431
J. Paleont., 83(3), 2009, pp. 431?447
Copyright  2009, The Paleontological Society
0022-3360/09/0083-431$03.00
ODONATAN ENDOPHYTIC OVIPOSITION FROM THE EOCENE OF
PATAGONIA: THE ICHNOGENUS PALEOOVOIDUS AND IMPLICATIONS FOR
BEHAVIORAL STASIS
LAURA C. SARZETTI,1 CONRAD C. LABANDEIRA,2,3 JAVIER MUZO? N,4 PETER WILF,5 N. RUBE? N CU? NEO,1
KIRK R. JOHNSON,6 AND JORGE F. GENISE1
1CONICET, Museo Paleontolo?gico Egidio Feruglio, Avenida Fontana 140, Trelew, Chubut 9100, Argentina, lsarzetti@mef.org.ar,
rcuneo@mef.org.ar and jgenise@mef.org.ar; 2Department of Paleobiology, National Museum of Natural History,
Smithsonian Institution, 20213-7012; 3Department of Entomology, University of Maryland, College Park, Maryland 20742, labandec@si.edu;
4Instituto de Limnolog??a ??Dr. Raul A. Ringuelet,?? Av. Calchaqu?? Km 23,5 712, Florencio Varela, Buenos Aires, Argentina, 1888, muzon@ilpla.edu.ar;
5Department of Geosciences, Pennsylvania State University, University Park, Pennsylvania, 16802, pwilf@psu.edu; and 6Department of Earth Sciences,
Denver Museum of Nature and Science, Denver, Colorado 80205, kirk.johnson@dmns.org
ABSTRACT?We document evidence of endophytic oviposition on fossil compression/impression leaves from the early
Eocene Laguna del Hunco and middle Eocene R??o Pichileufu? floras of Patagonia, Argentina. Based on distinctive mor-
phologies and damage patterns of elongate, ovoid, lens-, or teardrop-shaped scars in the leaves, we assign this insect
damage to the ichnogenus Paleoovoidus, consisting of an existing ichnospecies, P. rectus, and two new ichnospecies, P.
arcuatum and P. bifurcatus. In P. rectus, the scars are characteristically arranged in linear rows along the midvein; in P.
bifurcatus, scars are distributed in double rows along the midvein and parallel to secondary veins; and in P. arcuatum,
scars are deployed in rectilinear and arcuate rows. In some cases, the narrow, angulate end of individual scars bear a
darkened region encompassing a circular hole or similar feature indicating ovipositor tissue penetration. A comparison
to the structure and surface pattern of modern ovipositional damage on dicotyledonous leaves suggests considerable
similarity to certain zygopteran Odonata. Specifically, members of the Lestidae probably produced P. rectus and P.
bifurcatus, whereas species of Coenagrionidae were responsible for P. arcuatum. Both Patagonian localities represent an
elevated diversity of potential fern, gymnosperm, and especially angiosperm hosts, the targets of all observed oviposition.
However, we did not detect targeting of particular plant families. Our results indicate behavioral stasis for the three
ovipositional patterns for at least 50 million years. Nevertheless, synonymy of these oviposition patterns with mid-
Mesozoic ichnospecies indicates older origins for these distinctive modes of oviposition.
INTRODUCTION
OVIPOSITION ON vegetation is a common insect behavior, inwhich eggs are deposited either on the surfaces of plant
organs (exophytic oviposition) or, alternatively, inserted within
plant tissues (endophytic oviposition; Corbet, 1999). Endophytic
oviposition represents an ancient behavior, based on external ovi-
positor structure found in some Paleozoic insects (Carpenter,
1971; Labandeira, 1998) and additionally on stereotyped tissue
damage occurring within foliage and stems (Labandeira, 2002b;
Be?thoux et al., 2004, Beattie, 2007). Externally evident, often
sawlike ovipositors have been recognized in the Palaeodictyop-
tera, Protodonata, Dictyoptera, Archaeorthoptera, Hemipteroidea,
and early-derived holometabolous clades from the Late Carbon-
iferous through the Triassic (Labandeira, 2006). External ovipos-
itors currently are most prominent in the orders Odonata, Orthop-
tera, Hemiptera, Coleoptera, Lepidoptera and Hymenoptera (Zeh
et al., 1989).
Worldwide, evidence for leaf damage produced by ovipositors
is relatively abundant throughout the Phanerozoic. Several Paleo-
zoic examples have been documented on sphenopsid stems from
the Late Pennsylvanian of France (Be?thoux et al., 2004) and the
Middle Permian of Germany (Roselt, 1954), and especially on
glossopterid leaves from the Late Permian of Australia (Beattie,
2007), South Africa (Prevec et al., 2009), India (Bunbury, 1861;
Banerji, 2004), and Brazil (Adami-Rodrigues et al., 2004). For
the Mesozoic, oviposition has been recorded for a variety of sphe-
nopsids and seed plants from the Middle and Late Triassic of
Germany and France (Kelber, 1988; Grauvogel-Stamm and Kel-
ber, 1996), Chile (Gnaedinger et al., 2007) and Austria (Pott et
al., 2007); the Early Jurassic of Germany (Van Konijnenburg-van
Cittert and Schmein?ner, 1999); and the early Late Cretaceous of
Israel (Krassilov et al., 2007) and Germany (Hellmund and Hell-
mund, 1996c). Cenozoic occurrences are known from the Eocene
for Washington State in the United States (Lewis and Carroll,
1991; Labandeira, 2002a, 2002b) and Spain (Pen?alver and Del-
clo`s, 2004), from the Oligocene of Germany (Hellmund and Hell-
mund, 1991, 1993, 1996a, 1996b, 1998, 2002b; Van Konijnen-
burg-van Cittert and Schemie?ner, 1999), and from the early and
late Miocene of Germany (Hellmund and Hellmund, 1996c,
2002a, 2002c).
Complex patterns of ovipositional plant damage frequently al-
low reliable identification of the producers, particularly when fos-
sil ovipositor structure and egg-laying biology of modern coun-
terparts are well studied. Accordingly, most post-Paleozoic fossil
occurrences of oviposition are confidently attributed to the Odon-
ata (Hellmund and Hellmund, 1991; Grauvogel-Stamm and Kel-
ber, 1996; Van Konijnenburg-van Cittert and Schemie?ner, 1999;
Labandeira, 2002a, 2002b; Adami-Rodrigues et al., 2004; Pen?al-
ver and Delclo`s, 2004; Gnaedinger et al., 2007), including those
described below.
Despite the extensive fossil evidence, as well as autecological
knowledge of modern ovipositing insects (Wesenberg-Lund,
1913b; Schiemenz, 1957; Corbet, 1999), formal ichnotaxonomic
treatments of these trace fossils were lacking until Vasilenko
(2005) established the ichnogenus Paleoovoidus, the first ichno-
taxon created for ovipositional damage to fossil plants. In a more
recent and extensive documentation, Krassilov and Silantieva
(2008) defined several new ichnogenera and ichnospecies to in-
clude oviposition scars and presumably arthropod eggs from the
Late Cretaceous (Turonian) Gerofit locality in Israel. Recently,
Vasilenko (2008) redefined their ichnogenus Paleoovoidus and
also described two new ichnospecies, P. flabellatus and P. arcu-
atus, assigning them to endophytic ovipositional traces. He ad-
ditionally defined a new ichnogenus, Palaexovoidus, to refer to
exophytic oviposition on plant tissues. Before the establishment
of these taxa, trace fossils of oviposition in leaves and stems were
described in the literature without formal ichnotaxonomic analy-
ses (e.g., Hellmund and Hellmund, 1998; Labandeira, 2002a), in-
cluding the damage-type (DT) system of Labandeira et al. (2007),
432 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
FIGURE 1?Map of central Patagonia showing 50 Ma positions of both
localities under study, with modern coastlines: Laguna del Hunco (Chubut
Province) and R??o Pichileufu? (R??o Negro Province) (redrawn from Wilf et al.,
2005).
used for analyzing the paleoecology of plant-insect associational
patterns. Additionally, erection of ichnotaxa for ovipositional
trace fossils was not considered in several more encompassing
studies (Van Amerom, 1966; Straus, 1977; Givulescu, 1984; Vas-
ilenko, 2006, 2007), whereas in most of those same works, nu-
merous ichnotaxa were established for various other types of fo-
liar damage.
This contribution has three principal objectives. The first is to
formally define new ichnospecies and to provide additional evi-
dence for previously defined ovipositional damage from two di-
verse Eocene Patagonian paleofloras. Establishment of these new
ichnotaxa will be done by comparison with analogous oviposi-
tional damage by modern Odonata. The second is to ascertain
whether the ichnotaxa from these Eocene paleofloras have other
occurrences in the fossil record, providing a brief review of the
relevant fossil record of endophytic oviposition in leaves. The
third is to track long-term ovipositional behavior of odonatan lin-
eages inflicted on leaves about 50 million years ago, comparing
these oviposition patterns with earlier occurrences of the ichno-
taxa.
