New Miocene sulid birds from Peru and considerations on

their Neogene fossil record in the Eastern Pacific Ocean

MARCELO STUCCHI, RAFAEL M. VARAS-MALCA, and MARIO URBINA-SCHMITT

Stucchi, M., Varas-Malca, R.M., and Urbina-Schmitt, M. 2016. New Miocene sulid birds from Peru and considerations
on their Neogene fossil record in the Eastern Pacific Ocean. Acta Palaeontologica Polonica 61 (2): 417–427.

Boobies and gannets (family Sulidae) are the most specialized plunge divers among seabirds. Their fossil record along
the Pacific coast of South America extends to the early Middle Miocene. Here we describe three new species of sulids:
Sula brandi sp. nov., Sula figueroae sp. nov., and Ramphastosula aguirrei sp. nov., from the early Late Miocene of the
Pisco Formation (Peru). Two of them are relatives of the living genus Sula, which represents medium and large-sized
boobies. A new species of the extinct genus Ramphastosula is also described, adding to the discussion of possible alter-
native feeding strategies among sulids. The fossil record suggests that sympatric sulids exhibit different body sizes at
least since the Oligocene epoch, a strategy related with resource partitioning. Furthermore, we find current analysis and
knowledge of the fossil record unsuitable to evaluate properly seabird diversity changes through time.

Ke y w o rd s : Aves, Pelecaniformes, Sulidae, Sula, Ramphastosula, seabird evolution, Miocene, Peru.

Marcelo Stucchi [aicb.peru@gmail.com], Asociación para la Investigación y Conservación de la Biodiversidad (AICB),

Av. Vicús 538, Lima 33, Peru.
Rafael M. Varas-Malca [varas.m.r@gmail.com] (corresponding author) and Mario Urbina-Schmitt [mariourbina01@
hotmail.com], Departamento de Paleontología de Vertebrados, Museo de Historia Natural “Javier Prado”, Universi-
dad Nacional Mayor de San Marcos, Av. Arenales 1256, Lima 14, Peru.

Received 24 April 2015, accepted 16 September 2015, available online 12 October 2015.

Copyright © 2016 M. Stucchi et al. This is an open-access article distributed under the terms of the Creative Commons
Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited.

Introduction
The family Sulidae currently includes three genera of medi-
um-sized, strictly marine birds of pantropical distribution:
Papasula (the Abbott’s booby), Sula (other boobies) and
Morus (gannets). They are highly specialized for plunge
diving to catch fish, usually dropping from high altitudes
(40–100 m) and reaching depths up to 8 meters below
the water surface (Nelson 1979; Stucchi 2013). In South
America, Sula is the dominant genus on the Pacific coast
being particularly diverse though the west side, whereas
Morus is restricted to southwestern South America and is
completely absent in the East.
The Pacific coast of South America has yielded the most
abundant and extensive fossil record of sulids in the Southern
Hemisphere, from the Middle Miocene (Stucchi and DeVries
2003) to the Recent, recovered almost exclusively from the
Late Miocene beds of the Bahía Inglesa Formation (northern
Chile; Walsh and Hume 2001; Chávez-Hoffmeister 2007,
2008) and the Pisco Formation (Southwestern Peru; Stucchi
2003, Stucchi and Urbina 2004). In contrast to the relatively
scarce Chilean fossil record, sulids are the most abundant
birds in the Pisco Formation, and also the most diverse with
at least three described boobies, one gannet (Stucchi 2003)
and the extinct genus Ramphastosula (Stucchi and Urbina
2004). Furthermore, a high diversity of Miocene species
has also been observed in marine outcrops off the northern
Pacific Ocean, in which up to six species are recognized from
California, USA (Stucchi 2003; Warheit 1992; see Table 1).
Here, we describe two new species of Sula along with a
new species of the extinct genus Ramphastosula, all from
Late Miocene localities of the Pisco Formation, Peru (Fig. 1).
Additionally, we analyze the diversity of sulids from the
Eastern Pacific Ocean over the course of the Neogene. Sula
is diagnosed osteologically following the works of Tets et
al. (1988), and Stucchi (2003; 2013), which describe the dif-
ferences among Sula, Morus, and Papasula, and within the
genus Sula (Stucchi 2013).
For comparative purposes we assigned species to cate-
gories as follows: “small-sized boobies” for S. sula; “me-
dium-sized boobies” for S. variegata, S. leucogaster, and
S. nebouxii; and “large-sized boobies” for S. dactylatra,
S. granti, and S. tasmani (sensu Tets et al. 1988). Anatomical
terminology follows Howard (1980). All materials were col-
lected by one of the authors (MU-S) and are permanently

Acta Palaeontol. Pol. 61 (2): 417–427, 2016 http://dx.doi.org/10.4202/app.00170.2015

418 ACTA PALAEONTOLOGICA POLONICA 61 (2), 2016

Table 1. List of the described species of the family Sulidae showing their geological age and geographical distribution. * Stucchi (2003) proposed
nomina nuda for these species on the basis of classical morphometrical analysis; bold, species described in this study.
Miocene Pliocene Pleisto- Recent-
Species Geographical region Locality
Early Middle Late Early Late cene extinct
Sula universitatis Brodkorb, 1963* × Western Atlantic Ocean Florida, USA
Morus loxostylus (Cope, 1871) × × Western Atlantic Ocean Maryland, USA
Morus vagabundus Wetmore, 1930 × Northeastern Pacific Ocean California, USA
Morus avitus (Wetmore, 1938) × Western Atlantic Ocean Maryland, USA
Morus pygmaea (Milne-Edwards,
× Western Atlantic Ocean France
1874)
Sarmatosula dobrogensis
× Paratethys Romania
(Grigorescu and Kessler, 1977)
Morus olsoni (Grigorescu and
× Paratethys Romania
Kessler, 1988)
Sula brandi sp. nov. × Southeastern Pacific Ocean Peru
Sula figueroae sp. nov. × Southeastern Pacific Ocean Peru
Morus willetti (Miller, 1925) × × Northeastern Pacific Ocean California, USA
Sula pohli Howard, 1958 × × Northeastern Pacific Ocean California, USA
Morus lompocanus (Miller, 1925) × Northeastern Pacific Ocean California, USA
Morus media (Miller, 1925) × Northeastern Pacific Ocean California, USA
Morus stocktoni (Miller, 1935) × Northeastern Pacific Ocean California, USA
Morus magnus Howard, 1978 × Northeastern Pacific Ocean California, USA
Sula sulita Stucchi, 2003 × Southeastern Pacific Ocean Peru
Sula aff. variegata (sensu Stucchi
× Southeastern Pacific Ocean Peru
2003)
Morus peruvianus Stucchi, 2003 × Southeastern Pacific Ocean Peru
Sula magna Stucchi, 2003 × × Southeastern Pacific Ocean Peru
Sula guano Brodkorb, 1955* × × Western Atlantic Ocean Florida, USA
Sula phosphata Brodkorb, 1955* × × Western Atlantic Ocean Florida, USA
Ramphastosula ramirezi Stucchi
× Southeastern Pacific Ocean Peru
and Urbina, 2004
Ramphastosula aguirrei sp. nov. × Southeastern Pacific Ocean Peru
Morus peninsularis Brodkorb, 1955 × Western Atlantic Ocean Florida, USA
Bimbisula melanodactylus Benson South Carolina,
× × Western Atlantic Ocean
and Erickson, 2013 USA
Morus humeralis (Miller and
× Northeastern Pacific Ocean California, USA
Bowman, 1958)
Morus recentior Howard, 1949 × Northeastern Pacific Ocean California, USA
Sula clarki Chandler, 1990* × Northeastern Pacific Ocean California, USA
Morus reyanus Howard, 1936 × Northeastern Pacific Ocean California, USA
Sula tasmani Tets, Meredith,
× Southwestern Pacific Ocean Australia
Fullagar, and Davidson, 1988

