 3.11;
Covacevich and Frassinetti, 1990; DeVries, 2003b).
Small specimens of Incatella leptogramma are reported from
the Navidad Formation of central Chile (Philippi, 1887; Tavera,
1979, as T. breantiana var. nana). Tavera (1979) remarked upon
the size difference between specimens in the Navidad Formation
and on Isla de la Mocha. Current difficulties assessing the age of
transported molluscan material in the Navidad Formation (Nielsen
et al., 2003) make the phylogenetic significance of small Navidad
specimens of I. leptogramma uncertain.
Further comparative study is needed of Argentinian material to
determine the relationships of South Atlantic species assigned to
specimens of ‘Turritella breantiana’ figured by Ortmann (1902)
and Ihering (1907).
INCATELLA HUPEI Nielsen, new name
Figure 3.12–3.17
Turritella affinis HUPÉ, 1854, v. 8, p. 155; Conq., pl. 2, fig. 7, 7a; PHI-
LIPPI, 1887, p. 72, pl. 9, fig. 31; MÖRICKE, 1896, p. 555, pl. 11, fig. 3;
PHILIPPI, 1897, pl. 10, fig. 2.
Not Turritella affinis MÜLLER, 1849, p. 240 (nomen nudum); 1851, p.
31, pl. 3, fig. 11.
Not Turritella affinis ARCHIAC AND HAIME, 1853, p. 295, pl. 27, figs.
16–19.
Diagnosis.—Whorls strongly subquadrate, suture canaliculate.
Spiral sculpture of three to five strong beaded primary spirals.
Description.—Shell length to 40 mm, pleural angle 15⬚–16⬚.
Whorls strongly subquadrate. Sutures deeply impressed, canalic-
ulate, sutural platform horizontal to concave. Protoconch turbi-
nate. Early teleoconch with equally spaced spirals, spiral formula
c2B1a3. Adult spiral ‘A’ very strong, beaded, bordering suture;
spiral ‘C’ near base of whorl. Interspaces C/B and B/A each with
one secondary spiral, sometimes equal in size to primary spirals;
sometimes with tertiary threads; formula u7C2T5B1S4s6A3. Spacing
formula C20B49A87 (n ⫽ 23). Growth line orthocline; lateral sinus
inflected anteriorly, apex situated posteriorly. Base with five to
seven primary spirals and intervening secondary spirals; basal
growth line radial. Allmon formula 2-1/2-M/D-A-OR/OP.
Etymology.—Named after H. Hupé, who first described this spe-
cies.
Types.—According to Nielsen (personal commun., 2005), there
are 10 syntypes present in the MNHN-LG under number Gg2002/
49. Hupé (1854) gave Chiloé and Cahuil as localities of his ma-
terial, but according to Nielsen (personal commun., 2005), there
are no Cenozoic sediments near Cahuil and this locality name
usually refers to the Navidad Formation farther to the north. Since
all syntypes present in the collection of the MNHN-LG come
from Chiloé, the type locality is here restricted to the cliffs south
of Cucao, Chiloé Island, southern Chile. Both the Lacui Forma-
tion (Chiloé Island) and Navidad Formation (Navidad area) have
been dated as Late Miocene (Nielsen et al., 2003), but may range
into the Early Pliocene (Finger et al., 2003). However, as has been
noted by Nielsen et al. (2003), many of the mollusks may be
reworked from older units and the age of individual species re-
mains unresolved.
Material examined.—UWBM 97684, DV 937-1, L (15.1), W
(7.2); UWBM 97685, DV 542-1, L (23.0), W (9.8); UWBM
97686, DV 1428-1, mold, L (39.4), W 12.0; UWBM 97687, DV
1428-1, L 30.5, W 8.6; UWBM 97688, DV 1428-1, L (22.1), W
9.6; UWBM 97689, DV 590-1, L (21.3), W 10.7; UWBM 97690,
DV 590-1, L (35.7), W 10.2; UWBM 97691, DV 1647-1, L
(24.0), W 9.7; UWBM 97692, DV 590-1, L (31.8), W12.1;
MUSM INV 68, DV 590-1, L (30.1), W (11.0); MUSM INV 69,
DV 590-1, lot of 6; MUSM INV 70, DV 590-1, L (24.0), W 11.1;
MUSM INV 71, DV 590-1, L (22.2), W 9.3; MUSM INV 72,
DV 1647-1, L (22.56), W 9.8. DeVries collection: DV 590-1, lot
of 37; DV 1428-1, lot of 19; DV 1310-1, lot of 4 and block with
multiple molds. Type locality is Isla Chiloé, southern Chile.
Occurrence.—Lower to lower Middle Miocene, southern Peru
to southern Chile.
Discussion.—The coarsely beaded sculpture of Incatella hupei
resembles that of I. cingulata, a fact noted by Hupé (1854) and
Herm (1969), and I. trilirata. Individuals of I. hupei from south-
ern Peru vary widely in their spiral patterns: some exhibit five
strong beaded spirals (Fig. 3.13), while others have three, as seen
on specimens of I. cingulata and I. trilirata. Some specimens of
I. hupei have secondary spirals and tertiary threads characteristic
of I. leptogramma (Fig. 3.12). The primary spirals on specimens
of I. hupei, however, are not as broad as those of I. cingulata,
and tertiary threads are fewer than seen on specimens of I. lep-
togramma. Most specimens of I. trilirata lack the subquadrate
whorl profile and canaliculate sutures of I. hupei.
Peruvian specimens of Incatella hupei were found in coarse-
grained sandstone west of the Rı́o Ica (DV 590-1) in a channel
of coarse-grained sandstone cutting siltstones with a diatom flora
assigned to the C. elegans Zone (19.9–18.7 Ma) (H. Schrader,
University of Bergen, Norway, personal commun., 1989). A spec-
imen of I. hupei was also found beneath an angular unconformity
at Cerro Yesera de Amara (DV 542-1), where the oldest deposits
above the unconformity are estimated to be 16–13 Ma, based on
diatoms (Rønning, 1990). A specimen from the Filudo Depression
(DV 937-1), recovered from a horizon intercalated with sand-
stones bearing specimens of Turritella infracarinata, has an age
of about 14 Ma (DeVries, 1998).
INCATELLA TRILIRATA (Philippi, 1887)
Figure 3.18, 3.19
Turritella trilirata PHILIPPI, 1887, p. 71, pl. 9, fig. 8.
Turritella parvula PHILIPPI, 1887, p. 73, pl. 57, fig. 4.

←
FIGURE 3—1–3, Incatella cingulata (Sowerby, 1825). 1, OSU 37542, Early Pleistocene, Mancora Tablazo, ⫻1.9; 2, UWBM 97710, Middle Pleis-
tocene, ⫻1.5; 3, UWBM 97711, Middle Pleistocene, ⫻1.9. 4–7, 20, Incatella cingulatiformis (Möricke, 1896). All from the Pisco Formation. 4,
UWBM 97708, Upper Miocene, ⫻1.5; 5, MUSM INV 79, Upper Miocene, basal view, ⫻2.6; 6, MUSM INV 78, Pliocene, ⫻2.1; 7, MUSM INV
79, ⫻2.3; 20, UWBM 97706, Pliocene, ⫻2.5. 8, Incatella chilensis (Sowerby, 1846). UWBM 97704, Upper Miocene, Pisco Formation, ⫻2.1. 9–
11, Incatella leptogramma (Philippi, 1887). 9, MUSM INV 73, Middle Miocene, Pisco Formation, ⫻3.0; 10, UWBM 97699, Upper Miocene,
Pisco Formation, ⫻2.0; 11, UWBM 97702, Upper Miocene, Lo Abarca, Chile, ⫻5.0. 12–17, Incatella hupei Nielsen, new name. 12, MUSM INV
70, Lower Miocene, Chilcatay Formation, ⫻2.1; 13, UWBM 97692, Lower Miocene, Chilcatay Formation, ⫻2.2; 14, UWBM 97688, Middle
Miocene, Pisco Formation, basal view, ⫻3.1; 15, UWBM 97687, Middle Miocene, Pisco Formation, spire only, ⫻4.9; 16, UWBM 97691, Middle
Miocene, Pisco Formation, ⫻2.7; 17, UWBM 97687, entire shell, ⫻2.8. 18, 19, Incatella trilirata (Philippi, 1887). 18, UWBM 97758, Lower
Miocene, Chilcatay Formation, ⫻2.9; 19, UWBM 97696, Upper Oligocene, Camaná Formation, ⫻2.7.
338 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
Diagnosis.—Shell elongate. Three primary spirals prominent;
secondary spirals weak.
Description.—Length to 50 mm, pleural angle 12⬚–14⬚. Whorls
subquadrate to straight-sided. Sutures weakly to deeply im-
pressed, rarely canaliculate, sometimes with steeply inclined su-
tural platforms. Protoconch turbinate. Early whorls with formula
c2B1a3; later whorls with formula C2B1A3, followed by adult for-
mula dC2B1s4A3 and, rarely, dC2t5B1s4A3. Primary spirals promi-
nent, subequal in strength; crenulated or beaded. Secondary spi-
rals weak, tertiary threads variably developed. Spacing formula
C23B47A78 (n ⫽ 10). Lateral sinus of growth line inflected ante-
riorly and posteriorly, apex situated posteriorly between spirals
‘s’ and ‘B.’ Base with five to seven spirals; basal growth line
radial. Allmon formula 2-1-M-A-OR/OP.