GEOLOGICAL SETTING
Leaves bearing oviposition scars analyzed in this contribution
originate from the Laguna del Hunco (early Eocene, 51.91  0.22
Ma) and R??o Pichileufu? (middle Eocene, 47.46  0.05 Ma) sites
in central Patagonia (Fig. 1; Wilf et al., 2005a). These two classic
paleobotanical localities (Berry, 1925, 1928, 1935a, 1935b,
1935c; Frenguelli, 1943a, 1943b; Romero and Hickey, 1976; Gan-
dolfo et al., 1988), exhibit very diverse paleofloras (Wilf et al.,
2005a) and associated spectra of insect-mediated damage (Wilf et
al., 2005b), including oviposition, which was mentioned but not
analyzed by Wilf et al. (2005b).
The Laguna del Hunco (LH) paleoflora comes from a caldera-
lake deposit, the Tufolitas Laguna del Hunco (Arago?n and Maz-
zoni, 1997; Arago?n et al., 2001), and is located in western Chubut
Province, Argentina, 160 km southeast from the R??o Pichileufu?
(RP) locality, along the foothills of the Sierra de Huancache at
S42.5, W70.0 degrees (Fig. 1). The fossil material originates from
strata consisting of tuffaceous mudstones and sandstones, inter-
preted as a caldera-collapse lake unit within the expansive middle
Chubut River volcanic-pyroclastic complex (Arago?n and Romero,
1984; Arago?n and Mazzoni, 1997). The entombed plant remains
are preserved principally as impressions; angiosperm leaf impres-
sions are the most abundant plant organs, though there are also
significant gymnosperm and minor fern components (Wilf et al.,
2005a).
Recently, 25 distinct fossil plant localities (known as LH1-25)
were intensively collected and placed in a 170 m measured strati-
graphic section, from which six paleomagnetic reversals were de-
tected, and three tuffs interbedded with the fossils were dated
using the 40Ar-39Ar method (Wilf et al., 2003). All Ar-Ar ages
were near 52 Ma; and one tuff containing sanidine was reana-
lyzed, yielding an age of 51.91  0.22 Ma (Wilf et al., 2005a).
The combined evidence placed the LH flora within the sustained
period of globally warm temperatures known as the early Eocene
climatic optimum (Zachos et al., 2001). Leaf-margin and leaf-area
analyses indicate local mean-annual paleotemperatures near 16.6
C and annual rainfall of at least 1140 mm, though certain plant
taxa suggest much higher rainfall (Wilf et al., 2005a, 2008). A
number of botanical investigations have emerged from the recent
collecting effort, which has yielded 210 species, including stud-
ies of Myrtaceae (Gandolfo et al., 2006), Casuarinaceae (Zamaloa
et al., 2006), Proteaceae (Gonza?lez et al., 2007), and many other
groups (Wilf et al., 2007).
Several entomological studies have revealed the presence of a
diverse insect fauna, including representatives of the orders Odon-
ata, Blattodea, Orthoptera, Hemiptera, Coleoptera, Mecoptera,
Diptera, Hymenoptera and Trichoptera (Rossi de Garc??a, 1983;
Petrulevic?ius and Martins-Neto, 2000; Genise and Petrulevic?ius,
2001; Petrulevic?ius and Nel, 2003a; Petrulevic?ius, 2005). Addi-
tionally, pipoid anurans were described by Casamiquela (1961)
and Ba?ez and Trueb (1997) and a catfish species by Sa?ez (1941).
Nevertheless, the preserved fauna is less abundant and diverse
than the flora of this locality, though insect damage to fossil
leaves is abundant and extremely diverse (Wilf et al., 2005b).
The R??o Pichileufu? (RP) paleoflora originates from a probable
caldera-lake facies of the Ventana Formation (Arago?n and Ro-
mero, 1984) and is located in R??o Negro Province of Argentina,
approximately 40 km east from the city of San Carlos de Bari-
loche at S41.2, W70.8 degrees (Fig. 1). The RP flora was the
subject of an early monograph (Berry, 1938) that remains the
most extensive on Cenozoic macrofloras in South America,
though nearly all taxa are in need of revision. More recently, the
flora was extensively recollected from three quarries (known as
RP1-3), and tuffs immediately associated with the fossil floras
produced an Ar-Ar age of 47.46  0.05 Ma (Wilf et al., 2005a).
There is much less fossiliferous exposure at RP than at LH, and
preservation is not as good, but the floral diversity at RP3 is
similar to that of the richest LH quarries (Wilf et al., 2005a). Like
LH, the RP flora is dominated by angiosperm leaves but has sig-
nificant contributions from gymnosperms such as Araucaria, Po-
docarpaceae, and Ginkgo, as well as fern species. Early studies
of this locality (Berry, 1935a, 1935b, 1938) revealed a great di-
versity of fossil plant taxa that are only recently being taxonom-
ically updated. Based on preliminary leaf-margin analysis, the RP
mean annual paleotemperature is 19.2  2.4 C (Wilf et al.,
2005a).
Fossil frogs and insects, including large ants, have been noted
433SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
FIGURE 2?1, Paleoovoidus bifurcatus, isp. nov., holotype specimen (MPEF-IC-1385) on a sapindaceous leaf (LH13, morphotype indeterminate, MPEF-
Pb-1607), scale bar: 0. 5 cm; 2, specimen MPEF-IC-1385 enlarged from boxed area in A of Paleoovoidus bifurcatus, scale bar: 1 mm; 3, Paleoovoidus rectus
specimen (MPEF-IC-1376) on ??Myrcia?? chubutensis (LH13, morphotype TY21, MPEF-Pb-2216), enlarged from boxed area in D, scale bar: 0. 5 cm; 4,
specimen MPEF-IC showing P. rectus (boxed area) and P. arcuatum below (MPEF-IC 1392).
from the plant-bearing beds (Wilf et al., 2005a), although Archi-
myrmex piatnitzkyi, reported from the general area (Viana and
Haedo-Rossi, 1957; Dlussky and Perfilieva, 2003), can not be
correlated stratigraphically to the flora as suggested by those au-
thors, due to local faulting (Wilf and Johnson, personal observa-
tion, 2008). Also leaf-cutter bee (Megachilidae) foliar damage
was reported from RP (Sarzetti et al., 2008). Interestingly, Berry
(1938) figured an extensively insect-damaged leaf from RP he
named ??Anona?? infestans, one of the earliest explicit illustrations
of fossil insect damage from South America. Berry (1938, p.73)
reported that the damage ??might be the remains of scale insects.??
Our updated interpretation of this damage as oviposition scars
appears below.
SYSTEMATIC ICHNOLOGY
The specimens described here are deposited in the Museo Pa-
leontolo?gico Egidio Feruglio, Trelew, Chubut, under the paleo-
botanical collections (MPEF-Pb, for the leaf holding the trace),
with each trace assigned to a separate suite of ichnological col-
lection numbers (MPEF-IC).
Ichnogenus PALEOOVOIDUS Vasilenko, 2005
Figure 2
Paleoovoidus VASILENKO, 2005, p. 630, figs. 1?3.
Sertoveon KRASSILOV, 2008, partim, p. 69, figs. 1?5.
Paleoovoidus VASILENKO, 2008, p. 516, fig. 2, pl. 7.
434 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
435SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
FIGURE 4?Camera lucida drawing of a specimen of Paleoovoidus arcuatum, isp. nov. (MPEF-IC-1370), on the dicot ??Celtis?? ameghenoi (LH2, morphotype
?TY20, MPEF-Pb-1053), scale bar: 1 cm: 1, entire specimen, show a register of five rows of oviposition scars oriented across the blade from upper-left to
the center, and a differently juxtaposed register of perhaps five rows at the upper-right region of the leaf; 2, minor enlargement of six scars with central
ovoidal depressions from a circled region of the leaf margin at upper-left; 3, major enlargement of callus and emergence hole detail from a single scar,
indicated by a circular template from the leaf margin at center-left; 4, similar enlargement of three scars with central emergence holes, from the circled leaf
region at center-bottom, axial length of leaf: 4.5 cm, scale bar as in Fig. 3.5.
?
FIGURE 3?Paleoovoidus arcuatum specimens: 1, MPEF-IC-1380a, MPEF-IC-1380b and MPEF-IC-1380c specimens on an unknown dicot leaf (LH13,
morphotype indeterminate, MPEF-Pb-1591), showing a zigzag pattern, lettered arrows point to individual rows or ??files?? of oviposition marks; 2, MPEF-IC-
1371a-c showing consecutive and parallel rows on an indeterminate dicot leaf (LH6, morphotype indeterminate, MPEF-Pb-1054), scale bar: 1 cm; 3, linear
oviposition arcs (lettered arrows) covering MPEF-IC-1374a and specimens MPEF-IC-1374b, c on ??Cupania?? latifolioides (LH18, ?Cunoniaceae, MPEF-Pb-
1585), scale bar: 1 cm; 4, trace-fossil specimens MPEF-IC-1388 a-e (lettered lines) on an indeterminate dicot leaf (RP3, morphotype indeterminate, MPEF-
Pb-1611), scale bar: 1 cm; 5, specimens MPEF-IC-1370 a-g (lettered lines) on the dicot ??Celtis?? ameghenoi (LH2, morphotype TY20, MPEF-Pb 1053), scale
bar: 1 cm.