stored at the Museo de Historia Natural “Javier Prado” of
the Universidad Nacional Mayor de San Marcos (Lima,
Peru). The age of the Pisco Formation localities follows
Ehret et al. (2012).
Institutional abbreviations.—MUSM, Museo de Historia
Natural “Javier Prado”, Universidad Nacional Mayor de San
Marcos, Lima, Peru; USNM, National Museum of Natural
History, Smithsonian Institution, Washington DC, USA.
Nomenclatural acts.—The electronic edition of this article
conforms to the requirements of the amended International
Code of Zoological Nomenclature, and hence the new names
contained herein are available under that Code from the
electronic edition of this article. This published work and
the nomenclatural acts it contains have been registered in
ZooBank. The ZooBank LSID (Life Science Identifier) for
this publication is: urn: lsid: zoobank.org: pub: 12E70E13-
D7F9-43BB-A388-6A93FF0E4A52.
The electronic edition of this work was published in a
journal with an eISSN 1732-2421, and has been archived
and is available from the following digital repository: http://
www.app.pan.pl/article/item/app001702015.html
STUCCHI ET AL.—MIOCENE SULID BIRDS FROM PERU 419

76°00' W 75°50' W 75°50' W 75°00' W 74°50' W

Paracas
National Ica
Reserve
14°35' S
Pe

15°30' S

In Ba
ru

de y
pe
A

nd
B

en
Sacaco

ce
Ocucaje Montemar
14°40' S
Cerro Lomas

er iv
Colorado Poza

Yauca R
15°35' S
Pacific Ocean Negra

Ica
Pacific Ocean Poza
Roja

Riv
A 20 km
B 20 km

er
Cerro La Bruja Pisco Fm. rocks

Fig. 1. Maps of the Ocucaje (A) and Sacaco (B, modified from Brand 2001) areas, southestern Peru, indicating the type localities of fossil sulids described
in this paper.

