Geologica Acta: An International Earth Science Journal 1695-6133

Geologica Acta: an international earth science
journal
ISSN: 1695-6133
geologica-acta@ija.csic.es
Universitat de Barcelona
España
Labandeira, C.C.
The Four Phases of Plant-Arthropod Associations in Deep Time
Geologica Acta: an international earth science journal, vol. 4, núm. 4, december, 2006, pp. 409-438
Universitat de Barcelona
Barcelona, España
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Appendix I-IX
Av a i l a b l e o n l i n e a t w w w. g e o l o g i c a - a c t a . c o m
The Four Phases of Plant-Arthropod Associations in Deep Time
1 2
C.C. LABANDEIRA
1 Smithsonian Institution, National Museum of Natural History

P.O. Box 37012, MRC-121 Department of Paleobiology, Washington, D.C., USA 200137012. E-mail: labandec@si.edu
2 University of Maryland, Department of Entomology

College Park, Maryland, USA 20742
ABSTRACT
Vascular-plant hosts, their arthropod herbivores, and associated functional feeding groups are distributed spa-
tiotemporally into four major herbivore expansions during the past 420 m.y. They are: (1) a Late Silurian to
Late Devonian (60 m.y.) phase of myriapod and apterygote, hexapod (perhaps pterygote) herbivores on several
clades of primitive vascular-plant hosts and a prototaxalean fungus; (2) a Late Mississippian to end-Permian (85
m.y.) phase of mites and apterygote and basal pterygote herbivores on pteridophyte and basal gymnospermous
plant hosts; (3) a Middle Triassic to Recent (245 m.y.) phase of mites, orthopteroids (in the broadest sense) and
hemipteroid and basal holometabolan herbivores on pteridophyte and gymnospermous plant hosts; and (4) a
mid Early Cretaceous to Recent (115 m.y.) phase of modern-aspect orthopteroids and derived hemipteroid and
holometabolous herbivores on angiospermous plant hosts. These host-plant and herbivore associations are medi-
ated by seven functional feeding groups: a) external foliage feeding, b) piercing-and-sucking, c) boring (Phase 1
origins); d) galling, e) seed predation, f) nonfeeding oviposition (Phase 2 origins); and leaf mining (early Phase
3 origin). Within about 20 m.y. of each herbivore expansion, there is a biota that expresses the nearly full spec-
trum of later plant-arthropod associations. These four associational phases may be linked to the paleoclimato-
logic variables of greenhouse/icehouse cycles and atmospheric O2 and CO2 levels by uncertain causes, although
some relationship probably is present. The 7 functional feeding groups persist through most of the sampled
interval but harbor host-plants and arthropod herbivores that are spatiotemporally ephemeral. Poor understand-
ing of associations in Phases 1 to 3 is attributed to disproportionate focus on the angiosperm and holometabolan
insect associations of Phase 4.
KEYWORDS Plant-insect-associations. Herbivory Fossil-record. Land plants. Insects. Paleoclimate.
INTRODUCTION vided the basic evolutionary history and ecological struc-

ture to continental (terrestrial + fresh water) ecosystems.
The major macroscopic source for biodiversity on the Nowhere is the combination of taxonomic diversity and
planet is land plants and arthropods (Wilson, 1992; ecological richness better demonstrated than the varied
Schoonhoven et al., 2005). These two hyperdiverse plant-insect associations that have been documented for
clades, and to some extent, fungi, historically have pro- the fossil record during the latest wave of major terrestri-
C.C. LABANDEIRA Plant-Arthropod Associations in Deep Time
alization that commenced about 420 m.y. ago, during the 1999; Wilf et al., 2005) or quantitative assessments of
Late Silurian (Labandeira, 2005a). This major invasion of herbivorized versus original surface area (Beck and
terrestrial environments has continued unabated to the Labandeira, 1998; Adami-Rodrigues et al., 2004b), simi-
Recent and has experienced a dramatic expansion in taxo- lar to examinations of extant plants (Wint, 1983; Coley
nomic speciosity, major clade diversity, biomass abun- and Barone, 1996). To date, there have been a sufficient
dance, biogeographic colonization, and ecologic richness number of such studies regionally and from the latest Sil-
(Labandeira 2002a). Although these five measures are urian to the Pleistocene-Holocene boundary that a global
inextricably linked, it is ecological richness that provides data set now can be assembled and examined for coarse-
the single most effective way of assessing how terrestrial grained patterns involving vascular plants, herbivorous
ecosystems evolve in deep geologic time (Vermeij, 2004). arthropods, and their ecological associations. The results
In particular, it is an examination of plant-insect associa- of these data provide the broadest view of how plant-
tions—which provides the paleobiological evidence for arthropod associations have appeared, expanded, and con-
the fundamental interorganismic links of species within tracted in terrestrial environments, resulting in the bewil-
food webs, communities, and ecosystems—that is the dering ecological richness of today (Wilson, 1992).
central focus of this study.