436 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
437SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
?
FIGURE 5?Fossil and extant specimens of Paleoovoidus arcuatum isp. nov: 1, specimen MPEF-IC-1390 a-d (lettered lines) in an indeterminate dicot leaf
(RP3, morphotype indeterminate, MPEF-Pb-1612) with a consecutive and parallel pattern of rows, scale bar: 0. 5 cm; 2, extant oviposition of Acantagrion
oblutum (Coenagrionidae) on Cynoglossum amabile (Boraginaceae) showing a zigzag pattern and probably the immature aperture observed as a dark lineal
in the centre of the scar; 3, two individual eggs and associated callus tissue of Acantagrion oblutum, scale bar: 0.5 cm; 4, specimens MPEF-IC 1393b-d
(lettered lines) on an indeterminate dicot (RP3, morphotype indeterminate, MPEF-Pb-2407); scale bar: 0.5 cm; 5, specimens MPEF-IC-1368a-e (lettered lines)
showing a distinct zigzag pattern on ??Myrcia?? deltoidea (LH4, Myrtaceae, MPEF-Pb-1052), scale bar: 0.5 cm; 6, specimen MPEF-IC-1389d, a scar with a
circular breached area probably representing the emergence portion of the naiad (arrow) on Lomatia occidentalis (LH2, morphotype TY44, MPEF-Pb-988),
scale bar: 1 mm.
??Flea beetle egg deposition,?? LEWIS AND CARROLL, 1991, p. 335,
fig. 2.
??Flea beetle egg deposition,?? LEWIS AND CARROLL, 1992, p. 3,
fig. 2.
Emended diagnosis.?Medium-sized elongate, narrow, ovoid,
or lens-shaped structures, characterized by regular arrangement in
leaf lamina. These structures, defined by dark, surrounding reac-
tion tissue, are narrow at one end, each of which often bears a
dark spot.
Discussion.?Vasilenko (2005, p. 629) erected the ichnogenus
Paleoovoidus originally as ??oval or lentiform structures (eggs)
with regular distribution over substrate.?? More recently, Vasilen-
ko (2008) redefined the ichnogenus, providing a more precise
diagnosis. Nevertheless, the diagnosis is emended here because
we consider that the term ??substrate?? should be restricted to
leaves, which probably was the original intention of the author.
In addition, the mention of ??eggs?? in the diagnosis is dispensed
with herein, as there is no indication for the presence of eggs,
thus rendering moot the author?s inference. Alternatively, Kras-
silov and Silantieva (2008) defined the ichnogenus Sertoveon to
correspond with a structure and distribution over the leaf lamina
very similar to our material. However, according to the current
rules of zoological nomenclature, the valid name should be Pa-
leoovoidus because of date priority over Sertoveon. Other ich-
nogenera erected by Krassilov and Silantieva (2008), Costoveon
and Catenoveon, are comparable with Paleoovoidus, although the
disposition of the scars over the leaf are different. In Costoveon,
all scars are distributed over primary veins. By contrast, Caten-
oveon is similar to Paleoovoidus in that the scars are oriented in
the direction of stronger veins, but the sets of scars cover the
entire surface of the lamina.
PALEOOVOIDUS RECTUS Vasilenko, 2005
Figure 2.2, 2.4
??Odonata eggs?? VAN KONIJNENBURG-VAN CITTERT AND
SCHMEI?NER, 1999, p. 217.
??Egg scars?? KRASSILOV, SILANTIEVA, HELLMUND, AND HELL-
MUND, 2007, p. 806, fig. 3D.
Emended diagnosis.?Elongate to lens-shaped scars oriented in
a single, linear row, with long axes of scars aligned lengthwise,
mostly parallel to the long axis of the leaf and usually occurring
along the midrib.
Description.?The specimen of Paleoovoidus rectus (Fig. 2.3)
occurs in a leaf of ??Myrcia?? chubutensis (Myrtaceae, MPEF-Pb
2216). This leaf has two sets of leaf scars; those corresponding
to C. rectus are indicated by the inset box in Figure. 2.4. There
are twelve scars arranged rectilinearly near the leaf apex, aligned
approximately along the primary vein, with the scar long axis
parallel or subparallel to the vein. Two scars are on the primary
vein, whereas the other ten are closely adjacent to it in a single
row along the intersection of the laminar and veinal tissue, except
for the two closest scars at the tip of the leaf that appear on the
opposite side of the primary vein. The individual length of the
scars ranges from 1.0 to 1.2 mm, and the width ranges from 0.4
to 0.5 mm. The distance between adjacent scars varies from 0.7
to 1.4 mm. All scars have an ovoidal shape and rounded ends,
and they lack the dark spot observed in other specimens. In ad-
dition, we note that in this leaf a distinctive occurrence of Pa-
leoovoidus arcuatum (MPEF-IC 1392) (Fig. 2.3) appears basal to
the P. rectus trace, indicating that both patterns can occur on the
same leaf.
The P. rectus holotype described by Vasilenko (2005) exhibits
minor differences from the material presented here. Vasilenko?s
(2005) oviposition damage has a similar arrangement, orientation,
and measurements compared with the LH specimen, but the scars
are distributed in two linear rows adjacent to the central vein in
Pityophyllum sp. leaves (Coniferales: Pinaceae), and in up to four
subparallel rows on Ginkgoites (Ginkgoales: Ginkgoaceae), which
lack a midvein. None of these scars occurs within veins. Our LH
specimen also resembles the Lower Jurassic scar pattern docu-
mented by Van Konijnenburg-van Cittert and Schmei?ner (1999)
in leaves of Schmeissneria microstachys (Ginkgoales: Schmeis-
sneriaceae) and Podozamites distans (Coniferales: Podocarpa-
ceae); both species lack midveins. These authors described two
types of row arrangement, but in both cases, the scars are slightly
longer than the Patagonian material, with major-axis values ap-
proximating 3 mm. These multiple rows provide the most impor-
tant difference from the pattern described here, where scars are
distributed along a single row.
Material examined.?One specimen (MPEF-IC-1376) occurs
on a ??Myrcia?? chubutensis (MPEF-Pb-2216, from locality LH13
of Wilf et al., 2003), from the early Eocene Tufolitas Laguna del
Hunco, Chubut Province, Argentina (Fig. 2.3).
Discussion.?Vasilenko (2005) defined this ichnospecies as
??oval or lentiform relief structures, individual eggs oriented in
chain on leaf blades.?? The diagnosis is emended here to avoid
the use of an interpretative concept, such as ??eggs?? and to pro-
vide a more accurate description of the oviposition pattern. How-
ever, Pott et al. (2008) have described Late Triassic oviposition
scars from Austria with preserved ovoidal egg cuticles in ben-
nettitalean leaves, arranged in a near-circular pattern but lacking
the linearity of P. rectus. These early Mesozoic ichnospecies can
be distinguished from the ichnospecies P. rectus because of the
arrangement of scars in a single row. Krassilov and Silantieva
(2008) defined a similar ichnospecies, Catenoveon undulatum.
However, their definition included ??elliptical to fusiform egg
pits?? (page 68) distributed nearly parallel or oblique to veins but
distributed over most of the leaf. The principal difference between
C. undulatum and P. rectus is that oviposition scars cover most
of the laminar surface, whereas in P. rectus a single linear row
is present along the primary vein. We consider Vasilenkos ich-
nospecies Paleoovoidus rectus (2005) more germane for under-
standing the pattern in the Patagonian specimen. Other similar
evidence of P. rectus comes from the Mesozoic (Van Konijnen-
burg-van Cittert, 1999; Vasilenko, 2005). These occurrences ap-
pear to be associated with the great diversity of parallel-veined,
broadleaved gymnosperms from floras of Late Triassic to Early
Cretaceous age. Older documentation comes from Late Permian
floras, typically occurring in or adjacent to the robust midribs of
Glossopteris leaves. Paleozoic occurrences were probably pro-
duced by protodonatan dragonflies and related paleodictyopteroid
lineages (Labandeira, 2006).
438 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
PALEOOVOIDUS BIFURCATUS new ichnospecies
Figure 2.1, 2.2
??Galle Aceria nervesqua fagina?? STRAUS, 1977, p. 74, fig. 2, p.
78, fig. 50.
??Lestiden-Typ?? HELLMUND AND HELLMUND, 1991, p. 4, figs. 1,
2; HELLMUND AND HELLMUND, 1996c, p. 160, fig. 16; HELL-
MUND AND HELLMUND, 2002b, p. 49, figs. 1, 2.