Systematic palaeontology
Aves Linnaeus, 1758
Sulidae Reichenbach, 1849
Genus Sula Brisson, 1760
Type species: Sula sula (Linnaeus, 1766) or Sula leucogaster (Boddaert
1783), both Recent, no subsequent designation.
Remarks.—Generic definition follows Tets et al. (1988) and
Stucchi (2003, 2013).
Sula brandi sp. nov.
Fig. 2.
ZooBank LSID: urn:lsid:zoobank.org:act:B338FEDC-FDB1-4C17-88
DB-628776BCA613
Etymology: In honor to Leonard Brand (Loma Linda University, USA)
for his important contributions to Peruvian vertebrate paleontology,
and to the understanding of the geology, stratigraphy, and taphonomic
processes of the Pisco Formation.
Holotype: MUSM 2497; neurocranium lacking interorbital septum,
lacrimals, palatines, pterygoids, jugals, quadrates, and right quadrato-
jugal. Proximal portion of the beak, including most of the right dentary
and angular.
Type locality: MUSM 2497 was collected from the Cerro Colorado
locality (CCO), Pisco Formation, ~23 km west of Ocucaje District,
Ica, Peru (Fig. 1A). Coordinates: 14°21’37.90” S, 75°53’17.90” W.
The specimen (field number A3 of Bianucci et al. 2015) was found
1–2 km southwest of Cerro Colorado (Colorado hill), between the Wari
and Paracas marker beds of its lower allomember (sensu Di Celma
et al. 2015). The level from which the sulids were recovered (about
60 m of the stratigraphic column) is the second most fossiliferous in
terms of quantity and species diversity, having produced more than 50
specimens, among them osteichthyans, elasmobranchs (myliobatids,
Carcharocles, Cosmopolitodus), long-snouted crocodylians, delphi-
nidans (the pontoporiid Brachydelphis, indeterminate delphinidans),
ziphiids (Messapicetus), balaenopteroideans, cetotheriids, and physet-
eroideans (cf. Acrophyseter) (see Bianucci et al. 2015).
Type horizon: The age of this locality has been estimated according to
the presence of the arcid bivalve Anadara sechurana in late Middle
Miocene (ca. 12–13 Ma; Lambert et al. 2010). However, due to the
occurrence of early Late Miocene diatoms Denticulopsis hudstedtii
and Lythodesmium reynoldsii (9.9–8.9 Ma, Tortonian; Barron 2003) in
the referred level, the age is younger than previously thought (see Di
Celma et al. 2015). Thus, it is correlated with El Jahuay level (9.95 Ma;
Muizon and Bellon 1986) and Montemar Norte (about 10 Ma; Stucchi
2007) in the Sacaco area, and possibly with Laguna Seca locality in the
Ocucaje area (Stucchi et al. 2015).
Diagnosis.—Sula with braincase length similar to those of
medium-sized boobies S. variegata and S. leucogaster but
flattened, being the flattest among sulids; wide naso-frontal
hinge, greater laterofacial angle, smaller cerebellar dome,
and narrow sagittal crest. Shallow salt gland fossae and shal-
low postorbital fossae, as occurs in Morus and Papasula,
but in contrast to the deep condition found in all other Sula
species.
Measurements.—See SOM 1 (Supplementary Online Mate-
rial available at http://app.pan.pl/SOM/app61-Stucchi_etal_
SOM.pdf).
Description.—In lateral view (Fig. 2A) the length of the
braincase fits well within the range of medium-sized boobies
(SOM 1). The anterior region of MUSM 2497 has slightly
pronounced supraorbital ridges above the fused frontals,
as well as a straight dorsal profile, as it occurs in most
species of Sula (note that S. variegata has a slightly domed
profile, see Stucchi 2003, 2013). The middle region (pari-
etal, squamosal and laterosphenoid) of MUSM 2497 has the
same shape and disposition of Sula, but with a noticeable
dorso-ventral flattening (SOM 1: measurement 3). Indeed,
MUSM 2497 shows the most dorsoventrally compressed
skull among sulids (SOM 1: proportion 6). In the posterior
region, there is a large sagittal crest and a shorter and com-
pressed prominentia cerebelaris (supraoccipital) under the
posterior projection of the sagittal crest. However, it makes
a more acute angle with respect to the horizontal plane.
In caudal (occipital) view (Fig. 2B), the flattening of
the braincase is better appreciated. The whole region is
proportionally wider than higher (SOM 1: proportion 6),
differing from other sulids. The height at the vertical axis
of the supraoccipital (between the superior border of the
420
foramen magnum and the sagittal crest) represents 60% that
of S. variegata. Additionally, the medial parasphenoid pro-
cesses are more laterally placed than those of recent species.
In dorsal view (Fig. 2C), the face width of MUSM 2497
is wider than those of medium-sized boobies. This is seen
in both the breadth at the preorbital skull roof and the width
of the naso-frontal hinge. The width is greater than those of
S. variegata, S. leucogaster and S. sula, but lies within the
range of S. nebouxii (SOM 1: measurement 5). The breadth is
greater and reaches the range of large-sized boobies (SOM 1:
measurement 6). Therefore, the angle formed by the sagit-
tal axis and the lateral margin of the frontal (latero-facial
angle) is 10° in MUSM 2497, which contrasts with the an-
gle reached by medium-sized boobies and S. granti (20°
and 18°; Fig. 2C). The middle region exhibits a narrower
and longer sagittal crest in comparison to modern species
of Sula, except for S. leucogaster. In the posterior region,
MUSM 2497 has a rounded supraoccipital, which projects
posteriorly to the paroccipital processes of the exoccipitals.
In other species of Sula, the supraoccipital is triangular and
projects more posteriorly or is at the same level of the pa-
roccipital processes. Both latter characters are also distin-
guished in other specimens from the Pisco Formation, for in-
stance MUSM 249 (Aguada de Lomas locality, 8.0–8.8 Ma),
MUSM 309 and 202 (Montemar locality, 8.70–6.45 Ma) and
MUSM 361 (Sacaco Sur locality, 6.59–5.93 Ma), as well as in
Ramphastosula (Stucchi and Urbina 2004).
In ventral view (Fig. 2D), the otic region, the articular
with the quadrate, the basioccipital and medial parasphe-
noidal processes resemble those of living Sula. Likewise,
the salt gland fossae and postorbital fossae are slightly shal-
lower. The mandible is the same size and shape as those of
medium-sized boobies (Fig. 2E).
Remarks.—MUSM 2497 has the typical cone-shaped skull
of extant boobies, although it is more flattened and has
a wider nasofrontal hinge. Thus, we infer a similar feed-
ing ecology for this species. The sole character considered
primitive is the shallow depth of the salt gland fossae and
postorbital fossae which are both found in Morus (e.g.,
USNM 612654) and Papasula (e.g., USNM 560682), as well
as inmature individuals of Sula (Stucchi 2013). Indeed, the
derived condition is seen in adult Sula, which is more in
line with their early divergence (23 Ma) from other sulids
(according to Friesen and Anderson 1997).
Stratigraphic and geographic range.—Tortonian, Pisco
depositional sequence, Pisco Formation, South-Central Peru.
Sula figueroae sp. nov.
Fig. 3.
ZooBank LSID: urn:lsid:zoobank.org:act:F968A99A-AAF4-4066-B653
-B68E1B3E8319
Etymology: In honor to Judith Figueroa (Asociación para la Investi-
gación y Conservación de la Biodiversidad, Peru), for her valuable con-
tributions to the knowledge of the natural history of Peruvian seabirds.
Type material: Holotype: MUSM 2501; nearly complete, well-pre-
served cranium and mandible lacking pterygoids and right lacrimal;
ACTA PALAEONTOLOGICA POLONICA 61 (2), 2016
one partial thoracic vertebra (? 19–21), three free cervical vertebrae
(? 3–7), left scapula, right and partial left humeri, partial right and left
ulnae, partial left radius, left carpometacarpus, pelvis without pubis,
and partial tarsometatarsi. Paratype: MUSM 2502; associated furcula
and sternum, coracoids, scapulae, humeri, distal fragments of ulnae,
radius shaft, proximal fragment of left carpometacarpus, partial right
carpometacarpus, right and distal fragment of left femora, right tibio-
tarsus, and right tarsometatarsus.
Type locality: MUSM 2501 and MUSM 2502 (A1 and A2 field num-
bers, respectively, of Bianucci et al. 2015) were both collected in Cer-
ro Colorado locality, close to Sula brandi (Fig. 1A). MUSM 2501
coordinates: 14°21’47.30” S, 75°53’0.40” W, 596 masl. MUSM 2502
coordinates: 14°21’32.70” S, 75°53’21.90” W, 632 masl.
Type horizon: The same stratigraphic level as Sula brandi; early Late
Miocene (Tortonian).
Diagnosis.—Sula having similar dimensions to the large-
sized boobies S. dactylatra and S. tasmani, but with a more
robust and much shorter humerus.
Measurements.—See SOM 1–3.
Description.—Skull and mandible: In lateral view (Fig. 3A1),
the rostrum is approximately 50% larger than the braincase,
which corresponds to the range of all known Sula, although
S. sula shows a shorter rostrum (SOM 1: measurement 1).
The length of MUSM 2501 is similar to that of S. tasmani,
S. nebouxii, the largest specimens of S. dactylatra (sensu
Tets et al. 1988), and possibly S. granti. The mandible has
the same shape and proportions as those of extant species.
The braincase length lies in the same range of large-sized
boobies, but is shorter than that of S. tasmani (SOM 1:
measurement 2). The anterior region of MUSM 2501 has
slightly pronounced supraorbital edges above the fused
frontals, a slightly domed skull roof, rather smaller than
that of S. variegata (see Stucchi 2003, 2013). Posterior and
middle regions of the braincase are slightly flattened com-
pared with other species of Sula (SOM 1: proportion 3).
Paroccipital processes of exoccipitals are lower with respect
to the horizontal plane as in MUSM 220 (see Stucchi 2003).
The width of the skull at the postorbital processes in dorsal
view (Fig. 3A4) is proportionally similar to those of the more
robust specimens of large-sized boobies, but with a larger
postorbital process (SOM 1: measurement 8). A laterally
narrow skull is observed in caudal view (Fig. 3A 2), which
is more evident in its constricted foramen magnum (SOM 1:
proportion 4). Shallow salt gland fossae and postorbital fos-
sae, as observed in Morus and Papasula, differing from the
deep condition seen in Sula.
Postcranium: The humerus (Fig. 3B5–B7) is similar to that
of extant species of Sula (see Tets et al. 1988; Stucchi 2003)
and S. magna, but more robust (SOM 2). The humerus length
of both MUSM 2501 and 2502 falls within the range for S.
nebouxii, their proximal width approaches those of S. dactyl-
atra and S. tasmani, and their distal and mid-shaft maximum
width are greater than that of S. tasmani. In lateral view (Fig.
3B6), a deformity (healed injury) is observed in the right
humerus of MUSM 2502. The proximal width of the ulna is
also wider in relation to the distal width of the humerus. The
STUCCHI ET AL.—MIOCENE SULID BIRDS FROM PERU
A B
C
Fig. 2. Sulid bird Sula brandi sp. nov., holotype MUSM 2497 from Cerro Colorado locality, Peru, Pisco Formation, early Late Miocene. Skull in lateral
(A), posterior (B), dorsal (C), and ventral (D) views; mandible in dorsal view (E).
remaining postcranial elements share a similar morphology
with the other species of Sula as Warheit (1992) and Stucchi
(2003) have pointed out. Some details could be observed when
comparing S. figueroae with S. variegata in Stucchi (2011).
Moreover, their proportions notably correspond to different
species. Thus, carpometacarpus and tibiotarsus are similar
in length to that of S. tasmani, whereas the tarsometatarsus is
proportionally closer to S. dactylatra. The femur matches the
range of those of indeterminate Sula individuals (see Stucchi
2003), which is larger than those of both S. dactylatra and S.
tasmani (SOM 3). In addition, the sternum, pelvis and synsa-
crum reach the dimensions and proportions of large-sized
boobies (including the Sula spp. group), wider than those of
medium-sized boobies and considerably larger than that of S.
sula (SOM 3), but are smaller than those of S. magna. Based
on a small sample, we could tentatively say that the pelvis
shows a conspicuous lateral crest on the spina dorsolateralis
ilii, which is also found in S. leucogaster but shallow, and
completely absent in S. variegata.
Remarks.—Although S. figueroae has a notably robust skull
roof, it is narrow at the occipital region. This, however, does
421
E
10 mm
not affect the overall conical shape of the skull. Among the
postcranial elements, the humerus is remarkable because
of its proportions, being quite short relative to the proximal
and distal widths. Thus, the humerus length is the same of S.
nebouxii, even though its distal and proximal widths equal
those of S. tasmani and S. dactylatra, which in turn are
quite longer. The proportions concerning the length of the
other postcranial bones of S. figueroae in comparison with
other Sula species should be carefully addressed because
of the limited sampling of the fossil (only two incomplete
specimens). Warheit (1992) and Stucchi (2010) have already
noted the importance of the quantity of specimens when
evaluating the trends in dimensions and proportions (i.e. in-
traspecific variation) in a species. Despite that, the morphol-
ogy and morphometric data of this extinct booby are more
in line with the feeding and hunting strategies of its extant
relatives, having the same basic body plan of the genus.
Genus Ramphastosula Stucchi and Urbina, 2004
Type species: Ramphastosula ramirezi Stucchi and Urbina, 2004, Saca-
co Sur locality (Poza Roja), Pisco Formation (Peru), Messinian (Late
Miocene).
422 ACTA PALAEONTOLOGICA POLONICA 61 (2), 2016