Although the data provided herein are comparatively
The ecology of plant-insect associations is a signifi- coarse, it is my intention to provide a plant-arthropod asso-
cant aspect of modern ecological research, as illustrated ciational baseline for Phanerozoic terrestrial ecosystems,
by sections of ecological journals devoted to this topic. by which finer patterns, involving taxonomically more cir-
However, the intensity and broad scope of the study of cumscribed plant hosts and arthropod herbivores, can be
modern plant-insect associations has not been matched by further elucidated. In parts of this temporal-taxonomic-
a parallel examination of the fossil record (Labandeira, associational matrix where preservational potential is high
2005b). This disconnect is attributable mainly to an over- and intervals of geologic time are relatively short, more
whelming paleobiological focus on the description, clas- taxonomically circumscribed plant-host clades, or lineages,
sification, and phylogenetic analyses of taxa, rather on the can be specified and investigated for the evolution of their
paleoecological associations of plants and arthropods. component communities of arthropod herbivores (Root,
Recently, however, there has been a gradual and sus- 1973; Futuyma and Mitter, 1996). As an alternative to this
tained, albeit limited, trend toward examination of plant- lineage-by-lineage “vertical” approach, a “horizontal”
insect associations in the fossil record. These studies are approach would examine the topology of a herbivore radia-
characterizable into: (1) studies of a specific host plant tion or demise, such as the geochronologically sudden col-
interaction with an insect herbivore (Amerom and Boers- onization of plant-host lineages by clades of arthropod her-
ma, 1971; Zhou and Zhang, 1989; Waggoner and Poteet, bivores, leading to particular types of stereotyped
1996), typically involving one or perhaps a few associa- associations or specified lineages of insect gallers or leaf
tions; (2) the examination of a particular insect herbivore miners (Farrell, 1998; Wilf et al., 2000; Cook et al., 2002;
functional feeding group, such as leaf mining, on multiple Farrell and Sequeira, 2004). Fossil studies have the poten-
hosts within a flora of interest (Crane and Jarzembowski, tial to provide paleobiological ground-truthing, such as
1980; Lang et al., 1995); or alternatively (3) the compara- calibration rates of colonization or extinction by comparing
tively rare assessment of multiple herbivore associations occurrences of taxa to known time intervals, or tests of evo-
on a single host-plant genus or species, such as Early lutionary hypotheses regarding the origin and evolution of
Devonian Psilophyton DAWSON 1859 (Trant and Gensel, plant-insect associations (Fox, 1988; Boucot, 1990; Jermy,
1985; Banks and Colthart, 1993) or Late Pennsylvanian 1993; Farrell and Mitter, 1993; Thompson, 1994; Price,
Psaronius chasei MORGAN 1959 (Labandeira, 1998c; 1997; Vermeij, 2004; Schoonhoven et al., 2005). It should
Labandeira and Phillips, 1996a, 1996b); and (4) the com- be noted that historically the plant host (paleobotanical),
prehensive study typically of an entire flora and all of its arthropod herbivore (paleoentomological), and association-
herbivore associations, replete with a multitude of plant al (paleoecological) fossil records have been considered
hosts and herbivore damage types. This fourth approach separate (Labandeira, 2002a), and have had minimal cross
of evaluating bulk floras has variously focused on general links for understanding how plants and arthropods actually
qualitative examinations (Straus, 1977; Scott et al. 1985, have interacted in deep geologic time.