??Oviposition damage on secondary veins?? LABANDEIRA, WILF,
JOHNSON, AND MARSH, 2007, p. 10.
Diagnosis.?Elongate to lens-shaped scars arranged in pairs
along both sides of a primary vein, forming double rows and
sometimes a V-shaped configuration, with the arms of the V par-
allel to secondary veins and the vertex embedded in the midvein.
Oviposition scars may continue as a single row of scars located
in an acute angle along and parallel to one side of the vein. A
dark spot at one extremity of an individual scar very frequently
touches the primary vein.
Description.?The holotype occurs in a leaflet of ??Schmidelia??
proedulis, MPEF-Pb-1607. This specimen shows an upper pair of
scars arranged symmetrically about the midvein and parallel to
secondary veins characterized by a V-shaped configuration along
the primary vein. Another pair of scars occurs with a similar V
configuration, but their arrangement is less clear than for the up-
per pair. Two additional isolated scars occur at one side of the
vein in a more basal position with respect to the preceding two
pairs. The axial length of the scars ranges from 1.7 mm to 2.6
mm, and their width ranges from 0.5 mm to 0.7 mm. The pres-
ervation of the specimen is poor, and individual scars are only
partially preserved. The first scar occurs at the upper left side of
the lamina and is the best preserved; it has a length of 2.6 mm
that may indicate that the remaining scars were probably longer
than the values measured. The minimum distance between the
scars of the first pair is 0.8 mm and for the second pair is 1.3
mm. The longitudinal distance between two adjacent pairs ranges
from 5.5 mm to 10.7 mm. The greatest distance occurs between
the second pair of scars and the first scar of the single row.
The scars often occur in pairs or are isolated and oriented at
an acute angle with respect to the vein; their prominence is at-
tributable to the thickened scar tissue surrounding and probably
elicited by the inclined, secondary veins. Paired scars do not occur
at the same level on each side of opposite veins, or at the same
transverse position in the case of alternate veined leaves. MPEF-
IC 1385 (Fig. 2.2) is an example in which the upper pair of scars
is characterized by the left scar being slightly nearer the tip of
the leaf than the right one; by contrast, the lower pair shows the
opposite arrangement.
The specimens described by Hellmund and Hellmund (1991,
2002b) on Cinnamomum sp. (??Daphnogene??; Rott locality, Ger-
many, late Oligocene) and on a fragment of an unidentified leaf
(Geiseltal locality, Germany, middle Eocene) show a scarring ar-
rangement similar to that described above. In both examples, dou-
ble rows and single rows were found; in several instances, these
patterns appeared on the same leaf. In all cases for which a single
row was present, there was the continuation of a double row.
However, the double rows show some differences from the Pa-
tagonian specimen. These European specimens were composed of
12 to 16 scars that are separated for short longitudinal distances
between 0.1 to 1 mm, some of which are oriented 90 with respect
to the vein. Moreover, the average length, 1.20 mm, is shorter
than that of the LH specimen.
Etymology.?From the Latin noun, bifurcatus, the condition of
having two prongs or forks; masculine.
Types.?Holotype specimen MPEF-IC-1385 on the leaflet spec-
imen of ??Schmidelia?? proedulis (Sapindaceae) MPEF-Pb-1607
(Fig. 2.1, 2.2).
Examined material.?The same as the holotype.
Occurrence.?From the early Eocene Tufolitas Laguna del
Hunco, Chubut Province, Argentina, locality LH13.
Discussion.?This ichnospecies is comparable with Paleoovo-
idus rectus, as in both cases scars are located in relation to the
primary veins. However, in P. rectus there is a single row of scars,
their long axes approximately parallel with the long axis of the
leaf and consequently with the midrib or other major vein. The
combination of scar structure and position on the leaf separates
P. bifurcatus from P. rectus and P. arcuatum. This distinctive
ovipositional pattern corresponds to the damage type DT102 of
Labandeira et al. (2007). Patterns similar to P. bifurcatus de-
scribed here occur throughout the Cenozoic (Hellmund and Hell-
mund, 1991, 1996c, 2002b); this damage has a sporadic presence
on Mesozoic gymnospermous vegetation (Labandeira, pers. ob-
servation.), but is common on Glossopteris leaves from certain
Late Permian floras of South Africa (Prevec et al., 2009). Pres-
ervation of P. bifurcatus is variable across a broad range of com-
pression/impression floras, but all occurrences have the signature
ovoidal, ellipsoidal, or teardrop-shaped oviposition scars oriented
along or within the secondary venation and can occur at variable
distance from the midrib.
PALEOOVOIDUS ARCUATUM (Krassilov, 2008)
Figures 3, 4, 5.1, 5.4?5.6, 6, 7
??Coenagrioniden-Typ?? HELLMUND AND HELLMUND, 1991, p. 7,
fig. 3; HELLMUND AND HELLMUND, 1993, p. 349, fig. 1, p. 350,
fig. 2; HELLMUND AND HELLMUND, 1996a, p.109, fig. 1a, b;
HELLMUND AND HELLMUND, 2002a, p. 255, fig. 1a.
??Tra?ufelspitze?? HELLMUND AND HELLMUND, 1991, p. 291, pl. 1,
fig. 2.
??Coenagrioniden-Typ vom Bogenmodus?? HELLMUND AND
HELLMUND, 1998, p. 282, fig. 1.
??Concentric oviposition tracks?? LABANDEIRA, 2002a, p. 41.
??Radially oriented oviposition scars?? LABANDEIRA, JOHNSON,
AND LANG, 2002, p. 312, fig. 8o.
??Ovoposiciones de la Familia Coenagrionidae?? PEN? ALVER AND
DELCLO` S, 2004, p. 74, fig. 2.
??Zygopteran egg sets?? KRASSILOV, SILANTIEVA, HELLMUND,
AND HELLMUD, 2007, p. 806, fig. 3a, b, c.
??Endophytic oviposition probably of Calopterygina?? VASILENKO
AND RASNITSYN, 2007, p. 1156, figs. 4?6.
Sertoveon arcuatum KRASSILOV, 2008, p. 69, fig. 5.
Paleoovoidus arcuatus, VASILENKO, 2008 (new syn.), p. 516, fig.
2c, pl. 7, figs. 2, 3.
Emended diagnosis.?Elongate, lens-shaped to teardrop-shaped
scars arranged with the short axes aligned horizontally to each
other, either as straight rows or as arcs. Frequently the long axes
of scars are subparallel to each other. Occasionally, successive
rows are parallel or exhibit zigzag patterns.
Description.?The shape of the scars ranges from elongate
(Fig. 5.5), lens-shaped (Figs. 4.1, 4.2, 5.1, 6.5), to teardrop-shaped
(Figs. 4.1, 4.3, 4.4, 5.4). The specimens are from 0.9 to 1.6 mm
in length but only two specimens represent the highest values
(Fig. 3.5); widths range from 0.4 to 0.6 mm. The scars in the
rows are variously oriented within the leaves, with the long axis
parallel to subparallel to the primary vein (Figs. 3.4, 5.1, 5.4?
5.6), perpendicular to it (Figs. 3.1, 3.3), at a highly inclined angle
(Fig. 3.3), or at other inclinations (Fig. 6.1?6.3). The distance
between each scar within a row often is a relatively constant value
of 1 mm, but in some cases (e.g., Figs. 3.5, 4.1) the scars are
variably separated, with values that range from 0.4 to 2 mm (Fig.
6.3?6.5). Some leaves present distinct and well-defined arcuate
rows from three to 13 scars (Figs. 3.1, 3.2, 4.1, 5.1, 5.6, 6.1?6.3,
and 7.2?7.5), in which the acute end of the scar always is ori-
entated in the same direction (Figs. 3.1, 4.4, 5.4). In other cases
the arcuate rows are parallel to each other and are arranged en
echelon (Figs. 3.4, 3.5, 4.1, 5.1, 5.5, 6.2), with scars of each
439SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
FIGURE 6?Specimen of Paleoovoidus arcuatum on the holotype of ??Anona?? infestans from R??o Pichileufu? (Berry, 1938) (morphotype unknown, type
material, USNM 40389a,b): 1, entire leaf fossil showing the distributions of scars over the lamina, scale: 1 cm; 2, specimens USNM 40389a-b (lettered lines)
6.3?6.5, details of individual scars, scales: 1 cm (Fig. 6.1, 6.2), 1 mm (Fig. 6.3, 6.4), and 0.1 mm (Fig. 6.5).