A1 A2 A3

10 mm

A4 A5 B1 B2 B3 B4

10 mm

10 mm

B5

B6 10 mm

B7

A6 A7 A8 B9 B 10 B 11 B 12
B8

10 mm

Fig. 3. Sulid bird Sula figueroae sp. nov. from Cerro Colorado locality, Peru, Pisco Formation, early Late Miocene. A. Holotype MUSM 2501. Skull in lat-
eral (A1), posterior (A2), dorsal (A4), and ventral (A5) views. Isolated lacrimal in lateral view (A3). Left carpometacarpus in ventral view (A6). Synsacrum
and pelvis in ventral (A7) and lateral (A8) views. B. Paratype MUSM 2502. Sternum in lateral (B1) and ventral (B2) views. Coracoid in (B3) dorsal view.
Left scapula in (B4) lateral view. Right humerus in anterior (B5), lateral (B6), and posterior (B7) views. Proximal portion of ulna in ventral view (B8). Right
femur in anterior view (B9). Right tibiotarsus in anterior view (B10). Right tarsometatarsus in anterior (B11) and plantar (B12) views.
STUCCHI ET AL.—MIOCENE SULID BIRDS FROM PERU 423

A1 A2

10 mm

A3 A4 B C

10 mm D
Ramphastosula other sulids 10 mm
(R. aguirre)
(C–E)

A5 A6
E
10 mm

Fig. 4. Sulid bird Ramphastosula aguirrei sp. nov. from Poza Negra, Sacaco Sur (Peru), Pisco Formation, Late Miocene. A Holotype MUSM 665. Skull
in lateral (A1), posterior (A2), dorsal (A3), and ventral (A4) views. Left quadrate in (A5) anterior and (A6) posterior views. B. Section of rostrum at its mid-
length showing differences in the outline of Ramphastosula and other sulids. C–E. Comparison of the ear region in ventral view in the Peruvian booby
Sula variegata DPV AM P8 (C) from Isla Lobos de Afuera, Lambayaque (Peru), Recent; R. ramirezi MUSM 264 (D) from Poza Roja , Sacaco Sur (Peru),
Messinian (Late Miocene); and R. aguirrei MUSM 665 (E) from Poza Negra, Sacaco Sur (Peru), Messinian (Late Miocene).

Emended diagnosis.—Ramphastosula can be diagnosed Ramphastosula aguirrei sp. nov.