Material examined.—UWBM 97758, DV 1320-1, L (21.1), W
7.1; UWBM 97693, DV 1320-1, mold, (34.0), W (13.7); UWBM
97694, DV 1320-1, L (14.5), W 9.9; UWBM 97695, DV
1320-1, L (13.8), W 9.3; UWBM 97696, DV 1258-1, L 25.7, W
(8.0); UWBM 97697, DV 1258-1, L (11.5), W (5.6); UWBM
97698, DV 1648-1, L (11.3), W 6.0; MUSM INV 103, DV
1320-1, L (15.0), W 8.5; MUSM INV 104, DV 1320-1, L (11.3),
W 4.8; MUSM INV 105, DV 1648-1, L (15.5), W 6.7. DeVries
collection: DV 1320-1, lot of 9 and molds; DV 1316-1, numerous
molds; DV 1648-1, lot of 7. Type locality is Navidad, Chile.
Occurrence.—Uppermost Oligocene, southern Peru. Lower
Miocene, southern Peru to central Chile.
Discussion.—Specimens of Incatella trilirata from the Navidad
Formation of central Chile provided by S. Nielsen have growth
lines that vary from uninflected to inflected anteriorly and have a
narrower B/A interspace, rather than C/B. Specimens of I. trilirata
from Peru differ from those of I. hupei by being narrower and
consistently having only three strongly developed spirals, rather
than five or three strong primary spirals and two strong secondary
spirals (‘s,’ ‘t’). More variability in the differentiation of spirals
is seen in Chilean specimens provided by Nielsen. Specimens of
I. cingulatiformis and I. leptogramma have wider pleural angles
and a greater number of secondary spirals and tertiary threads.
Specimens of I. trilirata with secondary spirals, however, are very
difficult to distinguish from juvenile specimens of I. cingulatifor-
mis (Fig. 3.20), except in the aggregate of a population/assem-
blage.
Syntypes of Turritella parvula Philippi, 1887, examined in
1993 by the author, appear to be juvenile specimens of I. trilirata.
Incatella trilirata includes the oldest known examples of In-
catella n. gen. from Peru, found in a marine intercalation (DV
1258-1) within a continental volcanoclastic sequence near Car-
avelı́ (DeVries, 2003a), a few meters below beds dated at about
25 Ma (Noble et al., 1985). Other examples of I. trilirata were
found in beds of the Lower Miocene Chilcatay Formation in the
Pisco Basin (DeVries, 1998). The relationship of Chilean and Pe-
ruvian material with Argentinian specimens of Turritella iheringi
Cossmann, 1898 is unknown.
Genus CRISTISPIRA Allison, 1965
Type species (by original designation).—Cristispira pugetensis
Allison, 1965. Eocene, Washington State, USA.
Discussion.—The assignment of Turritella paracasensis Rivera,
1957 and T. cochleiformis Gabb, 1869 to Cristispira is based on
a shared ontogeny of spiral sculpture and growth-line pattern.
Differences in secondary spiral sculpture between northern and
southern Peruvian populations of C. paracasensis imply a broader
definition is needed for Cristispira, as do differences in the
strength and number of primary spirals between Peruvian taxa
and the western North American C. pugetensis.
CRISTISPIRA PARACASENSIS (Rivera, 1957)
Figure 4.1–4.9
Turritella paracasensis RIVERA, 1957, p. 174, pl. 1, fig. 2; pl. 2, fig. 11.
Turritella chira OLSSON, 1928. OLSSON, 1930, p. 67, pl. 12, figs. 1–4.
Not Turritella chira OLSSON, 1928, p. 66, pl. 14, fig. 5.
Diagnosis.—Shell small- to medium-sized, hypercampanulate,
with three keeled spirals clustered near periphery and few sec-
ondary spirals.
Description.—Shell length to 35 mm, whorls hypercampanu-
late, pleural angle 12⬚–13⬚. Sutural area deeply concave. Proto-
conch unknown. Early teleoconch whorls with spiral formula
dC2B1a3. Later whorls with appearance of secondary spirals, for-
mula dC2B1A3r4 followed by du5C2t5B1s5A3r4. Primary spirals
strongly keeled. Spacing formula C35B58A78 (n ⫽ 8), but quite
variable. Interspaces with tertiary threads. Lateral growth line pro-
socline to orthocline, inflected anteriorly. Lateral sinus moderately
deep, apex situated posteriorly between spirals ‘B’ and ‘A.’ Inner
lip with broad parietal callus. Base flattened, with five rectangular
primary spirals; bound by sharp spiral ‘d.’ Basal growth line ra-
dial to slightly concave. Allmon formula 3/2-1-M-A-P/OR.
Types.—R. Rivera (personal commun., 1986) confirmed that the
type specimen of Turritella paracasensis has been lost. The type
locality was the Paracas Peninsula. The neotypic locality is DV
634-1, 0.3 km north of La Mina, also on the Paracas Peninsula.
Paracas Formation, Upper Eocene. Specimen UWBM 97719 is
designated the neotype of Cristispira paracasensis (⫽Turritella
paracasensis). UWBM 97719, neotype, L 24.1, W 5.9.
Other material examined.—The following are from the neotypic
locality, DV 634-1: UWBM 97720, L (14.1), W 9.5; UWBM
97721, L (12.3), W (6.7); MUSM INV 75, L (10.7), W 7.2;
MUSM INV 98, L (7.5), W (21.1); MUSM INV 99, L (14.8), W
7.3. Additional specimens: UWBM 97746, DV 501-2, L (11.6),
W 8.1; DeVries collection: DV 1174-1, lot of 1; from northern
Peru: PRI 27284, L (34.5), W 9.9; PRI 2485, L (28.7), W 10.3;
PRI 27786, L (27.3), W 8.7; PRI 24287, L (22.1), W 6.2.
Occurrence.—Middle Eocene, northern to southern Peru.
Discussion.—Specimens of Cristispira paracasensis superfi-
cially resemble the holotype of ‘Turritella chiraensis’ [sic]
[⫽‘‘Turritella chira’’ Olsson (1928) of Rivera (1957)], but they
lack the biangulate frustate profile of the Saman holotype. They
do closely resemble specimens of T. chira figured by Olsson
(1930) from the Talara Formation at Yasila, which are herein re-
assigned to C. paracasensis. The Yasila specimens do have more
strongly developed secondary spirals near the posterior suture
(formula duC2B1sA3r4r4) than do southern Peruvian specimens.
CRISTISPIRA COCHLEIFORMIS (Gabb, 1869)
Figure 4.10–4.12
Turritella cochleiformis GABB, 1869, p. 29; 1878, p. 264, pl. 35, fig. 7,
7a; HANNA AND ISRAELSKY, 1925, p. 41, pl. 7, figs. 6, 7.
Diagnosis.—Shell small, hypercampanulate, with two keeled
primary spirals close to midpoint of whorls.
Description.—Length to 35 mm; pleural angle 10⬚–12⬚. Proto-
conch unknown. Earliest teleoconch whorls present with spirals
‘B’ and ‘C,’ spiral formula dC1B1. Later whorls with weak sec-
ondary spirals, formula du3C1B1r2. Primary spiral ‘A’ absent; spi-
rals ‘B’ and ‘C’ strongly keeled, close to midpoint of whorl. Spac-
ing formula C35B61 (n ⫽ 4). Interspaces with tertiary threads.
Lateral growth line prosocline to orthocline, inflected anteriorly;
lateral sinus moderately deep, apex situated posteriorly. Parietal
area without callus. Base flattened with three to four primary spi-
rals; basal growth line slightly concave. Allmon formula 3-1-M-
A-P/OR.
Material examined.—UWBM 97722, DV 1442-2, L (16.6), W
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU
9.7; UWBM 97723, DV 1442-1, L 25.3, W 7.8; UWBM 97724,
DV 1439-1, L (27.3), W 10.5; UWBM 97725, DV 1154-1, L
(8.3), W 4.0; MUSM INV 81, DV 1439-1, L (22.0), W 7.9;
MUSM INV 82, DV 1439-1, L 30.2, W 10.0; MUSM INV 83,
DV 1439-1, L (13.7), W 5.8. Type locality is Paita, Peru.
Occurrence.—Upper Eocene to Lower Oligocene, northern and
southern Peru.
Discussion.—Specimens of Cristispira cochleiformis differ
from those of C. paracasensis by lacking primary spiral ‘A’ and
having fewer secondary spirals on adult whorls. Specimens of C.
pugetensis have a single, strongly keeled primary spiral (‘B’),
rather than the two of C. cochleiformis.
Cristispira cochleiformis is the younger species of Cristispira
in southern Peru. Specimens occur abundantly at Bajada del Dia-
blo (DV 1442-1) and east of that valley (DV 1323-1) in near-
basal sandstones of the Otuma Formation. Some mollusks found
with C. cochleiformis suggest a Late Eocene age based on their
occurrence in the Chira-Verdun sequence of northern Peru
(DeVries, 2004). A single specimen of C. cochleiformis from Otu-
ma outcrops at Cerro Santa Cruz, near Paracas (DV 1154-1), pro-
vides a means of correlating outcrops from the northern Pisco
Basin with the Bajada del Diablo outcrop (DeVries, 2004).
Genus TURRITELLA sensu lato
Discussion.—Several species of Turritella from Peru are re-
ferred to Turritella s. l. for lack of compelling morphological or
biogeographical reasons to assign them to turritelline genera from
elsewhere in the world. Turritella chira and T. riverae are similar
to each other in many regards; so, too, are T. lagunillasensis and
T. woodsi, but the two pairs differ greatly from each other and
none are like specimens of T. salasi n. sp. and T. cruzadoi n. sp.
While new genera could be created for some of these species, no
biogeographical or evolutionary insight would result from a mul-
tiplication of monospecific or nearly monospecific genera.