440 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
FIGURE 7?Additional specimens of Paleoovoidus arcuatum isp. nov. exhibiting an overlapping distribution of oviposition scars (Fig. 7.1, 7.2, 7.4) and
clearly demarcated rows (Fig. 7.3, 7.5): 1, MPEF-IC-1367 in an unknown dicot leaf (LH4, morphotype TY122, MPEF-Pb-1051) [note continuous linear file
of oviposition marks at bottom of leaf (arrow)], scale bar: 1 cm; 2, MPEF-IC-1391 in an unknown dicot leaf (RP2, morphotype indeterminate, MPEF-Pb-
1613); 3, MPEF-IC-1375 in an unknown dicot leaf (LH2, morphotype indeterminate, MPEF-Pb-1588), scale bar: 1 cm; 4, enlargement from rectangular
template in Figure. 7.1, showing detail of oviposition scars exhibiting dark spots (arrows) at the narrow end of each ovoidal scar; 5, MPEF-IC-1377 on
??Cassia?? argentinensis (LH4, morphotype TY117, MPEF-Pb-1589); scale bar: 0.5 cm.
consecutive row arranged regularly with minimal deviation of
scars within a row. These rows may be subparallel to each other
(Figs. 4.1, 6.2) or convergent in one or more points that result
from projected lines drawn perpendicular to the multiply oriented
oviposition rows (Fig. 3.4) (Hellmund and Hellmund, 1991, fig.
6). In other cases (Figs. 3.1, 5.5) the arcuate rows are not parallel
to each other, the terminus of one approaching the terminus of
the next one, resulting in a zigzag pattern. In some specimens the
leaf tissue between two adjacent scars is broken or removed (Fig.
5.5). In other specimens (Figs. 3.2, 4.3, 4.4, 5.6, 6.4), the blunt
end of the scars displays a rounded hole, indicating the absence
of plant tissue, resulting in a whitish area. In these same speci-
mens, and in others (Figs. 5.6, 6.4), a dark spot at one end is
observed, which is most clearly distinguished at the acute ends
of teardrop-shaped scars (Figs. 3.5, 4.1).
Chaotic ovipositional patterns on leaves as in Figure. 7.1 may
441SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
result in uninterpretable behavior. However, closer examination
reveals that rows of distinctive oviposition scars have been over-
printed (Fig. 7.1, 7.4), with individual scars having an elongate-
to lens-shaped structure with an enveloping raised rim and a cen-
tral depression, often housing a dark circular region at one an-
gulated end (Fig. 7.4). There is discernable alignment of scars
into particular rows and arcs. Targeted host plants typically are
robustly constructed pinnate leaves with prominent midveins.
Material examined.?Specimen MPEF-IC 1367, on an unaffil-
iated dicot (LH4-1213, morphotype TY122 of Wilf et al., 2005a),
specimen MPEF-IC-1368a?e, on ??Myrcia?? deltoides, (LH4, Myr-
taceae, MPEF-Pb-1052); specimen MPEF-IC-1369, on ??Myrcia??
chubutensis, (LH13, Myrtaceae, MPEF-Pb-2217); specimen
MPEF-IC-1370a?f, on ??Celtis?? ameghenoi (LH2, MPEF-Pb-
1053); specimen MPEF-IC-1371a?c, on an unaffiliated dicot
(LH6, MPEF-Pb-1054, morphotype indeterminate); specimen
MPEF-IC-1372a?e, on an unaffiliated dicot (LH2, MPEF-Pb-
1584); specimen MPEF-IC-1373a?f, on ??Myrcia?? chubutensis
(LH13, Myrtaceae, MPEF-Pb-2215); specimen MPEF-IC-
1374a?c, on ??Cupania latifolioides?? (LH18, ?Cunoniaceae,
MPEF-Pb-1585); specimen MPEF-IC-1375, in an unaffiliated di-
cot (LH2, morphotype indeterminate, MPEF-Pb-1588); specimen
MPEF-IC-1377, on ??Cassia?? argentinensis (LH4, morphotype
TY117, MPEF-Pb-1589); specimen MPEF-IC-1378a?e, on Lo-
matia occidentalis, (LH4, Proteaceae, MPEF-Pb-998); specimen
MPEF-IC 1380a?c, on an unaffiliated dicot leaf (LH13, MPEF-
Pb 1591); specimen MPEF-IC-1381, on Malvaceae (LH6, mor-
photype TY23, MPEF-Pb-1592); specimen MPEF-IC-1383, on an
unaffiliated dicot (LH4, morphotype TY170 of Wilf et al., 2005a,
MPEF-Pb-1605); specimen MPEF-IC-1384a,b, on an unaffiliated
dicot (LH17, morphotype indeterminate, MPEF-Pb-1606); speci-
men MPEF-IC-1386a?c, on an unaffiliated dicot (RP3, morpho-
type indeterminate, MPEF-Pb-1608); specimen MPEF-IC-
1388a?f, on an unaffiliated dicot (RP3, morphotype
indeterminate, MPEF-Pb-1611); specimen MPEF-IC-1389a?f, on
Lomatia occidentalis (LH2, Proteaceae, MPEF-Pb-988); specimen
MPEF-IC-1390a?f, on an unaffiliated dicot (RP3, morphotype in-
determinate, MPEF-Pb-1612); specimen MPEF-IC-1391, on an
unaffiliated dicot (RP2, morphotype indeterminate, MPEF-Pb-
1613); specimen MPEF-IC-1392, on ??Myrcia?? chubutensis
(LH13, morphotype TY21, MPEF-Pb-2406; specimen MPEF-IC-
1393a?e, on an unaffiliated dicot (RP3, morphotype indetermi-
nate, MPEF-Pb-2407); and specimen USNM 40389a,b on the ho-
lotype of ??Anona?? infestans (type material).
Occurrences.?From the early Eocene Tufolitas Laguna del
Hunco Formation, Chubut Province, and from the middle Eocene
R??o Pichileufu? locality, R??o Negro Province, Argentina.
Discussion.?In his recent contribution, Vasilenko (2008) in-
cluded in the ichnogenus Paleoovoidus two new ichnospecies, P.
flavellatus and P. arcuatus. The former ichnospecies was de-
scribed as ??arched oviposition distributed in rows and separated
by distance about a maximum width of the eggs or somewhat
larger.?? This differs from P. arcuatus in having shorter distances
between scars. By contrast, P. arcuatus has slightly arched ovi-
position scars, with the eggs set in rows at considerable distance
from each other parallel to their long axes. This ichnospecies is
more similar to the patterns observed in the Patagonian leaves.
However, a few months prior to Vasilenko?s paper, Krassilov
(2008) established a similar ichnogenus named Sertoveon which
he described as ??egg scars in transverse rows or arches, inserted
at about right angles to the direction of the row.?? Likewise, he
erected two new ichnospecies, S. cribellum and S. arcuatum.
These two ichnospecies also resemble closely the pattern de-
scribed in our material. We consider that both ichnospecies, P.
arcuatus and S. arcuatum, represent the same oviposition pattern.
However, according to the ISCN rules, Paleoovoidus has nomen-
clatorial priority and is the valid ichnogenus; P. arcuatum is the
correct ichnospecies.
The diagnosis is emended herein to include scars of a broader
range of shape (elongate, lens-shaped and teardrop-shaped) that
are distributed both in linear rows and in arcs, which in some
cases can exhibit a zigzag trajectory. Oviposition specimens that
exhibited a zigzag pattern were referred by Krassilov and Silan-
tieva (2008) to a similar ichnospecies, Catenoveon undulatum.
Nevertheless, the principal difference between C. undulatum and
P. arcuatum is the relationship with the primary vein, which is
absent in the later. Moreover, P. undulatum is the only ichno-
species with scars distributed in the leaf lamina without any ap-
parent relation to the primary vein, a feature that differentiates it
from P. bifurcatus scars that are inclined to the primary vein and
from P. rectus scars that occur along major veins. Several authors
have documented scars exhibiting patterns similar to the material
described here under P. arcuatum. Hellmund and Hellmund
(1991, 1993, 1996c, 1998, 2002a), in particular have described
and figured numerous specimens with identifiable scar patterns,
including examples from Kounice and Vys?ehor?ovice (both Upper
Cretaceous) in Bohemia, Czech Republic; and from Messel (mid-
dle Eocene), Salzhausen (middle Miocene), Rott (upper Oligo-
cene), Hammerunterwiesenthal (upper Oligocene), and Berzdorf
and Randecker Maar (both lower Miocene) in Germany. They
distinguished between smaller scars, 1.3 to 1.4 mm in length,
present as an arcuate, parallel pattern, versus larger scars, 1.6 to
1.8 mm long, deployed in a zigzag pattern. The material docu-
mented by Pen?alver and Delclo`s (2004) from the ??La Rincon?ada??
site at the Ribesalbes locality (early Eocene, in Castello?n, Spain)
also show parallel rows occurring in a zigzag pattern from dif-
ferent species composed of ovoidal-oblong scars, with values that
range from 0.9 to 1.1 mm in length and from 0.2 to 0.3 mm in
width. Moreover, the traces documented by Labandeira (2002a),
from the middle Eocene Republic Flora of eastern Washington,
USA, show similar patterns that were defined as separated, equal-
sized, lenticular scars, arranged in semicircular arcs. Scars from
the latest Cretaceous Hell Creek Formation of North Dakota,
USA, show similar morphologies to the ones described here but
lack a clear pattern of arrangement (Labandeira et al., 2002). Re-
cently, Vasilenko (2007, 2008) described specimens on the float-
ing leaves of Quereuxia possessing elongate oviposition scars of
0.75 to 0. 85 mm in length and 0.13 mm in width, distributed in
parallel rows. The Quereuxia material is from the Maastrichtian
age and originates from the Udurchukan locality of the Amur
Region, Russia.