from other sulids by the following combination of char-
acters: (i) height of the rostrum at its mid-length is more
than half the height at the nasofrontal hinge; (ii) slightly to
strongly convex rostrum in all its dorsal edge; (iii) lerome-
dially compressed rostrum forming a triangle in anterior
view, in which the dorsal surface shows a strong ridge; (iv)
well-developed occipital and temporal regions; (v) broad pa-
roccipital procceses of the exoccipitals projecting laterally
to the level of the postorbital processes; (vi) broad temporal
fossae; (vii) robust frontal region.
Ramphastosula ramirezi Stucchi and Urbina, 2004
Emended diagnosis.—Height of the rostrum at its mid-
length is higher than the height at the nasofrontal hinge, and
all its dorsal edge is strongly convex.
Fig. 4.
ZooBank LSID: urn:lsid:zoobank.org:act:51F31FFF-4D44-45B6-A221
-07A128E4A5F3
Etymology: In honor to Walter Aguirre (MUSM, Peru), for his sub-
stantial work as a fossil preparator of the vertebrate paleontological
collection of the MUSM.
Holotype: MUSM 665; almost complete, well-preserved skull with
partial jugal bars, and lacking lacrimals and the right quadrate.
Type locality: MUSM 665 was collected from the Sacaco Sur (Sud-Sa-
caco or SAS) locality (Pisco Formation, Peru) which is dated to 6.59–
5.93 Ma (Ehret et al. 2012) (Fig. 1B). There are two areas (sublocal-
ities) recognized there: Poza Roja (red hole) and Poza Negra (black
hole). The first refers to the type locality described in Muizon and
DeVries (1985), where R. ramirezi was recovered. The second corre-
sponds to underlying beds (18 m lower in the stratigraphic column of
424
Leonard Brand, unpublished material) being older than the former one
and where R. aguirrei was recovered.
Type horizon: Sacaco Sur was formally dated as Pliocene age (Muizon
and DeVries 1985), but new radiometric data suggests a Late Miocene
age, (Messinian; see Ehret et al. 2012).
Diagnosis.—Height of the rostrum at its mid-length is 2/3
the height of the nasofrontal hinge, and its entire dorsal edge
is slightly convex.
Measurements.—See SOM 1.
Description.—Rostrum: R. aguirrei has a robust and large
rostrum, slightly larger than that of R. ramirezi. In lateral
view, the lateral sulci (Fig. 4A1, A3) are oriented parallel to
the culmen due to the ossification of primary nasal aper-
tures. These sulci divide the rostrum rather equally into dor-
sal and ventral parts as in R. ramirezi, but in Sula, Morus,
and Papasula the dorsal part comprises 1/3 of the rostrum
(see Fig. 4A1). Interestingly, we note that in S. figueroae
this division is closer to the condition in Ramphastosula
although this could be the product of lateromedial defor-
mation. Furthermore, the height of the rostrum at its mid-
length equals 2/3 the maximum height (at the level of the
nasofrontal hinge). In S. variegata and generally among
living sulids, this height equals 1/2 the maximum height,
and in R. ramirezi the height at the mid-length of its rostrum
is higher than the level of the nasofrontal hinge. Some indi-
viduals of the living species S. leucogaster and S. dactylatra
show a slender protuberance in the posterior part of the
rostrum (see Stucchi 2013: fig. 4). The rostrum is projected
downward with respect to the horizontal plane since the na-
sofrontal hinge towards the tip in R. aguirrei and Sula spp.,
whereas R. ramirezi shows an upward angle until the mid-
length of the rostrum where it begins to descend. In R. agu-
irrei the tip of the rostrum is ventrally curved in almost 40%
of its maximum height from approximately its mid-length.
In dorsal view (Fig. 4A3), R. aguirrei as well as Sula and
Papasula exhibit a rough surface across its length. Ventrally
(Fig. 4A4), the rostrum has the typical schizognathous con-
dition of the family. Palatines are separated in their cranial
extremes and fused in the caudal extreme as in other sulids.
Neurocranium: The only noticeable difference between
R. aguirrei and R. ramirezi in the braincase is located at its
dorsal surface. From the nasofrontal hinge to the mid-length
of the orbitals the frontals of R. aguirrei are lower than those
of R. ramirezi with respect of the horizontal plane. Posterior
to the mid-length of orbitals, the braincase is slightly in-
clined upward in R. aguirrei whereas that of R. ramirezi
remains flat.
Remarks.—The morphology of Ramphastosula suggests
closer affinities to Sula than to Morus or Papasula. Both
Sula and Ramphastosula have a roughened maxilla sur-
face in its posterior-most part, and both have reduced and
bifurcated postorbital processes. Conversely, Morus has a
smooth maxilla surface, but shows the same condition of the
postorbital processes. Papasula shows long, sharp and ven-
trally-inclined postorbital processes but retains the rough-
ACTA PALAEONTOLOGICA POLONICA 61 (2), 2016
ened maxilla surface (Stucchi 2003; Stucchi and Urbina
2004). In this respect, the primitive condition of both char-
acters is present in Papasula (according to Smith 2010).
The skull of R. aguirrei has an intermediate morphol-
ogy between other sulids and R. ramirezi. Interestingly, the
rostrum of R. aguirrei is more similar to that of Sula rather
than R. ramirezi. Taking all this into account, along with the
previously observed morphological stability in the cranium of
sulids (Stucchi 2003), allows us to suggest that the specimens
from Poza Negra and Poza Roja represent different species.
Stratigraphic and geographic range.—Messinian, Pisco
depositional sequence, Pisco Formation, Southwestern Peru.
Concluding remarks
Ecomorphology.—The acute morphology of the rostrum
in some seabirds is suitable for hunting by plunge-diving.
Among these, boobies typify this technique, although others
such as the tropicbirds (Phaethontidae) and some species of
terns (Sterninae, Laridae) and pelicans (Pelecanidae) also
use this strategy (Nelson 1979; Field et al. 2011).
Sulids can perform plunge diving from high altitudes be-
cause they have (i) developed a cone-shaped rostrum filled
with trabecular bone material and a modified mandibu-
lar symphysis, which allow them to withstand the plunge
impact without damage (e.g., bending); and (ii) completed
obliteration of the beak’s nasal fossae, which displaces
the respiratory function to a space located below the pre-
orbital fenestra and jugal protected by the ramphotheca
(MacDonald 1960; Stucchi 2013). However, the fossil re-
cord found in the Pisco Formation has revealed some spe-
cies with different morphologies, as is the case of the tou-
can-booby Ramphastosula, which probably did not exhibit
the same plunge-diving technique, not from high altitudes at
least (Stucchi and Urbina 2004).
The upper jaw in tropicbirds and terns curves toward the
tip as in R. aguirrei. These birds also swoop but from lower
altitudes than sulids (slightly less than 10 m; Nelson 1979).
Because of morphological superficial resemblance, R. agu-
irrei likely could have plunged in a similar way. Despite
the limitations imposed by its larger size and weight to
achieving a considerable altitude, it is possible that having
more developed frontals and a flat cranial roof would give
them greater resistance and aerodynamics, and thus attain-
ing greater altitudes.
Interestingly, the occurrence in northern Peru of a fe-
male adult extant individual of S. nebouxii (see Stucchi
2013: fig. 5) with a rostrum morphology similar to that of