TURRITELLA CRUZADOI new species
Figure 4.13–4.18
Diagnosis.—Shell small, early whorls with keeled medial spiral
‘B.’ Adult whorls concave to flat-sided with primary spirals ‘A’
and ‘C’ close to sutures; spiral ‘B’ usually reduced.
Description.—Shell length to 40 mm, pleural angle 12⬚. Early
whorls keeled, adult whorls slightly concave to flat-sided. Sutures
weakly impressed or appressed. Protoconch conic with three to
four whorls. First teleoconch whorl with medial keel, additional
primary triangular spirals appearing on second teleoconch whorl;
spiral formula dc2B1a3. Adult whorls with diminished spiral ‘B’
shifted anteriorly and situated medially or slightly anteriorly be-
tween strong, lightly beaded spirals ‘A’ and ‘C;’ spiral formula
dC2B1A3, dC2b1A3 or, rarely, dC2b1A3r4. Spacing formula
C19B44A77 (n ⫽ 9). Interspaces with numerous tertiary threads;
secondary spirals usually absent. Lateral growth line orthocline
to moderately opisthocline, inflected posteriorly and anteriorly;
lateral sinus deep, apex situated medially or slightly posteriorly
between spirals ‘A’ and ‘B.’ Base flattened, with four to six ter-
tiary threads; basal growth line concave. Allmon formula 3-2-D-
A/B-OR/OP.
Etymology.—Named in honor of Ing. José Cruzado, formerly
head of the paleontology laboratory at PetroPeru.
Types.—Type locality is Cerro Las Salinas (Fig. 5), east of Sa-
linas de Otuma (including collecting localities DV 775, DV 841,
and DV 1131). Chilcatay Formation, uppermost Oligocene to
Lower Miocene. The following specimens are syntypes: UWBM
97726, DV 775-1, L (18.6), W 8.3; UWBM 97729, DV 841-1, L
(26.5), W 7.3; UWBM 97730, DV 841-1, L (19.5), W 7.5;
UWBM 97731, DV 1131-1, L (23.4), W 6.8; UWBM 97732, DV
1131-1, L (8.6), W 3.4; UWBM 97733, DV 775-1, L (25.0), W
339
8.7; UWBM 97734, DV 841-1, L 23.2, W 7.1; UWBM 97735,
DV 841-1, L 10.7, W 3.7; MUSM INV 84, DV 841-1, L (23.0),
W 7.9; MUSM INV 85, DV 1131-1, L (20.1), W 6.8; MUSM
INV 86, DV 1131-1, L 18.6, W 5.7; MUSM INV 87, DV
1131-1, L (6.5), W 3.2; MUSM INV 88, DV 841-1, L (24.9), W
8.4; MUSM INV 89, DV 841-1, L (27.0), W 7.4.
Other material examined.—UWBM 97727, DV 631-12, L
(25.5), W (10.1); UWBM 97728, 80JM 016, block with numerous
specimens. DeVries collection: DV 631-12, block with numerous
specimens; DV 841-1, lot of 17; DV 775-1, lot of 7; DV
1131-1, lot of 9; DV 1403-1, lot of 2; DV 1611-1, lot of 6.
Occurrence.—Uppermost Oligocene to Lower Miocene, south-
ern Peru.
Discussion.—Specimens of Turritella cruzadoi are unlike any
other specimens of Turritella from Cenozoic deposits of northern
Peru or Chile. They do share several attributes with specimens of
the austral Eocene to Oligocene genus, Spirocolpus Finlay, 1927,
including a multispiral protoconch, concave basal growth line,
orthocline to opisthocline lateral growth line with a deep lateral
sinus, reduced spiral ‘B’ on adult whorls, and anterior and pos-
terior inflections on the lateral growth line. Spirocolpus specimens
differ, however, from those of T. cruzadoi in their spiral ontogeny
(C1B2A3) and spacing of primary spirals (C15B52A73, n ⫽ 1).
Specimens of Turritella cruzadoi most resemble those of Early
Miocene Indonesian turritellines assigned to T. herberti Archiac
and Haime, 1853 by Beets (1986), most notably in the extreme
reversal of spiral dominance from dcBa to dCbA or dCA, and
also in the shape of the lateral growth line, which is orthocline
to opisthocline with a deep sinus; the spiral ontogeny (dc2B1a3);
the concave adult whorl profile; and the presence of numerous
tertiary spirals in all the interspaces. The two species differ prin-
cipally in the spacing formula, which for Indonesian T. heberti is
C25B53A81 (n ⫽ 4). A poorly preserved, Early Miocene specimen
of Turritella from Fiji (Ladd, 1972, p. 17, pl. 1, fig. 16) might
bridge the geographic gap between T. cruzadoi and populations
of T. heberti from Indonesia (Beets, 1986) and India (Archiac and
Haime, 1853; Vredenburg, 1928).
Turritella cruzadoi specimens from the type locality at Cerros
Las Salinas occur below a horizon dated with foraminifera at
about 18 Ma and above an uppermost Oligocene/Lower Miocene
angular unconformity marking the base of the Chilcatay Forma-
tion (DeVries, 1998). Specimens are found farther south in basal
sandstones of the Chilcatay Formation between Salinas de Otuma
and the Rı́o Ica.
TURRITELLA LAGUNILLASENSIS Rivera, 1957
Figure 6.1–6.7
Turritella lagunillasensis RIVERA, 1957, p. 174–176, pl. 1, figs. 3, 4; pl.
2, fig. 11.
Diagnosis.—Whorls weakly concave medially. Two primary
spirals near anterior suture; one primary spiral towards posterior.
Numerous secondary spirals and tertiary threads.
Description.—Shell length to 90 mm, elongate, pleural angle
variable, 6⬚–15⬚. Protoconch unknown. Earliest preserved whorls
straight-sided or slightly convex to imbricate, with spiral ‘C’ bor-
dering anterior suture and primary spiral ‘A’ bordering posterior
suture; spiral formula C1A1. Later whorls with addition of spiral
‘B’ and secondary spirals; formula C1B2s3s4A1r4. Adult whorls im-
bricate to concave; formula d5C1T5B2S3S4A1R4r5; order of inser-
tion of numerous secondary spirals and tertiary threads varies.
Spirals ‘C’ and ‘A’ slightly more prominent than all other spirals.
Spacing formula C17B37A73 (n ⫽ 17). Lateral growth line usually
orthocline, inflected anteriorly and, rarely, posteriorly. Lateral si-
nus moderately deep, apex situated posteriorly at posteriormost
340 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 341

spiral ‘S.’ Base flattened on juveniles, convex on adults. Basal

growth line concave. Allmon formula 3/4-1-M/D-A-OR.
Types.—R. Rivera (personal commun., 1986) confirmed that the
type specimen of Turritella lagunillasensis has been lost. The type
locality was the Paracas Peninsula. The neotypic locality is DV
501, Punta Arquillo, Paracas Peninsula. Paracas Formation, Upper
Eocene. Specimen MUSM INV 94 is designated the neotype of
Turritella lagunillasensis. MUSM INV 94, DV 501-2, L (37.9),
W 12.5.
Other material examined.—MUSM INV 95, DV 501-2, L
(27.4), W 10.6; UWBM 97747, DV 585-1, L (32.3), W 13.7;
UWBM 97748, RICE QH-A1, L (48.6), W (16.0); UWBM
97749, DV 544-1, L (14.3), W 8.8; UWBM 97750, DV 637-7, L
(19.3), W (7.0); UWBM 97751, DV 501-2, L (21.5), W 8.4;
UWBM 97752, DV 394-2, L (22.1), W (10.6); UWBM 97753,
DV 501-2, L (9.9). W 4.0; UWBM 97754, RICE QH 87A-1-53m,
L (41.2), W 12.7; UWBM 97755, DV 501-2, L (22.4), W 11.6;
UWBM 97756, DV 501-2, L (26.3), W 15.0; MUSM INV 90,
DV 501-2, L (14.9), W 12.7; MUSM INV 91, DV 394-2, L
(24.0), W (10.8); MUSM INV 92, RICE QH-A1, L (42.0), W
13.3; MUSM INV 93, DV 394-2, L (26.7), W 7.5. DeVries col-
lection: DV 501-2, lot of 14; DV 394-2, lot of 8; DV 585-1, lot
of 1; QH87A-1, lot of 15.
Occurrence.—Middle to Upper Eocene, southern Peru.
Discussion.—Turritella lagunillasensis was thought to be syn-
onymous with T. gilbertharrisi Hodson, 1926 (or T. olssoni Clark
in Clark and Durham, 1946) by DeVries (2001a), but closer in-
spection of Olsson’s (1931) specimens from the Eocene Chira
shales of northern Peru shows otherwise. Both species have sim-
ilar basal growth-line patterns but the lateral growth-line sinus in
T. olssoni is medial, not posterior. The spiral formula for T. ols-
soni is DCbA or even DCA, with spiral ‘b’ being situated me-
dially on the whorl and spiral ‘D’ being entirely exposed and
robust. For T. lagunillasensis, the spiral formula is dCBA or
FIGURE 5—Type locality (DV 775) for Turritella cruzadoi n. sp. Other
dCbA, with spiral ‘B’ located at the anterior third or quarter of locality-samples are mentioned in the text. The neotypic locality for
the whorl. Cristispira paracasensis is north of La Mina in basal orange sandstones
The only turritelline taxa noted by Allmon (1996) to have spiral that include locality-sample DV 634-1.