Berry (1938) described scars on the holotype of ??Anona?? in-
festans from the R??o Pichileufu? paleoflora that clearly represent
P. arcuatum ovipositional damage (Fig. 6.1?6.5). However, he
suggested that these marks could represent the bodies of scale
insects (Hemiptera: Coccoidea). This leaf (Fig. 6) shows many
arcuate rows, either parallel or in a zigzag pattern, of lenticular
to teardrop scars distributed over most of the lamina. The tem-
poral occurrence of distinctive Paleoovoidus arcuatum, unlike P.
rectus, and P. bifurcatus, probably is confined to the Cenozoic
and Late Cretaceous; no known occurrences antedate the appear-
ance of angiosperms during the mid Early Cretaceous.
DISCUSSION
Endophytic oviposition has considerable potential for the study
of dragonfly host-plant use and behavioral evolution. This is at-
tributable to the fact that the insertion of the ovipositor into plant
tissue results in a persistent, fossilizable scar of callus tissue that
surrounds inserted, typically nonpreserved eggs. This distinctive,
convex, ovoidal damage has an extensive fossil record on various
plant tissues, displaying a characteristic pattern of linear, curvi-
linear or arcuate rows that occur in plant organs as dark, oval, to
teardrop-shaped scars (Labandeira, 2002a, 2006). In many cases,
ovipositional patterns occur in a parallel or zigzag fashion, or
alternatively, in no pattern at all. By contrast, in exophytic ovi-
position, eggs often are distributed in tight packets or as looser
442 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
clusters on the leaf surface (Corbet, 1999; Labandeira, 2002a;
Labandeira et al., 2007; Vasilenko and Rasnitsyn, 2007; Krassilov
and Silantieva, 2008; Vasilenko, 2008). Moreover, exophytic ovi-
position is done by insects either lacking an external ovipositor
or possessing a modified, nonslicing ovipositor (Labandeira,
2006), and any resulting effects on plant surface features rarely
preserve as fossils. In addition, egg morphology also can be useful
to distinguish the type of oviposition, when infrequently present
(e.g., Sahle?n, 1995; Vasilenko and Rasnitsyn, 2007; Vasilenko,
2008; Pott et al., 2008; Krassilov and Silantieva, 2008). Corbet
(1999) characterized endophytic odonatan eggs as rather elongate,
often flattened, and several times longer than wide, whereas exo-
phytic eggs are more ellipsoidal to subspherical.
Fossil record of insect oviposition.?Ichnofossils of oviposi-
tional traces have been recognized principally in sphenopsid stems
and seed-plant leaves (Roselt, 1954; Kelber, 1988; Schaarschmidt,
1992; Johnston, 1993; Grauvogel-Stamm and Kelber, 1996; Zher-
ikhin, 2002; Banerji, 2004; Beattie, 2007). The oldest record of
oviposition is from an arborescent Pennsylvanian age sphenopsid,
Calamites cistii, bearing pronounced, ellipsoidal scars that were
attributed to the Archaeorthoptera or Palaeodictyoptera (Be?thoux
et al., 2004). The oldest trace fossils assigned to odonatan ovi-
position, referred to the Protodonata, comes from the Lower
Permian flora of the R??o Bonito Formation, Brazil (Adami-Ro-
drigues et al., 2004). Later Paleozoic endophytic oviposition, such
as that from the Upper Permian of Australia, is recognizable but
difficult to assign to a particular producer. For example, an upper
Permian flora from southeastern Australia reveals small circular
insertion scars grouped into several clusters in the foliage of the
extinct sphenopsid cladode Phyllotheca (Beatty, 2007). Presum-
ably, these scars were induced by a member of the Protodonata
or Palaeodictyopteroidea, paleopterous clades that largely became
extinct at the end Permian (Labandeira, 2006).
Pre-Cretaceous, Mesozoic evidence for odonatan ovipositional
damage predominantly originates from biogeographically dispa-
rate Middle and Upper Triassic sites worldwide, and from the
European Lower Jurassic. The most important odonatan ovipo-
sition occurrences, unaffiliated to family, are from the Middle and
Upper Triassic of France and Germany (Grauvogel-Stamm and
Kelber, 1996). In particular, Pott and colleagues (2008) docu-
mented spectacular material, replete with microstructural details
of insect egg and adjacent plant cuticle, from the Upper Triassic
of Austria. Gondwanan occurrences from the Upper Triassic of
Chile (Gnaedinger et al., 2007) and especially the Molteno For-
mation of South Africa (Labandeira, 2006), have documented sev-
eral distinctive ovipositional types (Labandeira et al., 2007). Van
Konijnenburg-van Cittert and Schemie?ner (1999) have docu-
mented younger, new types of ovipositional damage from the
Lower Jurassic of Bavaria (Germany) that may represent an ex-
pansion of dragonfly egg-laying strategies from the earlier Tri-
assic.
More recent, better-defined, endophytic oviposition damage, at-
tributable to particular modern lineages, becomes abundant during
the Cretaceous and especially the Cenozoic (Hellmund and Hell-
mund, 1991, 1993, 1996a, 1996b, 1996c, 1998, 2002a; Lewis,
1992; Labandeira, 2002a, 2002b; Labandeira et al., 2002; Pen?al-
ver and Declo`s, 2004; Vasilenko, 2005, 2008; Vasilenko and Ras-
nitsyn, 2007; Krassilov et al., 2007; Krassilov and Silantieva,
2008). As in Triassic and Jurassic examples of oviposition, Cre-
taceous and Cenozoic material is attributed commonly to the
Odonata, or occasionally to other groups such as the Diptera,
Lepidoptera, Coleoptera, Hymenoptera and Hemiptera (Krassilov
and Silantieva, 2008). Specimens referred originally as coleop-
terous eggs in Alnus (Betulaceae) leaves (Lewis and Carroll,
1991, 1992; Lewis, 1992), later were reidentified as odonatan ovi-
positional damage (Labandeira, 2002a). Many of these docu-
mented descriptions come from the Cenozoic of Germany and
were evaluated by Hellmund and Hellmund (1991, 1993, 1996a,
1996b, 1996c, 1998, 2002a), and typically involve damselfly ovi-
position. The German mid-Cenozoic occurrences reveal consid-
erable behavioral information and are directly comparable with
extant oviposition patterns and associated behaviors.
The biology and paleobiology of odonatan oviposi-
tion.?Odonatan oviposition patterns have received considerable
attention in ecological studies, particularly those emphasizing en-
dophytic placement and behavior (Wesenberg-Lund, 1913a,
1913b, 1943; Schiemenz, 1957; Jurzitza, 1974; Grunert, 1995;
Hellmund and Hellmund, 1998; Corbet, 1999). Matushkina and
Gorb (2007) have emphasized that females of most odonate spe-
cies deposit their eggs into a plant substrate. Females use the two
sets of valves of the ovipositor to penetrate and cut leaf tissue,
inserting eggs within surface or deep tissue (Matushkina and
Gorb, 2007; Corbet, 1999; Miller and Miller, 1988; Labandeira,
2006). Typically, one egg is deposited for each insertion (Tillyard,
1917; Ando, 1962; Hellmund and Hellmund, 2002a; Sahle?n,
1995; Lutz and Pittman, 1968).
The Anisoptera (dragonflies) display a wide variety of ovipo-
sitional modes, involving both exophytic and endophytic egg
placement (Corbet, 1999). However, patterns of endophytic plant
damage are poorly described in the literature for this group, and
the behaviors and mechanisms of egg insertion within leaves or
other plant organs are largely unknown. Considerably more is
known of extant zygopteran ovipositional behavior, in which scars
typically are narrow and have an oval to ellipsoidal surface ex-
pression. For example, the eggs of Lestes eurinus were described
by Lutz and Pittman (1968) as elongate, cylindrical, and uniform
in size. Modern field and laboratory observations at Horco Molle,
in Tucuma?n Province, Argentina, were made for oviposition of
Acanthagrion ablutum (Zygoptera: Coenagrionidae) (Calvert,
1909) on Cynoglossum amabile (Boraginaceae) leaves and for
oviposition of the same zygopteran species on Polygonum punc-
tatum (Polygonaceae) at Dique Escaba, in southern area of Tu-
cuma?n Province. Both of these modern examples display eggs
with a general ellipsoidal shape, whose ends are acute, rounded,
or blunt in shape. The egg structure is characterized by a yellow-
ish to brownish region, which in a few cases bears a small, linear
aperture (Fig. 5.2, 5.3). However, in some cases a white aperture
at one end also is observed. This last pattern matches the ovi-
positional scars in the fossils (Fig. 5.6).