R. ramirezi suggests this particular beak shape do not hin-
der them from feeding similarly to their extant relatives.
Even though boobies can dive from altitudes up to 100 m,
they generally dive from lower heights (a few meters), and
reach less than a meter above the sea surface when they dive
diagonally (MS, personal observation). In contrast, peli-
STUCCHI ET AL.—MIOCENE SULID BIRDS FROM PERU
cans have their own plunge strategy as described Forbes
(1914) for Pelecanus thagus, as well as Field et al. (2011)
and Schreiber et al. (1975) for P. occidentalis, which are not
morphologically comparable with R. aguirrei.
Body size.—In the Eastern Pacific Ocean, Morus was the
predominant genus in the Northern Hemisphere (reported
in Mexico and California, USA), whereas Sula was in the
Southern Hemisphere (southern Peru and northern Chile) in
past communities. Stucchi (2003) has noticed the sympatric
occurrence of these species with disparate body sizes at
least during the Late Miocene. In California, there are three
big-sized gannets (M. magnus, M. stocktoni, and M. lompo-
canus) coexisting with the medium-sized M. media, and the
small-sized M. willetti. Likewise, we found in the Messinian
age the big-sized S. magna, the medium-sized M. peru-
vianus and S. aff. variegata, and the small-sized S. sulita
co-occurring in southern Peru. This study also provides
evidence that during the Tortonian age the medium-sized
S. brandi and the large-sized S. figueroae cohabited in this
southern area. In modern sulid communities, body size
plays an important role in segregating sympatric seabirds
with similar feeding strategies offering different solutions
to foraging challenges and thus facilitating coexistence (i.e.,
allowing them feeding on different prey size items and in
different areas), as exemplified by Duffy (1987), Jahncke et
al. (1997), Jahncke and Goya (2000) and Young et al. (2010)
in the case of congeneric species which live together but
with different body sizes. This also occurs even in sexually
dimorphic species, in which bigger individuals (females)
feeds on bigger and heavier prey (García-R. and López-
Victoria 2008). Of course, these differences do not prevent
them from terrestrial habitat competition (see Duffy 1984
and Figueroa 2004). Body size could thus be tentatively
used as ecomorphological proxy to indicate resource parti-
tioning in either northern or southern extinct assemblages.
Interestingly, differences in body size could be trace as far
back as the late Oligocene in the sulid community of the
Chandler Bridge Formation (South Carolina, USA) (Ksepka
2014), the oldest reliable sulids reported to date.
Sulid paleodiversity.—The fossil record of the genus Sula
ranges from the Early to Middle Miocene Chilcatay forma-
tion (Stucchi and DeVries 2003) to the present, representing
the largest timespan in this region. Since the Late Miocene
age, several species have evolved including the largest sulid
(S. magna), some large forms (Sula spp.), and even some re-
lated to modern species (S. aff. variegata) (Stucchi 2003). The
set assigned to Sula spp. is undoubtedly the most abundant,
occurring in all horizons. It is also one of the most complex
in terms of its systematics due to size, overlapping among
species such as S. tasmani or S. dactylatra and a paucity of
morphological characters (Stucchi 2003). We cannot rule out
the possibility that some material corresponds to more than
one species, to an undescribed species, to some inmature
material of S. magna, or even to Ramphastosula (only cranial
material has been recovered). Considering the morphological
425
affinities, the progressive change through time seen in the
skull of Sula and Ramphatosula, and the relative abundance
of Sula in layers under and overlying the Sacaco Sur level,
and the late and restricted appearance of Ramphastosula,
anagenetic evolution between these two genera is possible
(contra Benson and Erickson 2013). Additional material is
therefore necessary to clarify their affinities.
Biological drivers of seabirds diversity are difficult to
test, at least with the available data. Although the diver-
sity though time of marine primary producers such a dia-
toms have been used to explain the rise of top consumers
(see Marx and Uhen 2010 for cetaceans), such correlations
have several concerns, as diatom species richness is not
necessarily related with their abundance (Pyenson et al.
2010), because diversity or richness are not related to bio-
mass in temperate coastal upwelling ecosystems. In fact,
their trophic dynamics is controlled not from bottom-up
(i.e., diatoms) or top-down (i.e., seabirds, marine mammals)
but rather from the intermediate trophic level (i.e., pelagic,
planktivore fish) toward both sides (Cury et al. 2001; Bakun
2006). The Peruvian anchovy Engraulis ringens and the
sardine Sardinops sagax currently dominate the produc-
tivity in the Peru–Chile Current ecosystem, although they
are not ecological analogues as the anchovy prefer colder,
coastal waters than the more oceanic sardine (Bakun 2006;
Gutiérrez et al. 2007). Thus, it must be expected that similar
fish (see Lambert et al. 2015 for the Miocene occurrence of
the genus Sardinops in Peru) have driven the ecology in the
region at least since the onset of the Antarctic Circumpolar
Current (about 30 Ma; Scher et al. 2015), which mainly gen-
erated the conditions of upwelling (Marx and Uhen 2010).
Unfortunately, the fossil record of fish is largely unknown
in terms of both diversity and quantity in the southeastern
Pacific Ocean preventing any assessment. Even assuming a
dense fish fossil record, high species richness is commonly
found not in temperate but in tropical latitudes, which coin-
cides with the current sulid high species richness.
In similar manner, Stucchi (2003) has already pointed
out that the sulid diversity peak occurred during the Late
Miocene, but this observation could be misinterpreted.
The relative abundance of sulids in some Peruvian locali-
ties rather than others could be explained by preservation
bias due to different paleo-environments (see Marocco and
Muizon 1988; Stucchi and Figueroa 2011), but it could also
be affected by local conditions (e.g., volcanism). In short,
analyzing global-scale diversity patterns require taking into
account regional considerations of biotic and physical fac-
tors in an ecological context (Pyenson et al. 2014) to prop-
erly address biological questions.
Acknowledgments
We thank Walter Aguirre (MUSM) for materials preparation. Giovanni
Bianucci (Università di Pisa, Italy) funded the fieldtrip and preparation
of Sula brandi with partial support of a National Geographic Society
426
Committee on Research Exploration grant (9410-13). Leonard Brand
(Loma Linda University, Loma Linda, CA, USA) kindly shared his un-
published stratigraphic work on the Sacaco basin. We also thank editor
Stephen Brusatte (University of Edinburgh, UK), and reviewers Daniel
Ksepka (Bruce Museum, USA) and Daniel Field (Yale University, New
Haven, CT, USA) for their constructive comments on the manuscript.
Rodolfo Salas-Gismondi (MUSM) and Storrs Olson (USNM) allowed
access to specimens in their care. Luciano Stucchi (Universidad del
Pacífico, Lima, Peru), Manuel Plenge and Mauricio Bustamante (both
Lima, Peru) provided inaccesible references.
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 http://app.pan.pl/SOM/app61-Stucchi_etal_SOM.pdf