‘B’ appear last among the three primary spirals are Colpospira
Donald, 1900 and Colpospira (Acutospira) Kotaka, 1959, both
with ranges from the Eocene to present. Enough dissimilarities of whorl and secondary spiral adjacent to posterior suture (spiral
exist to question whether the Australasian and Peruvian taxa are formula C1B1A1r1). Later whorls keeled, telescoped, or concave,
closely related. with spiral ‘A’ as a strong posterior keel. Spiral ‘d’ exposed, ad-
ditional secondary spirals present; formula d2C1t2B1s2A1r1r2; or
TURRITELLA WOODSI Lisson, 1925
with reduction in spirals: formulae d2C1b1A1 or d2c1A1. Strength
Figure 6.8–6.14
of keel ‘A’ and slope of sutural platform vary greatly. Interspaces
Turritella woodsi LISSON, 1925, p. 29–30, pl. 3, fig. 9; RIVERA, 1957, p. with tertiary threads or smooth. Spacing formula C19B37A68 (n ⫽
176–177, pl. 1, fig. 5; pl. 6, fig. 43. 29, 9, 15 for ‘C,’ ‘B,’ and ‘A,’ respectively). Lateral growth line
Turritella portaroi LISSON, 1925, p. 30, pl. 3, fig. 10. prosocline to orthocline, inflected anteriorly and sometimes pos-
Turritella conquistadorana HANNA AND ISRAELSKY, 1925, p. 41, pl. 7, teriorly. Lateral sinus deep, apex situated medially just posterior
fig. 5; OLSSON, 1931, p. 74–75, pl. 12, figs. 1, 2, 3, 5, 7; MARKS, 1951, to spiral ‘B.’ Base strongly convex with five broad but weak
p. 99, pl. 6, fig. 13.
spirals; spiral ‘d’ keeled in juveniles, subdued or obsolete in
Diagnosis.—Strong posterior keel. Anterior portion of whorl adults. Basal growth line concave to sinuous. Allmon formula
straight-sided, usually with reduced spiral sculpture. 5/4-1/2-D-B-P/OR.
Description.—Shell length 60–110 mm, pleural angle 14⬚–18⬚. Material examined.—UWBM 97736, DV 638-1, L (62.8), W
Protoconch unknown. Early teleoconch whorls straight-sided with (24.3); UWBM 97737, DV 621-1, L (41.0), W (16.3); UWBM
three strong spirals evenly spaced across anterior three-quarters 97738, DV 621-1, L (37.2), W (13.0); UWBM 97739, DV

←
FIGURE 4—1–9, Cristispira paracasensis (Rivera, 1957). All specimens from Upper Eocene. All from the Paracas Formation except PRI specimens
from northern Peru. 1, MUSM INV 99, ⫻4.1; 2, MUSM INV 99, basal view, ⫻4.3; 3, UWBM 97746, ⫻3.0; 4, PRI 27786, ⫻3.4; 5, MUSM INV
75, ⫻3.1; 6, PRI 24287, ⫻3.6; 7, PRI 27284, ⫻3.1; 8, UWBM 97719, neotype, ⫻4.1; 9, UWBM 97720, ⫻2.6. 10–12, Cristispira cochleiformis
(Gabb, 1869). All specimens from Late Eocene to Early Oligocene, Otuma Formation. 10, UWBM 97722, ⫻2.8; 11, UWBM 97723, ⫻2.9; 12,
MUSM INV 83, ⫻4.4. 13–18, Turritella cruzadoi n. sp. All specimens are syntypes from uppermost Oligocene to Lower Miocene, Chilcatay
Formation. 13, UWBM 97726, ⫻3.8; 14, UWBM 97733, ⫻5.7; 15, UWBM 97734, ⫻3.9; 16, MUSM INV 89, ⫻3.6; 17, UWBM 97735, ⫻5.7;
18, UWBM 97733, ⫻3.4.
342 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 343

631-8, L (42.9), W (19.5); UWBM 97740, DV 509-2, L (36.9),
W (17.5); UWBM 97741, DV 638-1, L (47.2), W (25.1); UWBM
97742, DV 638-1, L (57.2), W 21.8; UWBM 97743, DV 638-1,
L (32.6), W 11.9; UWBM 97744, DV 502-1, L (12.3), W 6.4;
UWBM 97745, DV 621-1, L (25.7), W (17.7); MUSM INV 106,
DV 638-1, L (42.8), W 22.6; MUSM INV 107, DV 638-1, L
(49.4), W (18.1); MUSM INV 108, DV 638-1, L (28.0), W 12.9;
MUSM INV 109, DV 621-1, L (23.9), W (15.8); MUSM INV
110, DV 621-1, L (38.4), W 16.8; MUSM INV 111, DV 502-1,
L (11.1), W 4.7; MUSM INV 112, DV 1272-1, L (24.8), W 23.7;
PRI 2036, Caleta Mero, L (34.9), W 13.0; PRI 2037, Caleta Mero,
L (31.1), W 13.6; PRI 2038, Caleta Mero, L (23.1), W 12.3; PRI
2040, Caleta Mero, L (30.4), W 13.5. DeVries collection: DV
638-1, lot of 9; JM 314, lot of 32; DV 621-1, lot of 28; DV
502-1, lot of 4; DV 627-1, lot of 6; DV 1442-1, lot of 6. Type
locality is Caravelı́, Peru.
Occurrence.—Uppermost Eocene: southern Peru. Oligocene to
Lower Miocene: southwestern Ecuador to southern Peru.
Discussion.—Lisson (1925) recognized two species from Car-
avelı́, one smaller with a pronounced keel (Turritella woodsi), the
other larger with a less pronounced posterior spiral (T. portaroi).
Later in the same year (July), Hanna and Israelsky (1925) briefly
described a strongly keeled turritelline, T. conquistadorana, from
northern Peru. New material from the Heath and Mancora For-
mations permitted Olsson (1931) to better characterize T. con-
quistadorana, but he overlooked similarities with Lisson’s T.
woodsi. Marks (1951) extended the geographic range of T. con-
quistadorana into Oligocene and Lower Miocene beds in south-
western Ecuador. Rivera (1957) placed T. portaroi in synonymy
with T. woodsi, but with little explanation rejected a relationship
between T. woodsi and T. conquistadorana. Specimens of the
latter, however, are indistinguishable from many smaller speci-
mens of T. woodsi.
Specimens of Turritella woodsi are encountered from the Par-
acas Peninsula to Bajada del Diablo in basal sandstones of the
uppermost Eocene to Lower Oligocene Otuma Formation (De-
Vries, 1998, 2004). Specimens from the Poroma hills, southwest
of Nazca (DV 638-1), and from the type locality at Caravelı́ (DV
1256-1) are associated with gastropods signifying a latest Oligo-
cene to early Middle Miocene age (DeVries, 2001a; DeVries and
Frassinetti, 2003), consistent with a superjacent ash bed at Car-
avelı́ dated radiometrically at 25 Ma (Noble et al., 1985). The
youngest known specimen of T. woodsi, with an estimated age of
22 Ma, was found east of Yesera de Amara, several tens of meters
above the base of the Chilcatay Formation.
A poorly preserved specimen of Turritella (Fig. 6.8) from the
Eocene Paracas Formation, east of Bahı́a de la Independencia
(DV 631-8), has a smaller pleural angle and a less pronounced
posterior spiral keel than younger specimens of T. woodsi, and
lacks inflections on the lateral growth line. Its spiral sculpture
(spiral formula dCtBsArr; spacing formula C18B36A66) and medial
lateral sinus centered on spiral ‘s,’ however, are nearly identical
with those of T. woodsi from the Otuma Formation. The Paracas
specimen may represent a new species, closely related but older.
In his masters thesis, Tavera (1947, p. 7, pl. 2, fig. 1) described
a subquadrate posteriorly carinate turritelline (Turritella ‘trongo-
lensis’) from Upper Eocene to Oligocene strata of Arauco (Fig.
6.15, 6.16). The spiral formula of early whorls (C1B1s1A1r1) and
spacing formula (C18B38A74, n ⫽ 4) are consistent with formulae
of T. woodsi. In older juveniles of T. ‘trongolensis,’ the spiral
sculpture becomes more complex (spiral formula dC1t2B1s1A1r2)
and interspaces are filled with tertiary threads. Adult specimens
have a reduced sculpture (dc1b1s1A1r2). The lateral sinus is in-
flected anteriorly and situated posteriorly with the apex centered
just posterior to spiral ‘s.’ The base is convex and spiral ‘d’ ob-
solete on the ultimate whorl of adult specimens. The many char-
acter states shared with specimens of T. woodsi suggest some
degree of affinity, but the absence of a second spiral ‘r’ and pres-
ence of numerous tertiary threads on Chilean specimens suggest
T. ‘trongolensis’ is a separate taxon.
The heavy posterior keel on specimens of Turritella woodsi
invites comparison with two similarly keeled fossil Turritella
from California. Specimens of the Eocene T. meganosensis pro-
tumescens Merriam and Turner, 1937 have a posterior spiral that
becomes pronounced much later in their ontogeny and have bet-
ter-developed secondary spirals. Specimens of T. carrisaensis An-
derson and Martin, 1914, from Middle to Upper Miocene strata
of California (Merriam, 1941), have a lateral growth-line pattern
and adult spiral sculpture like that of T. woodsi, including a ten-
dency towards gerontism and an obsolete spiral sculpture. Spec-
imens of T. carrisaensis differ in exhibiting greater variability in
the development of spirals ‘B’ and ‘C’ and secondary spirals.
TURRITELLA SALASI new species
Figure 6.17
Diagnosis.—Shell elongate, flat-sided. One primary spiral bor-
dering anterior suture, another forming weak posterior carina.
Description.—Shell size indeterminate, at least 20 mm long,
elongate, pleural angle 14⬚. Whorl flat-sided; sutures impressed,
exceedingly shallow. Protoconch and earliest whorls unknown.