Fossil ovipositional scars described herein and in the literature
share with extant odonatan egg insertions a similar general shape.
This similarity ranges from scars with two similar extreme, round-
ed, or more tapering points, but in other cases with a blunt and
acute end resulting in a teardrop shape. Hellmund and Hellmund
(2002a) noted that several scars possess raised encircling rims and
have a smooth concavity in the central area. They also argued
that small protrusions in the interior depression of the scars could
be the remnants of egg envelopes. These features were not ob-
served on our material. In some specimens (Figs. 3.3, 5.5), a
visible dark spot was observed at the angulate scar end. The same
feature was observed by Krassilov et al. (2007) but described as
preserved callus tissue. By contrast, Krassilov and Silantieva
(2008) show a slender dark band inside the egg impression that
was interpreted as a dried yolk sac or an exuvium of an embryonic
larva.
In other cases (Figs. 4, 5.2, 5.6), scars display a blunt end
cradling a more centrally positioned, rounded or more linear hole
that is distinguishable as a clear zone because of the absence of
plant tissue. This distinctive zone, observed both in the fossils
and in one leaf from the Dique Escaba locality, may represent an
emergence aperture, indicating that the immature nymph had
abandoned the foliar microenvironment for life in water (Fig. 5.6).
Ovipositional behavior varies among the major groups of
Odonata, resulting in a diversity of patterns that characterize ex-
tant and fossil damage on leaf surfaces. Observations of extant
443SARZETTI ET AL.?EOCENE ODONATAN OVIPOSITION FROM PATAGONIA
Zygoptera show that there are differences in ovipositional behav-
ior, for example between Coenagrionidae and Lestidae (Gower
and Kormondy, 1963; Lutz and Pittman, 1968; Hellmund, 1991,
2002b; Matushkina and Gorb, 2007). The endophytic oviposition
of Coenagrionidae includes more-or-less a 1 mm spaced distance
between the oviposition scars that form linear rows, which are
either slightly crescentic, semicircular, or parallel, or alternatively
deployed as a zigzag pattern (Wesenburg-Lung, 1913a, 1913b;
Grunert, 1995; Hellmund and Hellmund, 1991, 1998). Paleoo-
voidus arcuatum clearly resembles this range of pattern. Hell-
mund and Hellmund (1991) were the first to attribute similar pat-
terns in fossil leaves to ovipositional traces of Odonata from the
upper Oligocene of Rott, Germany. These authors originally
named the arcuate and parallel rows as Bogenmodus, and the zig-
zag rows as Zickzackmodus. These alternative styles were com-
pared to extant ovipositional damage in leaves made by damsel-
flies, particularly the Coenagrionidae, and this clade was
determined to be the producer of this pattern. Later, new fossil
evidence from the Hell Creek Formation (Upper Cretaceous) of
North Dakota, U.S.A., attributed oviposition damage on leaves of
Marmarthia pearsonii (Laurales) and Erlingdorfia montana (Pla-
tanaceae) to the Coenagrionidae (Labandeira et al., 2002). More-
over, Pen?alver and Delclo`s (2004) documented Miocene ovipo-
sition trace fossils of Coenagrionidae from La Rincon?ada, Spain,
on leaves of Laurophyllum (Lauraceae), Caesalpinia (Fabaceae),
and Populus (Salicaceae). Similar ovipositional patterns from the
Eocene of Washington were observed by Labandeira (2002a) for
Betula (Betulaceae), Crataegus, Paracrataegus, Sorbus (all Ro-
saceae), and Ginkgo (Ginkgoaceae), some of which initially and
incorrectly were attributed to beetles (Chrysomelidae) by Lewis
and Carroll (1991, 1992). Likewise, Vasilenko and Rasnitsyn
(2007) and Vasilenko (2008) described several examples of elon-
gate and oval endophytic oviposition on leaves of Quereuxia that
were assigned to the ichnotaxonomic group, Paleoovoididae, from
the Maastrichtian of the Udurchukan locality in the Amur Region
of Russia.
Body fossils of Coenagrionidae are known from several Early
Cretaceous localities (Jarzembowski, 1990; Jarzembowski et al.,
1998; Jell, 2004). It is unclear if Paleoovoidus arcuatum origi-
nated within the Coenagrionidae or possibly from an earlier Me-
sozoic ancestor shared with the sister-clade Platycnemidae (Be-
chley, 1996), which generates a similar pattern of oviposition
(Watanabe and Atachi, 1987). A similar pattern of damage, found
in Agrion (Wesenberg-Lund, 1943, fig. 57), a member of the dis-
tant zygopteran clade Calopterygidae, may represent behavioral
convergence.
A different mode of oviposition is represented by species of
Lestidae that lay eggs in double rows at both sides of a primary
vein or occasionally in a single row along one side of the vein.
At insertion, lestid eggs normally are oriented from an acute to a
perpendicular angle relative to a major vein. Structural details of
the ovipositional process were provided by Matushkina and Gorb
(2007), including the ovipositional behavior of Lestes sponsa and
L. barbarus, indicating that the major axis of paired or single
eggs is parallel to plant fibers. Similarly, the fossil oviposition
pattern described here and assigned to Paleoovoidus bifurcatus
probably was produced by damselflies of the family Lestidae.
Hellmund and Hellmund (1991, 1996c, 2002b) described the
same type of oviposition that we attribute to P. bifurcatus in a
leaf of Cinnamomum sp. (??Daphnogene,?? Lauraceae) from the
late Oligocene and in a unidentified leaf from the middle Eocene
of Germany, a pattern they named Doppelreihenmodus.
Ovipositional damage distributed in a single row, assigned here
to Paleoovoidus rectus, also has been attributed to the Lestidae
based on extant foliar damage patterns. Modern lestid oviposi-
tional scars have been observed to occur within major veins form-
ing a single concatenated row, characterized by parallel alignment
of long axes of the elongate scars (Gower and Kormondy, 1963).
The same pattern has been observed by Hellmund and Hellmund
(1991, 1996a) in various dicot leaves. Some specimens (Fig. 2.2,
2.3) display subparallel ovipositional insertion in a single row, of
which individual scars may occur in both midrib vascular tissue
and in the leaf-blade. This pattern also consists of scars that ap-
pear in a single, straight row described by Vasilenko (2005) and
herein, but it has not been documented extensively in modern
foliar material.
Some of our specimens (Fig. 7.1?7.5), included under Paleoo-
voidus arcuatum, show some disorder in the distribution of scars
in contrast to the distinctive patterns described previously. This
more chaotic distribution may be related to the high density of
scars, which would preclude identification of individual rows,
possibly because of superimposed, multiple ovipositional events
by one or several females in the same leaf. Alternatively, elevated
scar density may be attributable to the normal reproductive be-
havior of an unknown, ovipositing odonatan. This latter alterna-
tive is less likely because distinct linear files of scars were de-
tectable and apparently were overprinted with subsequent waves
of oviposition.
Analyses of the distribution of oviposition scars on fossil leaves
and their comparison to patterns obtained from extant autecolog-
ical studies is important for the interpretation of these ichnofos-
sils. This allows for an understanding of the ovipositional behav-
ior that produced the original damage. Especially in Paleoovoidus
arcuatum, the orientation of each scar in an arcuate or zigzag
trajectory can indicate the position of the female on the leaf dur-
ing oviposition (Hellmund and Hellmund, 1991). Accordingly, if
lines are drawn perpendicular to each linear file of scars, the point
where these lines converge, through triangulation, should be the
abdominal pivot point represented by an anchored thorax. When
the scars are teardrop-shaped, the abdominal pivot point is more
precisely identifiable because the shape of eggs allows a sharper
definition of the perpendicular lines. From this pivot point, the
female lays eggs by swinging her abdomen from one side to the
other, depositing from three to fifteen eggs during each swipe.
The next row likewise is produced after she moves a single step
forward and inscribes another swinging movement of her abdo-
men (Hellmund and Hellmund, 1991; Pen?alver and Delclo`s,
2004). The abdominal pivot point potentially is moved along the
same axis when producing different arcs of scars, resulting in a
distinctive and successive series of en echelon files (Hellmund
and Hellmund, 2002a). These parallel and zigzag patterns may be
combined in the same ovipositional behavior, as shown by field
observations at Horco Molle and Escaba in northwestern Argen-
tina. Oviposition begins with well-defined and independent
curved rows, continues with increasingly arcuate files, and results
finally in a zigzag configuration. This observation indicates that
the same female can produce both types of patterns, Bogenmodus
and Zickzackmodus, from the same ovipositional pivot point,
which should discourage the possibility of separating ichnospecies
based on this modern behavioral character.