SUPPLEMENTARY ONLINE MATERIAL FOR

New Miocene sulids from Peru and considerations on their Neogene fossil record in the
Eastern Pacific Ocean

Marcelo Stucchi, Rafael M. Varas-Malca, and Mario Urbina-Schmitt

published in Acta Palaeontologica Polonica 61 (2): 417-427.

http://dx.doi.org/10.4202/app.00170.2015

Supplementary Online Material

SOM 1. Measurements (in mm) of the skull of sulid extant and extinct species. X: incomplete.

SOM 2. Measurements (in mm) of the postcranial elements of Sula figueroae sp. nov. MUSM 2501 (holotype) and
MUSM 2502 (paratype).

SOM 3. Measurements (in mm) of the postcranial elements of sulid extant and extinct species.

References
SOM 1. Measurements (in mm) of the skull of sulid extant and extinct species. X: incomplete. Sula variegata: n=30, Sula nebouxii: n=30, Sula dactylatra: n=8,
Sula sula: n=1, Sula leucogaster: n=2, Sula granti: n=4, Papasula abbotti: n=2, Morus bassanus: n=1, Morus capensis: n=2, Morus serrator: n=2. See details
in Stucchi (2013).
R. R. S. S. S. S. S. S. P. M. M. M.
Measurements S. dactylatra S. leucogaster
ramirezi aguirrei brandi figueroae variegata nebouxii sula granti abbotti bassanus capensis serrator
84.3 (91.4- 93.6 (100.1- 88.5 (88.9- 94.1 (97- 94.5 (96.3- 96.5 (98.1- 91.3 (94.3-
Rostrum length ~101 104.7 x 100.2 93.2 (97-90) 71.8 111.3
76.5) 84.5) 88.1) 92.4) 92.6) 94.8) 88.3)
58.7 (63.3- 62.3 (65.7- 66.6 (69.1- 59.3 (59.7- 64.8 (68.9- 67.6 (68.5- 70.8 (71.6- 67.7 (68.7-
Neurocranium length 68.3 75.6 60 68,4 54,3 77,1
56.1) 57.6) 64.7) 58.9) 61.8) 66.7) 70) 66.6)
26.9 (29.2- 27.5 (29.8- 28.7 (29.2- 26.1 (26.9- 28.2 (29.2- 32.8 (33.2- 34.4 (34.5- 31.9 (31.9-
Neurocranium height 33,6 33,7 23,1 27,5 26,8 34,5
25.7) 26.4) 27.6) 25.4) 27.6) 32.4) 34.3) 31.9)
Width at the level of 35.2 (37.8- 38.3 (40.1- 38.7 (40.4- 38.6 (40.7- 40.4 (41.1- 42.9 (43- 44.6 (45.3-
47,7 45,4 35,1 36,7 31,2 33,4 49,1
paroccipital processes 33.3) 34.8) 37.5) 37.3) 39.6) 42.8) 43.8)
Width at the level of the 25.7 (27.6- 27.3 (29.4- 30.8 (32.7- 27.5 (27.8- 29.1 (33.1- 29.7 (30.3- 28.8 (29.6-
~33.9 ~33.9 28,8 30,9 26,2 29,1 33,5
preorbital processes 23.8) 25.6) 29.7) 27.1) 26.7) 29.1) 28)
19.8 (21.2- 23.4 (24- 21.4 (21.9- 22.3 (25.1- 22.5 (22.7-
Nasofrontal hinge width 28,7 25 24,5 24 20.7 (21.7-19) 20,2 27,2 25,9 22.6 (23.1-2)
18.4) 22.2) 20.9) 20.5) 22.3)
15 (16.8- 15.8 (17.1- 17 (18.2- 14.4 (14.5- 16.7 (17.4- 19.2 (19.4- 17.3 (17.4- 18.7 (19.3-
Breadth at the exoccipitals 22,7 23,4 14,8 15,6 14,1 21,8
14.2) 14.2) 16.5) 14.2) 16.1) 18.9) 17.2) 18.1)
Breadth at the postorbital 41.7 (44.5- 42.9 (44.7- 45.1 (50.4-
50,2 ~51.7 x 48,6 x x 42.2 (42.3-41) x 56,5 x x
processes 39.9) 40.8) 41.8)
Postorbital process size 9,4 9,8 x 12 6.7 (8-5.3) 7.8 (9.6-6) x x 8 (8.9-7) 8.4 (9.6-7.4) x 11,2 x x
Occipital condyle width 5,5 7 4,7 5 4.4 (4.9-4) 4.8 (5.5-4) 5 (6.6-4.4) 3,7 4.4 (4.6-4.2) 5 (5.4-4.8) 5 (5-5) 6,7 5.1 (5.1-5) 5.6 (5.7-5.5)
Occipital condyle height 5 5,5 3,5 4 3.4 (3.7-3.2) 3.6 (4.2-3) 3.9 (4.4-3.8) 2,9 3.3 (3.4-3.1) 4 (4.1-3.8) 4 (4-3.9) 5,1 4.5 (4.9-4.1) 4 (4-4)
Foramen magnum width 10 10,1 8,3 6,3 9 (9.8-8.4) 9 (9.6-8.4) 9.6 (10-9.3) 8,2 8 (8.2-7.7) 9.4 (9.7-9.2) 9.4 (9.4-9.3) 10,2 9.9 (10-9.8) 9.5 (9.6-9.3)
10.4 (10.6-
Foramen magnum height 10 11,2 7 8,2 8.1 (8.9-7.1) 8.4 (9.2-7.7) 8.4 (8.6-8.2) 6,8 8.2 (8.4-7.9) 8.5 (8.7-8.1) 9.3 (9.3-9.2) 11,7 9.5 (9.5-9.4)
10.2)
13.3 (14.9- 15.5 (16.8- 14.5 (14.9- 15.6 (17.4- 16 (16.3- 14.1 (16.1- 14 (16.3-
Quadrate width x 19,6 x 17,8 14 (15-12.6) 14 19,2
12.5) 14.3) 14.1) 14.6) 15.6) 12.1) 11.7)
17.1 (18.7- 18.3 (19.4- 20 (21.1- 19.5 (20.7- 21.5 (21.6- 20.7 (20.9- 21.1 (21.5-
Quadrate height x 24,9 x 21,7 15,9 17.8 (18-17.5) 24
15.9) 16.4) 18.5) 18.9) 21.3) 20.4) 20.6)
21.6 (23.1- 21.3 (23.4- 22.2 (23.4- 21.9 (22.6- 27.5 (27.9- 25.9 (26.4-
Orbital fossa size 27,1 25,7 22,8 23,2 18,3 21 25 28,2
20.2) 19.4) 21.1) 21.2) 27.1) 25.4)
12.1 (12.6- 11.7 (12.7- 12.9 (13.9- 13.6 (15- 15.2 (16.2- 14.1 (14.9-
Fossa temporalis size 11,5 13 13 14,3 11 (12.5-9) 13 (14.6-11.6) 11 15,8
11.2) 10.7) 12.1) 12.2) 14.1) 13.2)
Mandible width at the 12.9 (14.4- 13.9 (14.9- 15.8 (17.1- 13.4 (13.5- 15.8 (17.4- 18.4 (18.6- 16.1 (16.3- 15.4 (15.7-
x x x 17,7 12,1 18,1
coronoid process 11.2) 12.8) 14.8) 13.3) 14.6) 18.1) 15.9) 15)

Breadth between the

30 (33.6- 34.4 (36.8- 30.5 (30.8- 34.3 (36.2- 33.5 (34- 34.9 (35.6- 35.3 (36-
coronoid proc. and the x x x 32,5 33.5 (35.5-29) 25,9 41,4
26.9) 33.2) 30.2) 33.6) 32.9) 34.2) 34.5)
medial mandibular proc.
Proportions
Rostrum length /
1,48 1,38 x 1,46 1,44 1,50 1,40 1,32 1,49 1,45 1,40 1,44 1,36 1,35
Neurocranium length
Rostrum length /
3,52 4,19 x 4,18 4,26 4,52 3,98 3,55 4,14 4,22 3,47 4,30 4,27 4,06
Nasofrontal hinge width
Neurocranium length /
2,03 2,24 2,60 2,49 2,18 2,27 2,32 2,03 2,27 2,30 2,06 2,23 2,06 2,12
Neurocranium height
Neurocranium length /
Paroccipital processes 1,43 1,67 1,71 1,87 1,67 1,63 1,72 1,74 1,78 1,68 1,67 1,57 1,65 1,52
width
Neurocranium length /
Breadth at the postorbital 1,36 1,46 x 1,41 1,41 1,45 x x 1,41 1,44 x 1,36 x x
processes
Neurocranium height /
Width at the level of 0,70 0,74 0,66 0,75 0,76 0,72 0,74 0,86 0,78 0,73 0,81 0,70 0,80 0,72
paroccipital processes