Early whorls with spiral formula C1B1A1 and hints of secondary
spiral ‘r;’ later whorls with reduction in spiral sculpture, formula
C1A1. Spiral ‘C’ at anterior suture; spiral ‘A’ carinate; both prom-
inent on all whorls; spacing formula C13B45A69 (n ⫽ 1). Lateral
growth line poorly preserved; probably prosocline, uninflected,
with lateral sinus deep, posteriorly situated, apex on spiral ‘A.’
Base flattened, mostly missing. Allmon formula ?-4-D-A-P.
Etymology.—Named in honor of Rodolfo Salas-Gismondi, a Pe-
ruvian vertebrate paleontologist with the Museo de Historia Nat-
ural, Universidad de San Marcos, Lima.
Type.—Type locality is a bioclastic sandstone horizon above an
angular unconformity at the foot of Cerros Tinajones, west of
Ullajalla (locality DV 1608; Fig. 7). UWBM 97757, holotype,
DV 1608-1, L (19.7), W 7.2.
Occurrence.—Lower Middle Miocene, southern Peru.
Discussion.—Among the few species from Peru with a well-
developed primary spiral at the anterior suture is Turritella bifas-
tigata Nelson, 1870, which has a growth-line pattern similar to
that of T. salasi but also a spiral adjacent to the posterior suture
and an intervening, finely threaded concave whorl. Specimens of

←
FIGURE 6—1–7, Turritella lagunillasensis Rivera, 1957. All specimens from Upper Eocene, Paracas Formation. 1, MUSM INV 91, ⫻2.4; 2, MUSM
INV 93, ⫻3.2; 3, MUSM INV 90, basal view, ⫻2.1; 4, UWBM 97753, ⫻6.0; 5, UWBM 97752, ⫻3.3; 6, MUSM INV 92, ⫻2.3; 7, UWBM
97748, ⫻1.5. 8–14, Turritella woodsi Lisson, 1925. 8, UWBM 97739, Upper Eocene, Paracas Formation, ⫻1.3; 9, UWBM 97740, Upper Eocene
or Lower Miocene, ⫻1.7; 10, MUSM INV 107, Uppermost Oligocene, Chilcatay Formation, ⫻1.8; 11, MUSM INV 110, Eocene/Oligocene, Otuma
Formation, basal view, ⫻1.9; 12, PRI 2036, Oligocene, Caleta Mero, ⫻1.8; 13, UWBM 97737, Eocene/Oligocene, Otuma Formation, ⫻2.4; 14,
UWBM 97738, Eocene/Oligocene, Otuma Formation, ⫻2.0. 15, 16, Turritella ‘trongolensis’ Tavera, 1947. Tavera collection, Arauco, Eocene. 15,
⫻3.5; 16, ⫻1.5. 17, Turritella salasi n. sp. UWBM 97757, holotype, Lower Middle Miocene, Pisco Formation, ⫻3.7.
344 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
FIGURE 7—Type locality (DV 1608) for Turritella salasi n. sp. Other
locality-samples are mentioned in the text, including those with the
youngest-known (DV 1611-1) and possibly oldest-known (DV 509-2)
specimens of Turritella woodsi.
T. filicincta Gryzbowski, 1899 and T. varicosta Olsson, 1932 have
an anterior sutural keel but several additional spirals posteriorly.
Specimens of T. woodsi have a similar posterior carina or keel,
but their spirals ‘A’ and ‘B’ are located closer to the anterior
suture and they have three early primary spirals, not two.
TURRITELLA CHIRA Olsson, 1928
Figure 8.1–8.5
Turritella chira OLSSON, 1928, p. 66, pl. 14, fig. 5; CLARK AND DURHAM,
1946, p. 24, pl. 23, fig. 24.
Not Turritella chira OLSSON, 1928. OLSSON, 1930, p. 67, pl. 12, figs. 1–4
(referred herein to Turritella paracasensis RIVERA, 1957).
Turritella prenuncia cruziana OLSSON, 1932. DEVRIES, 1998, fig. 4d.
Turritella sp. DEVRIES, 2004, fig. 7.
Diagnosis.—Shell small- to medium-sized, frustate, broadly
biangulate. Lateral growth line with anterior inflection, posterior
apex.
Description.—Shell length under 40 mm, pleural angle 13⬚–16⬚.
Whorls frustate to convex on latest whorls. Sutures moderately
impressed, deeply excavated posteriorly. Protoconch unknown.
Early whorls with spiral formula c2B1a3, followed by C2B1a3 and
C2B1A3. Later whorls usually broadly biangulate anteriorly with
strong spirals ‘C’ and ‘B’ and with additional secondary spirals
and tertiary threads; spiral formula C2t6B1s5A3r4r4 and finally
u7C2T6B1S5A3r4r4. Spacing formula C28B57A71 (n ⫽ 6). Lateral
growth line orthocline to slightly prosocline with anterior inflec-
tion. Lateral sinus moderately deep, apex slightly posterior, be-
tween spirals ‘B’ and ‘s.’ Base with about six primary and ad-
ditional secondary spirals; basal growth line radial. Allmon
formula 2-1-M-A-P/OR.
Material examined.—PRI 3633, holotype, Casa Saman, north-
ern Peru, L (33.7), W 13.0; UWBM 97716, DV 1442-1, L (25.1),
W 8.6; UWBM 97717, DV 1174-1, L (14.7), W 9.0; UWBM
97718, DV 1442-2, L (21.1), W 12.3; MUSM INV 74, DV
1442-1, L (25.3), W 12.0. DeVries collection: DV 1442-1, lot of
8; DV 1442-1, lot of 6.
Occurrence.—Upper Eocene to Lower Oligocene, southern
Peru.
Discussion.—Four specimens of ‘Turritella chira’ from Yasila
have been reassigned to Cristispira paracasensis, leaving only the
holotype from Casa Saman, northern Peru, and material from the
Eocene of Colombia (Clark and Durham, 1946). Specimens of T.
chira from the Paracas Formation east of Cerros Canastones (DV
1174-1) and Otuma Formation at Bajada del Diablo (DV
1442-1) always have a spiral ‘t,’ which is missing in the holotype,
and have more strongly developed spirals ‘s,’ ‘A,’ and ‘r.’ Spec-
imens of T. chira are smaller than those of T. riverae n. sp., (40
mm vs. 100 mm long) and have the apex of the lateral sinus
situated posteriorly between spirals ‘B’ and ‘s,’ not medially be-
tween spirals ‘t’ and ‘B.’
TURRITELLA RIVERAE new species
Figure 8.6–8.11
Diagnosis.—Shell large, frustate, biangulate. Lateral sinus of
growth line with medial apex.
Description.—Shell length to 100 mm, pleural angle 16⬚.
Whorls frustate; sutures moderately impressed. Protoconch and
earliest whorls of teleoconch unknown. Early whorls with spiral
formula C1B1, becoming d6u5C1t3B1a2r4r4. Spirals ‘C’ and ‘B’
prominent, rounded, producing biangulate anterior profile; spiral
‘B’ at periphery of whorl. Interspace C/B wider than B/A; spacing
formula C30B55A71 (n ⫽ 10). Lateral growth line orthocline to
prosocline, with anterior inflection and, rarely, posterior inflection.
Lateral sinus shallow, with medial apex between spirals ‘B’ and
‘t.’ Base convex, with six strong spirals, sometimes with inter-
calated threads. Basal growth line radial. Allmon formula 2-1/2-
S-B-P/OR.
Etymology.—Named in honor of Dr. Rosalvina Rivera, a Pe-
ruvian paleontologist active for many decades.
Types.—The type locality is a bioclastic cross-bedded sandstone
on Pampa Alto Gramadal, south of Caravelı́ (locality DV 1256;
Fig. 9). Camaná Formation, uppermost Oligocene. UWBM
97712, holotype, L (47.4), W 18.6. All other specimens are para-
types from DV 1256-1. UWBM 97713, L (7.7), W 4.8; UWBM
97714, L (31.2), W 16.5; UWBM 97715, L (43.8), W (25.0);
MUSM INV 100, L (16.9), W 7.8; MUSM INV 101, L (52.0),
W 24.9; MUSM INV 102, L (27.5), W 11.5.
Other material examined.—DeVries collection, DV 1256-1, lot
of 6.
Occurrence.—Uppermost Oligocene, southern Peru.
Discussion.—Specimens of Turritella riverae are very similar
to those of T. chira from Bajada del Diablo in the Pisco Basin
and the holotype of T. chira from the Talara Basin of northern
Peru, differing principally by being much larger and having the
apex of the lateral sinus situated medially. Several specimens of
T. riverae lack a spiral ‘t,’ as does the holotype of T. chira.
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 345

FIGURE 8—1–5, Turritella chira Olsson, 1928. 1, PRI 3633, Upper Eocene, Casa Saman, northern Peru, ⫻2.5; 2, MUSM INV 74, Eocene/Oligocene,
Otuma Formation, ⫻2.7; 3, UWBM 97718, Eocene/Oligocene, Otuma Formation, ⫻2.6; 4, UWBM 97716, Eocene/Oligocene, Otuma Formation,
⫻2.5; 5, UWBM 97717, Upper Eocene, Paracas Formation, ⫻3.4. 6–11, Turritella riverae n. sp. All from Upper Oligocene, Camaná Formation.
6, UWBM 97714, paratype, ⫻1.6; 7, MUSM INV 102, paratype, ⫻2.7; 8, UWBM 97712, holotype, ⫻1.8; 9, UMBM 97715, paratype, basal view,
⫻1.7; 10, UWBM 97713, paratype, ⫻4.9; 11, UWBM 97715, paratype, ⫻2.0.