Other behavioral traits are reflected in the ichnospecies de-
scribed herein. Some members of the Lestidae insert their eggs
on plant stems near water, instead of leaves. Hellmund and Hell-
mund (1991) mentioned that Lestidae lay eggs at the sides of a
primary leaf vein, usually a midrib, and inferred that they used
this anatomic structure as a guide. (The use of a prominent vein
as a guide may have an evolutionary relationship with linear ovi-
position on stems, extending to the late Paleozoic.) The eggs are
inserted into foliar tissue by moving the abdomen once to the
right and then to the left for each oviposition point, producing
often paired files along the vein (Hellmund, 1991, 2002a). Ob-
servations made on the reproductive habits of extant Lestes rec-
tangularis Say 1839 on leaves of Rumex (Polygonaceae), Typha
(Typhaceae) and Scirpus (Cyperaceae), from Pennsylvania,
U.S.A., showed that females rotate the last eight to ten abdominal
444 JOURNAL OF PALEONTOLOGY, V. 83, NO. 3, 2009
segments 90 degrees to produce this pattern (Gower and Kor-
mondy, 1963). By contrast, coenagrionid oviposition appears not
to be influenced by vein architecture during selection of the foliar
area for egg insertion. This type of oviposition targets different
regions of the leaf without restriction and produces canted ori-
entations of scar files.
Host-plant preferences.?Observations regarding host-plant
preferences for endophytic oviposition have been made for sev-
eral extant zygopteran species (Gower and Kormondy, 1963; Bick
and Hornuff, 1965). Corbet (1963) showed that some zygopteran
species chose one type of plant for oviposition among several
available alternatives. Similar results were obtained by Lutz and
Pittman (1968) who concluded that Lestes eurinus Say, 1839 ovi-
posited only in Sparganium americanus (Sparganiaceae). How-
ever, Kumar and Prasad (1977) did not note any host-plant pref-
erence in their study of the reproductive behavior of Neurobasis
chinensis (Calopterygidae), which oviposited endophytically in
several species of riparian plants.
Only a few of our leaf specimens, each with oviposition scars
at LH or RP, represent the ichnospecies Paleoovoidus rectus and
P. bifurcatus. Given the paucity of occurrences, it is impossible
to determine if ovipositional specialization was present for P. rec-
tus and P. bifurcatus. By contrast, comparatively abundant Pa-
leoovoidus arcuatum is represented by ovipositional damage on
a wide variety of foliage, including specimens of Lauraceae, Pro-
teaceae, Fabaceae, Malvaceae, Cunoniaceae, and Myrtaceae. Sup-
porting this broad spectrum of plant hosts, P. arcuatum only oc-
curs more than once per species in two specimens each of
??Myrcia?? chubutensis and Lomatia occidentalis. These observa-
tions indicate that no preferential host-plant preference was pres-
ent for damselflies in their selection of oviposition leaf substrates.
The South American odonatan body fossil record.?South
America has a significant body-fossil record of Odonatoptera, par-
ticularly because the wings of dragonflies and damselflies pre-
serve well. From the Paleozoic and Mesozoic of Brazil, several
families are known (Martins-Neto, 2005; Moura et al., 2006), par-
ticularly from the Early Permian of the Parana? Basin and the Early
Cretaceous of the Araripe Basin. By contrast, in Argentina odo-
natopteran body-fossil specimens overwhelmingly originate from
Cenozoic deposits (Petrulevic?ius, 2001; Petrulevic?ius et al., 1999;
Petrulevic?ius and Nel, 2002a, 2002b, 2003a, 2003b, 2004a,
2004b, 2005, 2007). There are a few records from the Paleozoic
of Argentina, including two members of the Geroptera in the Ear-
ly Pennsylvanian (Riek and Kukalova?-Peck, 1984; Wootton et al.,
1998; Gutierrez et al., 2000), and multiple occurrences of the
Odonata from the Middle and Late Triassic (Carpenter, 1960;
Martins-Neto et al., 2003). Additionally, Petrulevic?ius and Nel
(2003a, 2005, 2007) previously documented body fossils (wings)
of Odonata from the Laguna del Hunco locality. One zygopteran
specimen was assigned to the extinct family Austroperilestidae
(Austroperilestes hunco) and possibly is related to the extant fam-
ily Perilestidae. The other specimen was assigned to the new ex-
tinct family Frenguelliidae (Frenguellia patagonica). The Austro-
perilestidae and Frenguelliidae records currently represent the
only body-fossil evidence for Odonata from LH; neither family
has been documented at RP. These taxa conceivably could have
produced any of the ichnotaxa observed on LH leaves, but there
is no evidence for or against this other than the presence of the
body fossils. It bears re-emphasis that neither the Coenagrionidae
nor Lestidae are known from body fossils at either locality.
Currently the oldest reliable evidence worldwide of Lestidae is
ovipositional traces from the middle Eocene of Germany (Hell-
mund and Hellmund, 2002b). Additionally, Coenagrionidae ovi-
positional patterns have been documented from the upper Oligo-
cene and upper Miocene of Germany (Hellmund and Hellmund,
1996a, 1998). By contrast, specimens of Paleoovoidus bifurcatus
and P. arcuatum from the LH and RP localities of Patagonia
represent the oldest records of ovipositional damage for lineages
likely to be Lestidae and Coenagrionidae. Based on these Pata-
gonian occurrences and evidence for distinctive damselfly ovi-
positional damage, we infer that, by the early to middle Eocene
boundary interval, species of the Lestidae and Coenagrionidae
and their ovipositional behaviors probably were present at the LH
and RP localities.
CONCLUSIONS
1. Evidence of endophytic odonatan oviposition is documented
from the Laguna del Hunco (LH, early Eocene) and R??o Pichi-
leufu? (RP, middle Eocene) floras of Patagonia, Argentina. Three
types of oviposition are described from LH, assigned to the three
newly defined ichnospecies Paleoovoidus rectus and Paleoovo-
idus bifurcatus, as well as the previously established Paleoovo-
idus arcuatum of Krassilov and Silantieva (2008). By contrast,
only P. arcuatum is recorded from the RP paleoflora. Ichnospe-
cies of Paleoovoidus include elongate to lens-shaped or teardrop-
shaped scars expressed on the surfaces of foliar tissue that are
arranged in distinctively regular or otherwise random patterns. In
some cases, the narrow end of the scar bears a dark spot, but in
other instances, there is a broader, circular, small depression
lodged at the wider end. Both of these features suggest apertures
for the emergence of dragonfly (odonatan) immatures.
2. By comparison with similarities in morphology with extant
oviposition patterns, ichnospecies of Paleoovoidus are attributable
to recognizably modern types, specifically to the Suborder Zyg-
optera. Paleoovoidus rectus and P. bifurcatus. probably were pro-
duced by members of the Family Lestidae; P. arcuatum was pro-
duced by members of the Family Coenagrionidae.
3. The same patterns of egg insertion, particularly for the more
abundant Paleoovoidus arcuatum, were present for several leaf
morphotypes, but no host-plant specificities were observed.
4. Evidence from the Eocene of South America demonstrate
that certain odonatan taxa have maintained similar patterns and
behaviors of endophytic oviposition, represented by scars distrib-
uted in linear, zigzag, or otherwise parallel traces over leaf lam-
inae for approximately a 50 million-year interval. These behav-
iors, by virtue of their ichnotaxonomic synonymy with earlier
Early Jurassic to Late Cretaceous occurrences, considerably an-
tedate the mid Paleogene, and extend deeper into the Mesozoic.
ACKNOWLEDGMENTS
We are grateful to Roberto Straneck for his excellent assistance
with German translation. We thank Alejandra Molina for her help
with field research in collecting extant odonatan oviposition, and
to Alberto Slami for his help with the plant identifications. For
helpful reviews of the manuscript, we thank J. Petrulevic?ius and
D. Vasilenko. In addition, we acknowledge M. Caffa, L. Canessa,
B. Cariglino, I. Escapa, M. Gandolfo, C. Gonza?lez, R. Horwitt,
P. Puerta, E. Ruigomez, H. Smekal, and S. Wing for invaluable
assistance in the field and laboratory. This work was supported
by the Proyecto de Investigacio?n Cient??fica y Tecnolo?gica (PICT)
13286 to J. F. Genise; National Science Foundation Grant DEB-
0345750 to PW, CCL, NRC, and KRJ; National Geographic So-
ciety grant 35229-G2 to PW and NRC; the Evolution of Terres-
trial Ecosystems Consortium of the National Museum of Natural
History to LCS; and the Smithsonian Institution Small Grants
Fund to CCL. Thanks go to Finnegan Marsh for the final draft of
the figures, and to the Nahueltripay family and INVAP (Instituto
de Investigaciones Aplicadas) for land access. This is Contribu-
tion 163 of the Evolution of Terrestrial Ecosystems Consortium
at the National Museum of Natural History, Washington, D.C.
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ACCEPTED 10 FEBRUARY 2009