SOM 2. Measurements (in mm) of the postcranial elements of Sula figueroae sp. nov. MUSM 2501 (holotype) and MUSM 2502 (paratype).
MUSM 2501 (holotype) MUSM 2502 (paratype)
Left Right Left Right
Total length x x 186,5 185,6
Proximal width 30,9 x 29,3 28,4
Humerus
Distal width 24,9 24,9 23,8 24,3
Mid-shaft maximum width x 11,3 10,8 10,6
Proximal width 20,4 x x x
Carpometacarpus
Distal width 12,7 x x x
Proximal width x x x 15,2
Femur
Distal width x x x 16,8
Tibiotarsus Distal width x x x 14,2
Antero-posterior proximal width x x x 14,3
Tarsometatarsus
Latero-medial proximal width x x x 14,8
Total length x x 73,3 73,8
Sternal facet length x x 16,7 15,9
Coracoides Head length x x 14,4
Glenoid facet length x x 13,6 13,1
Distance head-glenoid facet x x 26,3 26,7
Total length x x 88,7 x
Scapula
Proximal width x 18,9 20,7 x
Total length x 109,3
Sternon
Breadth at the sternocoracoid processes x 63,3
Total length x 116,7
Pelvis
Breadth at the antitrochanteric processes 48,7 47
Ulna Proximal width 21,8 x x 16,2
SOM 3. Measurements (in mm) of the postcranial elements of sulid extant and extinct species. Fossil taxa measurements are shown in original descriptions unless those
indicated by a letter. A: Wetmore (1938); B: USNM 177935; C: USNM 23721; D: USNM 412010; E: Brodkorb (1955); F: USNM 412070; G: USNM 411963; H: USNM
426031; I: USNM 193364; J: USNM 252310; K: Grigorescu and Kessler (1977); L: USNM 178033; M: USNM: 501511; N: Howard (1978); O: USNM 215722; P: USNM
426057; Q: USNM 181029; R: USNM 215761; S: Miller (1935); T: Chandler (1990); U: USNM 501514; V: USNM 215716; W: USNM 411981; Y: Olson and Rassmussen
(2001); Z: USNM 412017; AA: USNM 426023; BB: USNM 426061; CC: USNM 177910; DD: USNM 412054; EE: USNM 367093. Sula variegata and S. nebouxii are after
Stucchi (2003); S. leucogaster and S. sula are the mean from Warheit (1992); S. dactylatra and S. tasmani from Tets et al. (1988); Sula indet. sensu Stucchi (2003). (*) Sula
indet. from Pisco Formation (Stucchi 2003).

Humerus Ulna Carpometacarpus Tarsometatarsus Femur Tibiotarsus Coracoid

Sulid species Mid-shaft
Proximal Total Proximal
Total length Distal width maximum Total length Total length Total length Total length Total length
width length width
width
Sarmatosula x
158 x x 127 x x 50 x 110 x
dobrogensis
Morus loxostylus x x 21.1A x x 13.1B 70.6C 55.6D x x 51.3/48.2E
x 12.2H /
Morus atlanticus 166.5F 22.5F 17.6G /17.5F 146.2F 71I / 69.6F 49F 57.6J / 50F 80.1F x
12.8F
Morus vagabundus 170K x 18,3 x x x x x x x x
Morus avitus x x 15.3L /15.2 x x 9.7M 75N /65 /56.3O 52.7P / 48.7Q 52.9R x x
Morus pygmaea 128K x x x x x x x x x x
Morus olsoni x x x x x x 70,2 x x x x
Morus willetti 156 x x x 147 x 70 41 52 71 45
Morus stocktoni 264 x x x 175 x 102 x x x x
x 201S / 96.6N / 95.7N /
Morus lompocanus 245 / 230S 28 x x x 69 x 62
107.6T 94.7N
Morus magnus x x 29,2 x x x 116,1 x x x x
Morus media 181S / 180 x x x 140 x 81 64 x x x
22V / 22W / x 64.2EE/
Morus peninsularis x 26.6U x 15.1Z 78.8AA 57.5BB 59.1CC x
21Y 59.8DD/55.6-54
Morus humeralis x 28T 19.8 /19.7T x x 14.7T x x 58.2T x x
Morus recentior x x x x 248.3T 18.2T x x x 114 64.7T
Morus reyanus x x x x x x x x x x 56,6
Morus peruvianus x x x x 165 14,8 76,8 x x x x
Sula pohli 150.5-145 22,1 17,3 x 170-148.5 x 69,3 x x x 55
209 11-9.0 212
Sula tasmani 30 23 (23-21) 17 (18-16) 92 64 (65-62) 68 (70-67) 100 75
(210-206) (216-208)
Sula magna 251.5-245 38.5-34.5 30.2-28 13.7 (14- 271 21.5-19.6 123.9-115.2 x x x 93-91
 13.4)
Sula sulita x x 15,8 x x x x x x x 45,7
Sula aff. variegata x x 20 x x 15.5-13 x x x x x
Sula indet. (*) 240-210 34.2-29.1 27.8-21.8 x 247-230 19.1-17 113.7-101.9 73.2-62.5 75-68.5 x 88.1-74.1
Sula figueroae 191-185 30.9 -28.4 25-24.4 11.3-10.6 x 16,2 92,9 58,8 72,8 98,1 73,3
193 9.1-7.8 204
Sula dactylatra 29 (30-28) 20 (21-19) 15 (15-14) 86 (90-85) 60 (63-57) 64 (66-61) 95 68 (73-67)
(202-186) (211-196)
Sula leucogaster 156,3 22,7 18,3 x 169,2 12,5 73,5 46,7 50,4 76,3 51,9
182.35 24.92 19.74 8.82
Sula nebouxii 210-172 14.5-12.1 85.8-70.9 57.6-49.8 56.7-49.6 93.4-79.9 55.5-47.7
(195.5-161.5) (26.3-22.5) (20.72-18.14) (9.9-7.8)
161.94 23.27 18.3 186.3-
Sula variegata 8.45 (9-7.7) 14.3-11.9 78.6-69.4 52.8-45.6 56.8-49.7 90.9-81.1 55.3-48.2
(175-161) (25.6-21.3) (19.8-17.2) 158.5
Sula sula 161,5 22,3 18,3 x 181,2 12,9 71,6 36,2 48 65,9 47,2

References
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Grigorescu, D. and Kessler, E. 1977. The middle Sarmatian avian fauna of south Dobrogea. Revue Roumaine de Géologie, Géophysique et Géographie 21: 93–102.
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