DISCUSSION
The Incatella clade.⎯Incatella turritellines, which are mostly
endemic to Peru and Chile, appear to have evolved prior to the
latest Oligocene on the southwestern or perhaps southeastern
coast of South America [I. trilirata; also Argentinian species doc-
umented by Ortmann (1902), Ihering (1907), and del Rı́o (2004)].
Their evolutionary history in western South America can be in-
terpreted in two ways, depending on the emphasis given to strati-
graphic occurrences of different morphotypes, and the manner in
which characters (pleural angle, depth of suture, and differentia-
tion of primary, secondary, and tertiary spirals) and character var-
iability within and between populations are defined.
Employing existing taxonomy, the Incatella lineage may be
seen as passing gradually from smaller and less massive taxa in
the Early to Middle Miocene (I. trilirata, I. hupei new name,
possibly smaller I. leptogramma) to a succession of larger and
thicker taxa during the Late Miocene, Pliocene, and Pleistocene
(larger I. leptogramma, I. chilensis, I. cingulatiformis, I. cingu-
lata). This taxonomy gives greater weight to stratigraphic occur-
rence as a criterion for recognizing different populations of a
gradually evolving lineage and incorporates an expectation of
modest temporal spans for individual species. Alternatively, one
can view all elongate tricordate specimens as a single species
ranging from the latest Oligocene to earliest Pleistocene, subsum-
ing all Incatella taxa except I. cingulata under the oldest available
name, I. chilensis. Specimens assigned to I. leptogramma, a spe-
cies that spans the Middle to Late Miocene, would be considered
only multicordate ecophenotypic variants of this hypothetically
346 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
FIGURE 9—Type locality (DV 1256) for Turritella riverae n. sp., south
of Caravelı́, and locality-sample (DV 1258-1) with oldest Peruvian
specimen of Turritella trilirata (Fig. 3.19).
long-lived species. This alternative taxonomy incorporates a
greater degree of intraspecific variability and does not reject a
duration of 23 million years for an Incatella species.
The first alternative is favored here, thereby giving emphasis
to character differences typically observed between stratigraphi-
cally discrete assemblages. Within-assemblage character variation
for Incatella taxa is substantial, however, which does obscure tax-
onomic boundaries when too few individuals are examined.
Other species groups.⎯Another species group consists of Cris-
tispira paracasensis and C. cochleiformis. With the discovery of
the latter in southern Peru and the assignment of northern Peru-
vian specimens once identified as Turritella chira to the former,
the genus can now be considered to have been present along the
entire coast of Peru from the Late Eocene to Early Oligocene.
Both species of Cristispira are found in near-basal transgressive
sandstones from comparable inner shelf to foreshore paleoenvi-
ronments.
Two pairs of Turritella species from southern Peru exhibit sim-
ilar patterns of taxon longevity and geriontic gigantism. Within
the first group, T. chira ranged from northern to southern Peru
during the Late Eocene and possibly earliest Oligocene. Individ-
uals of T. riverae n. sp. from uppermost Oligocene beds near
Caravelı́ are nearly identical with those of T. chira except for
their very large size and medial position of the lateral sinus apex.
Within the second group, smaller specimens of T. woodsi from
Eocene/Oligocene Otuma beds of southern Peru are replaced by
unusually large (⬎100 mm) waveworn specimens of T. woodsi
from Upper Oligocene beds near Caravelı́.
Two specimens of T. woodsi in upper Eocene beds of the Par-
acas Formation (Fig. 6.8, 6.9) exhibit features in common with
younger specimens of T. woodsi and older specimens of T. la-
gunillasensis, particularly with respect to the spacing of spirals
and lateral and basal growth-line pattern, suggesting that T. wood-
si may have evolved during the Late Eocene from T. lagunilla-
sensis.
Biogeographic patterns, paleoceanographic explanations.⎯The
most striking biogeographic feature of the western South Amer-
ican turritelline fauna is the contrast between Paleogene and Neo-
gene provincial affinities. During the Eocene and Oligocene, most
turritelline species from southern Peru (Turritella chira, T. wood-
si, Cristispira paracasensis, C. cochleiformis) were also present
in northern Peru, some even in southwestern Ecuador (Marks,
1951) and Colombia (Clark and Durham, 1946). The only mem-
ber of this group in central Chile may have been T. ‘trongolensis.’
The extinction of these species in southern Peru coincided with
a global decline in sea-surface temperatures at the end of the
Eocene and Early Oligocene (Miller et al., 1987), which may have
provoked an equatorward contraction of a paleo-Panamic-Peru-
vian Faunal Province.
Miocene, Pliocene, and Pleistocene turritellines from southern
Peru belonging to Incatella n. gen. (I. trilirata, I. hupei, I. lep-
togramma, I. chilensis, I. cingulatiformis, I. cingulata) also occur
in deposits of comparable age in Chile but, except for I. cingulata,
not in northern Peru. Those Neogene species from southern Peru
not found in Chile (Turritella infracarinata, T. cruzadoi n. sp., T.
salasi n. sp.) are not part of the Incatella clade. Turritella cru-
zadoi is unique in western South America, but shares many char-
acters with the Early Miocene Indonesian T. heberti. Turritella
infracarinata is an early Middle Miocene addition to the southern
Peruvian fauna from northern Peru; its assignment to Turritella
(Turritella) also implies an Indo-Pacific affinity. Turritella salasi
may have arrived from northern Peru, if it is related to the mor-
phologically similar T. filicincta. These turritellines not belonging
to Incatella were extinct by the end of the early Late Miocene.
The shift from equatorial to high-latitude provincial affinities
in the southern Peruvian fauna is evident for the entire molluscan
fauna from southern Peru. Most molluscan species from the Par-
acas and Otuma formations are also found in Eocene and Early
Oligocene deposits of northern Peru (Rivera, 1957; DeVries,
2004), whereas a majority of molluscan species in upper Miocene
and Pliocene deposits of southern Peru are also found in Chile
(DeVries and Frassinetti, 2003). Pleistocene and modern mollus-
can faunas from southern Peru and Chile continue to be quite
similar (DeVries, 2001b; DeVries and Frassinetti, 2003).
The emergence of a late Neogene turritelline fauna in southern
Peru with Chilean biogeographic affinities resulted from the ex-
tinction of equatorial turritellines in southern Peru throughout the
late Paleogene and early Neogene; the failure of species from a
diverse Neogene turritelline fauna in northern Peru to establish
themselves in southern waters, except during the early Middle
Miocene, when Turritella infracarinata and T. salasi appeared;
and the persistence until today of the Incatella clade in western
South America. From the late Late Miocene onward, the Peruvian
Faunal Province has supported just one turritelline species, first
I. cingulatiformis (Late Miocene to Early Pleistocene) and later
I. cingulata (Middle Pleistocene to Recent).
The biogeographic behavior of a few turritelline taxa during
the latest Oligocene to earliest Miocene is problematic. The last
of the Paleogene species, the giants Turritella woodsi and T. riv-
erae, dominated high energy shoreface environments until 22 Ma,
then disappeared, as did the small endemic T. cruzadoi, just as
the first populations of the Incatella clade were appearing. Later,
species with equatorial affinities, T. infracarinata and T. salasi,
moved poleward into southern Peru during the early Middle Mio-
cene. The former thrived despite contemporaneous global cooling,
but disappeared by 9 Ma.
Turritella diversity, upwelling, and productivity.⎯Allmon
(1988, 1992) has proposed that turritellines thrive in the presence
of nutrient-rich seasonally upwelled waters. Menge et al. (1997)
linked productivity in modern coastal upwelling zones to the
abundance of invertebrate grazers, although others have been
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU 347

TABLE 2—Temporal range of turritelline species from southern Peru and species richness by two-million-year increments. Question marks are assigned to
presumed occurrences that unite older and younger known occurrences of the same or related species.

Age (Ma) Species of Turritellines Sum of Species

0–2 cingulata 1
2–4 cingulatiformis 1
4–6 cingulatiformis chilensis 2
6–8 cingulatiformis chilensis 2
8–10 leptogramma 1
10–12 leptogramma infracarinata 2
12–14 leptogramma infracarinata 2
14–16 hupei new name leptogramma salasi n. sp. infracarinata 4
16–18 hupei new name leptogramma 2
18–20 hupei new name 1
20–22 trilirata 1
22–24 trilirata cruzadoi n. sp. woodsi 3
24–26 trilirata cruzadoi n. sp. riverae n. sp. woodsi 4
26–28 chira (?) woodsi (?) 2
28–30 chira (?) woodsi (?) 2
30–32 chira (?) woodsi (?) 2
32–34 cochleiformis chira woodsi 3
34–36 cochleiformis chira woodsi 3
36–38 cochleiformis chira woodsi 3
38–40 lagunillasensis paracasensis chira woodsi 4
40–42 lagunillasensis paracasensis chira 3

equivocal on this point (Wooten et al., 1996). The inferred con- 1993), and dominated today by high productivity associated with
nection between neritic primary productivity and both turritelline the Gulf of Guayaquil and Guayas Estuary (Twilley et al., 2001),
and molluscan diversity has been used to explain Pliocene ex- has supported an approximately equal or greater number of tur-
tinction patterns in the northeastern Pacific Ocean and western ritelline species throughout the Cenozoic (Table 3).
North Atlantic Ocean (Allmon, 1992, 1993) and the existence of
refugia for Pliocene species in the Caribbean (Vermeij and Petuch, CONCLUSIONS
1986). Data from western South America challenge the generality The Cenozoic turritelline fauna of southern Peru consists of 15
of these conclusions. species, six of which belong to an endemic South American Neo-
The Peruvian margin has supported exceptionally high levels gene clade (Incatella n. gen.) with a midlatitude austral origin.
of primary and secondary productivity since at least the Middle Most Paleogene turritellines, in contrast, range from southern Peru
Eocene. Key indices of productivity—diatomaceous sediments, towards the equator. Turritelline species richness is less than or
dolomite remineralization, phosphatic concretions, and fish scales
comparable to that of northern Peru and Ecuador, but higher than
of anchoveta and sardines (Suess et al., 1987; Dunbar et al., 1990;
that of Chile. Species richness was at a maximum during the Early
Rønning, 1990; DeVries, 1998)—are present in strata of all sub-
epochs in the Pisco Basin. At present, a strong seasonal cycle of Miocene, when some Paleogene holdover taxa coexisted with the
coastal upwelling and high primary productivity exists along the oldest species of Incatella and turritellines of possible Indo-Pa-
coast of southern Peru (Puillat et al., 2005). Nevertheless, the cific origin, and has held at a long-term minimum since the early
southern Peruvian margin has never been inhabited simultaneous- Late Miocene. Decreasing species richness coincides with a broad
ly by more than three or four turritelline species since the early decline in Paleogene sea-surface temperatures and later with a
Late Miocene and only one or two gradually evolving species thermal isolation of the Neogene Peruvian-Chilean coastline by
from the same clade have occupied the region since that time cold coastal upwelling and the northward flow of the cold water
(Table 2). The coast of northernmost Peru and southwestern Ec- in the Humboldt Current.
uador, characterized as much by siliciclastic shelf and deltaic de-
posits during the Cenozoic as by organic-rich sediments associ- ACKNOWLEDGMENTS
ated with upwelling (DeVries, 1986; Carrozi and Palomino, I wish to thank M. Urbina and R. Salas (Laboratorio de Pa-
leontologı́a de Vertebrados, Lima, Peru), V. Alleman (Universidad
Ricardo Palma, Lima, Peru), the scientific custodians of the Re-
TABLE 3—Species richness of Cenozoic turritellines from northern and south- serva Nacional de Paracas (Peru), D. Frassinetti and H. Nuñez
ern Peru. Data from Olsson (1928, 1930, 1931, 1932), DeVries (1986,
2004), and Alamo and Valdivieso (1997). (Museo Nacional de Historia Natural, Santiago, Chile), the late
E. Valenzuela and R. Martinez-Pardo (both formerly of the De-
Species Numbers Peru partamento de Geologı́a, Universidad de Chile), the late V. Cov-
Time Interval North South acevich (Servicio Nacional de Geologı́a y Minerı́a), and S. N.
Nielsen (Freie Universität Berlin) for their hospitality, logistical
Quaternary 5 1 support, and scientific advice. Photographs of museum specimens
Pliocene 3 1
Late Miocene 2 4 were kindly provided by P. Jeffery (Natural History Museum,
Middle Miocene 3 4 London) and F. Wesselingh (National Museum of Natural History,
Early Miocene 5 5 Leiden, The Netherlands). Reproductions of Dumont d’Urville’s
Late Oligocene 4 5
Early Oligocene 3 2 account of his expedition to the Southern Hemisphere were pro-
Late Eocene 8 4 vided by the Beinecke Rare Book and Manuscript Library, Yale
Middle Eocene 2 2 University. I would like to thank the Conchologists of America
Early Eocene 3 ? for a travel grant for work in Chile in 1993 and the Fulbright
Paleocene 2 ?
Commission for a research scholarship for study in Peru in 1999.
348 JOURNAL OF PALEONTOLOGY, V. 81, NO. 2, 2007
S. N. Nielsen and W. D. Allmon provided helpful critiques of the
manuscript.
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ACCEPTED 18 NOVEMBER 2005
APPENDIX. LOCALITY—SAMPLES
CAS 328.01 Near top of a small hill south of Corona Peak, northern
Peru (G. C. Gester).
CAS 329.02 Ridgeline near Corona Peak, northern Peru (G. C. Gester).
CAS 337.01 West of La Cuesta, near trail to Tumbes, northern Peru (G.
C. Gester).
RICE QH A1 Quebrada Huaricangana. 14⬚57⬘S, 75⬚16⬘W (Palpa
1:100,000 quadrangle). Chilcatay Formation, uppermost
Oligocene.
80JM 016 ‘‘This locality is noted for Lower Pliocene diatomites and
has not been associated with mollusks in other samples.
The sandstone block attributed to this locality probably
was collected elsewhere’’ (J. Macharé). 13⬚42⬘S, 76⬚10⬘W
(Pisco 1:100,000 quadrangle).
82JM 017 Cerro El Huevo, northwest of San Juan de Marcona. Pleis-
tocene marine terraces. 15⬚18⬘S, 75⬚09⬘W (San Juan
1:100,000 quadrangle).
84JM 314 15⬚01⬘S, 74⬚58⬘30⬙W (Acarı́ 1:100,000 quadrangle). Chil-
catay Formation, uppermost Oligocene.
DV 333-21 Cuesta del Alto, near El Alto, northern Peru; gravel road
to wells on west side. Coquina of Early Pleistocene Man-
cora Tablazo exposed on north side of road where it begins
to descend western side of ridge. See DeVries (1986) for
map.
DV 381-1 San Juan-Lomas road, km 47.5. Low on southwest flank
of ridge protruding westward along road. 15⬚22⬘59⬙ S,
75⬚03⬘11⬙ W (San Juan 1:100,000 quadrangle). Pliocene.
DV 381-5 San Juan-Lomas road, km 47.5. Uppermost coquina.
15⬚22⬘59⬙ S, 75⬚03⬘11⬙ W (San Juan 1:100,000 quadran-
gle). Middle Pleistocene.
DV 394-2 Bahia de la Independencia-Comotrana road, 2–3 km in-
land, in valley about 200 m north of road. 14⬚11⬘54⬙S,
76⬚08⬘41⬙W (Punta Grande 100,000 quadrangle). Paracas
Formation, Upper Eocene.
DV 419-4 Filudo Depression, north of Pozo Santo-Laguna Grande
road, low ridge at bottom of eastern side, midsection, silty
sandstone. 13⬚52⬘52⬙S, 76⬚0721⬙W (Pisco 1:100,000 quad-
rangle). Pisco Formation, Middle Miocene.
DV 420-1 Filudo Depression, south end, gray sandstone beds.
13⬚53⬘12⬙S, 76⬚07⬘15⬙W (Pisco 100,000 quadrangle). Pis-
co Formation, Middle Miocene.
DV 465a-1 Terrace outcrop at abandoned San Juan-San Nicholas
bridge, northeast of San Juan de Marcona. 15⬚21⬘S,
75⬚08⬘W (San Juan de Marcona 1:100,000 quadrangle).
Late Pleistocene.
DV 501-2 West side of Punta Arquillo. 13⬚54⬘39⬙S, 76⬚21⬘29⬙W (Pis-
co 1:100,000 quadrangle). Paracas Formation, Upper Eo-
cene.
DV 502-1 Playa Talpo, north side of Paracas Peninsula, west side of
valley. 13⬚48⬘02⬙S, 76⬚20⬘43⬙W (Pisco 1:100,000 quad-
rangle). Otuma Formation, Eocene/Oligocene.
DV 509-2 About 12 km south of Yesera de Amara, sandstones near
base of section, above bedrock. 14⬚39⬘45⬙S 75⬚37⬘44⬙W
(Lomitas 1:100,000 quadrangle). Chilcatay Formation, Up-
per Eocene? Lower Miocene?
DV 523-2 Cerro Terrestral, near red bed on southeastern flank of hill.
14⬚48⬘02⬙S, 75⬚23⬘02⬙W (Palpa 1:100,000 quadrangle).
Pisco Formation, Upper Miocene.
DV 523-4 Cerro Terrestral, just below red bed on southeastern flank
of hill. 14⬚48⬘02⬙S, 75⬚23⬘02⬙W (Palpa 1:100,000 quad-
rangle). Pisco Formation, Upper Miocene.
DV 532a-3 Quebrada Huaricangana, east of Molde de Queso diato-
maceous siltstones. 14⬚55⬘06⬙S, 75⬚15⬘33⬙W (Palpa
1:100,000 quadrangle). Pisco Formation, Upper Miocene.
DV 542-1 Yesera de Amara. Coarse-grained sandstone below an un-
conformity. 14⬚35⬘31⬙S, 75⬚40⬘15⬙W (Lomitas 1:100,000
quadrangle). Chilcatay Formation, Lower Miocene.
DV 544-1 Nine to ten km south-southeast of Yesera de Amara. Basal
sandstones. 14⬚39⬘44⬙S, 75⬚37⬘50⬙W (Lomitas 1:100,000
quadrangle). Chilcatay Formation, Lower Miocene.
DV 571-5 Alto Grande (⫽El Jahuay of Muizon publications), hillside
west of Panamerican Highway, south of intersection with
road northwest to San Juan de Marcona. 15⬚26⬘57⬙S,
74⬚52⬘06⬙W (Yauca 1:100,000 quadrangle). Pisco Forma-
tion, Upper Miocene.
DV 579-2 One km east of double-knobbed hill, south of mouth of
Quebrada Gramonal, east side of the Rı́o Ica. 14⬚46⬘30⬙S,
75⬚30⬘06⬙W (Lomitas 1:100,000 quadrangle). Pisco For-
mation, Middle Miocene.
DV 585-1 About 100–200 m south of the mouth of Quebrada Santa
Cruz. 14⬚55⬘33⬙S, 75⬚28⬘53⬙W (Palpa 1:100,000 quadran-
gle). Paracas Formation, Upper Eocene.
DV 590-1 Cerro Tiza, northernmost of two points, south face.
14⬚40⬘05⬙S 75⬚41⬘18⬙W (Lomitas 1:100,000 quadrangle).
Chilcatay Formation, Lower Miocene.
DV 599-2 South side of Montemar hill, Sacaco Basin. Over Upper
Miocene sandstones overlain by Pleistocene marine terrac-
es. 15⬚32⬘05⬙S, 74⬚47⬘39⬙W (Yauca 1:100,000 quadran-
gle).
DV 621-1 Three to four km south along road from intersection of
roads to Salinas de Otuma and the Paracas Peninsula.
Coarse-grained sandstone unit near base of Otuma For-
mation. 13⬚53⬘37⬙S, 76⬚15⬘45⬙W (Pisco 1:100,000 quad-
rangle). Eocene/Oligocene.
DV 627-1 0.5–1 km west of Necropolis, Paracas Peninsula. 13⬚52⬘S,
76⬚17⬘W (Pisco 1:100,000 quadrangle). Otuma Formation,
Eocene/Oligocene.
DV 631-8 Thin-bedded sandstone at 120 m in section. Northwest of
Loma Cuesta Chilcatay, north of Ica-Comotrana road.
14⬚11⬘S, 76⬚07⬘W (Punta Grande 1:100,000 quadrangle).
Paracas Formation, Upper Eocene.
DV 631-12 Basal sandstone above angular unconformity, northwest of
Loma Cuesta Chilcatay, north of Ica-Comotrana road.
14⬚11⬘S, 76⬚07⬘W (Punta Grande 1:100,000 quadrangle).
Chilcatay Formation, Uppermost Oligocene.
DV 634-1 Road to Las Minas, Paracas Peninsula. Sandstone outcrops
in parking lot at end of road and along low east-facing
cliff. 13⬚54⬘32⬙S, 76⬚18⬘58⬙W (Pisco 1:100,000 quadran-
gle). Paracas Formation, Upper Eocene.
DV 637-7 Rı́o Ica, 0.5 km north mouth of Quebrada Gramonal, east
side. 14⬚45⬘25⬙S, 75⬚01⬘03⬙W (Lomitas 1:100,000 quad-
rangle). Paracas Formation, Upper Eocene.
DV 638-1 Southwest of Nazca, in third gully in from mouth on
southwest corner of low ridge to south of Quebrada Los
Espinos. 15⬚01⬘53⬙S, 74⬚58⬘52⬙W (Acarı́ 1:100,000 quad-
rangle). Chilcatay Formation, uppermost Oligocene.
DV 769-1 San Juan-Lomas road, km 47.5. Terrace capping ridge pro-
truding westward along road. 15⬚22⬘59⬙S, 75⬚03⬘10⬙W
(San Juan 1:100,000 quadrangle). Middle Pleistocene.
DV 775-1 West side Cerro Las Salinas, eastern edge Salinas de Otu-
ma. Sandstone bed about 100 m southeast of small knoll
 DEVRIES—TURRITELLIDAE FROM SOUTHERN PERU
at base of larger hill. 13⬚59⬘27⬙S, 76⬚13⬘33⬙W (Punta
Grande 1:100,000 quadrangle). Chilcatay Formation, Low-
er Miocene. DV 1323-1
DV 841-1 Cerro Las Salinas, east of Salinas de Otuma, sandstone
bed on western face of long outcrop of tilted strata, near
northern end. 13⬚59⬘48⬙S, 76⬚13⬘38⬙W (Pisco 1:100,000
quadrangle). Chilcatay Formation, Lower Miocene. DV 1403-1
DV 937-1 Filudo Depression, northwest to southwest side, below
sandstone horizon with Anadara and above unconformity
with hackly siltstone. 13⬚53⬘12⬙S, 76⬚07⬘15⬙W (Pisco
1:100,000 quadrangle). Pisco Formation, Middle Miocene. DV 1428-1
DV 1021-4 East side of Quebrada Gramonal. 14⬚44⬘19⬙S, 75⬚31⬘02⬙W
(GPS; Lomitas 1:100,000 quadrangle). Pisco Formation,
Middle Miocene.
DV 1131-1 Western face of Cerro Las Salinas, sandstone bed. DV 1439-1
13⬚59⬘48⬙S, 76⬚13⬘38⬙W (GPS; Pisco 1:100,000 quadran-
gle). Chilcatay Formation, Lower Miocene.
DV 1154-1 Lower knoll, north side of Santa Cruz pass, northeast of
Salinas de Otuma. 13⬚59⬘39⬙S, 76⬚11⬘53⬙W (GPS; Pisco DV 1442-1
1:100,000 quadrangle). Otuma Formation, Eocene/Oligo-
cene.
DV 1174-1 Valley’s southeast flank, southeast of Cerro Canastones.
14⬚08⬘37⬙S, 76⬚10⬘16⬙W (GPS; Punta Grande 1:100,000 DV 1442-2
quadrangle). Paracas Formation, Upper Eocene.
DV 1235-4 Northeast side of Quebrada Usaca, northwest of Nazca.
14⬚50⬘S, 75⬚12⬘W (Palpa 1:100,000 quadrangle). Pisco
Formation, Upper Miocene. DV 1608-1
DV 1256-1 South flank of Quebrada del Carrizal. 15⬚56⬘27⬙S,
73⬚19⬘40⬙W (GPS; Caravelı́ 1:100,000 quadrangle). Ca-
maná Formation, Upper Oligocene. DV 1611-1
DV 1258-1 Coquina block along route of INGEMMET section J, Alta
Gramadal, south of Caravelı́. 15⬚56⬘35⬙S, 73⬚19⬘19⬙W
(GPS; Caravelı́ 1:100,000 quadrangle). Camaná Forma- DV 1647-1
tion, Upper Oligocene.
DV 1272-1 Roadcut with basement section, road to Quilca.
16⬚42⬘09⬙S, 72⬚26⬘57⬙W (GPS; Camaná 1:100,000 quad- DV 1648-1
rangle). Camaná Formation, Lower Miocene.
DV 1310-1 Pampa Corre Viento. 14⬚27⬘56⬙S, 75⬚44⬘37⬙W (GPS; Ica
1:100,000 quadrangle). Chilcatay/Pisco Formation, Lower
to Middle Miocene. DV 1653-1
DV 1316-1 Southwest corner of Corre Viento Depression, base of hill,
on floor of quebrada. 14⬚29⬘03⬙S, 75⬚43⬘57⬙W (GPS; Ica
1:100,000 quadrangle). Pisco Formation, Middle Miocene. WJZ 343
DV 1320-1 Pampa Colorado, along road between Ocucaje and Bajada WJZ 346
351
del Diablo. 14⬚25⬘59⬙S, 75⬚49⬘19⬙W (GPS; Ica 1:100,000
quadrangle). Chilcatay Formation, Lower Miocene.
Road between Corre Viento and Bajada del Diablo, where
bluff overlooks plains leading to coastal mountains.
14⬚27⬘08⬙S, 75⬚52⬘19⬙W (GPS; Ica 1:100,000 quadrangle).
Otuma Formation, Eocene/Oligocene.
East of Cerro Sombrero, near road between Laguna
Grande and Pozo Santo. 14⬚05⬘49⬙S, 76⬚11⬘08⬙W (GPS;
Punta Grande 1:100,000 quadrangle). Chilcatay Forma-
tion, uppermost Oligocene.
North end of Corre Viento depression, brown sandstone
beds on northeast corner. 14⬚26⬘41⬙S, 75⬚45⬘34⬙W (GPS;
Ica 1:100,000 quadrangle). Pisco Formation, Middle Mio-
cene.
Bajada del Diablo, head of valley, on hillsides south of
road to beach, soft weathering sandstones. 14⬚28⬘02⬙S,
75⬚54⬘17⬙W (GPS; Ica 1:100,000 quadrangle). Otuma For-
mation, Eocene/Oligocene.
Bajada del Diablo, bluff overlooking ocean, lower part of
section with indurated sandstone beds. 14⬚28⬘59⬙S,
75⬚57⬘05⬙W (GPS; Ica 1:100,000 quadrangle). Otuma For-
mation, Eocene/Oligocene.
Bajada del Diablo, bluff overlooking ocean, upper part of
section with soft sandstone beds. 14⬚28⬘59⬙S, 75⬚57⬘05⬙W
(GPS; Ica 1:100,000 quadrangle). Otuma Formation, Eo-
cene/Oligocene.
Cerros Tinajones, foot of steep hill, bioclastic coarse-
grained sandstone bed. 14⬚35⬘24⬙S, 75⬚38⬘18⬙W (GPS;
Lomitas 1:100,000 quadrangle).
Campsite, Ullujaya West, depression northeast of Cerros Ti-
najones. 14⬚34⬘52⬙S, 75⬚38⬘40⬙W (GPS; Lomitas 1:100,000
quadrangle). Chilcatay Formation, Lower Miocene.
Northeast corner of Cerros Colorado, tuffaceous siltstones.
About 14⬚21⬘S, 75⬚54⬘W (Punta Grande 1:100,000 quad-
rangle). Pisco Formation, Middle Miocene.
Valley southwest of Cerro Colorado; southwest-facing
flank. 14⬚22⬘25⬙S, 75⬚53⬘52⬙W (GPS; Punta Grande
1:100,000 quadrangle). Chilcatay Formation, Lower Mio-
cene.
‘Sula’ site, between Cerro Terrestral and the Rı́o Ica, east
of the ‘labyrinth.’ 14⬚50⬘16⬙S, 75⬚27⬘29⬙W (GPS; Lomitas
1:100,000 quadrangle). Pisco Formation, Middle Miocene.
Coquimbo, Chile. Pleistocene.
Coquimbo, Chile. Pleistocene.