Stephenson and Scott, 1992; Grauvogel-Stamm and Kel-
ber, 1996; Ash, 1997; Castro, 1997; Glasspool et al.,
2003), quantitative use of presence-absence data such as METHODS, DATA STRUCTURE, AND RATIONALE
Scott and Taylor (1983) from a comparatively limited
number of specimens representing a single plant-host tax- This is the first compilation of published or otherwise
on, to diverse bulk floras with explicitly described dam- personally observed associational data from the fossil
age types (Labandeira et al., 2002a; Wilf and Labandeira, record to provide plant hosts, types of associations, and
insect herbivores that are partitioned into discrete slices of arthropod-mediated damage on plants; (4) a temporal
time. The associational part of these data is expressed by framework in which these three records are geochrono-
the major types of feeding, known as functional feeding logically related, including recent absolute-age calibra-
groups (Coulson and Witter, 1984; Labandeira, 1998a; tions of the geologic time scale and stage-level correla-
Wilf and Labandeira, 1999). The plant-host taxa generally tions of relevant strata; and (5) a preliminary assessment
are provided at the conventionally accepted ordinal rank of the role between global phases of herbivory and envi-
and the inferred arthropod herbivores are delimited by ronmental variables such as atmospheric oxygen abun-
ordinal and superordinal ranks. The 7 functional feeding dance and surface temperature.
groups of plant-arthropod associations are a) external
foliage feeding, b) piercing-and-sucking, c) boring, d) Initial evaluation of the data was done by recording
leaf mining, e) galling, f) seed predation, and g) oviposi- only those publications that had explicit and compelling
tion. The first six are distinctive feeding strategies; the photographic documentation. Personal observation by the
last one is not a feeding type, but rather a distinctive egg- author of unpublished plant-insect associational data also
laying strategy that involves endophytic insertion of eggs was included. Published and observational data included
into plant tissues by a lancet-like device, the ovipositor in the data base (see Appendix at www.geologica-
(Wesenberg-Lund, 1913), and thus is included for com- acta.com) required demonstration of herbivory, often
pleteness since it has a significant fossil record of plant using multiple criteria, as opposed to detritivory (Laban-
damage (Béthoux et al., 2004). This examination of the 6 deira, 1998a; Scott and Titchener, 1999). For most func-
functional feeding groups and oviposition is supplement- tional feeding groups, one or more of the following four
ed by an earlier, separate study of palynivory and nec- criteria were present (Labandeira, 1998a). First, the pres-
tarivory (Fig. 1; Labandeira, 2000). The additional func- ence of callus or other types of reaction tissue induced by
tional feeding groups of palynivory and nectarivory trauma to a live plant should be demonstrated (Tovar et
(surface fluid feeding) are not considered further in this al., 1995). A second line of evidence, particularly for
study. The plant taxa and phylogenetic framework to external foliage feeding, is the presence of micromorpho-
which the 7 associations were assigned is based mostly logical structures such as veinal stringers, necrotic tissue
on the recent work of Anderson et al. (in press) and flaps, and contiguous cuspules within larger excisions
Hilton and Bateman (2006). that are caused by mandibulate chewing, or other features
that indicate active feeding (Gangwere, 1966; Kazikova,
A context for this contribution is provided by several 1985; Araya et al., 2000). Third is the presence of stereo-
studies of both plant diversity (Knoll and Niklas, 1987; typed feeding patterns that are consistent with known
Niklas et al., 1985) and insect diversity (Labandeira and modern feeding types (Bodnaryk, 1992; Puplesis, 1994;
Sepkoski, 1993; Jarzembowski and Ross, 1996; Dmitriev Heron, 2003). The resulting plant-host damage types
and Ponomarenko, 2002) through time that provide broad involve both a single, unique pattern by a host-specialist
patterns of major clade origination, turnover and extinc- clade as well as a pattern that is convergently arrived at
tion during the past 420 m.y., based on taxa that are by different, often host-generalist, clades (see discussion
assumed to be or are monophyletic from then-current in Labandeira et al. 2002a). Last, within diverse floras,
evaluations (See Labandeira [2005a] for the validity and distinctive plant-host specificity patterns and the targeting
indispensability of the taxic approach in understanding of particular tissue types can reveal herbivory that would
major patterns of insect diversity through time). Although not be expected from physically induced leaf injury such
there has been discussion regarding the more specific and as wind damage or particle impact abrasion (Katterman,
focused associational relationships of these two hyperdi- 1990; Vincent, 1990). In certain instances, criteria includ-
verse and interacting clades through time (Howe and ed links to known, highly distinctive, extant damage types
Westley, 1988; Jolivet, 1998; Shcherbakov, 2000; Ver- (Opler, 1973), sometimes involving both conspecific lar-
meij, 2004; Schoonhoven et al., 2005), modest attempts val and adult instars on the same host-plant species (Wilf
have demonstrated mostly anecdotal evolutionary patterns et al., 2000). Collectively or singly, these four criteria
(Willemstein, 1987; DiMichele and Hook, 1992; Scott et effectively segregated herbivory from detritivory in
al. 1992; Labandeira, 2002a; Zherikhin, 2002a, 2002b). almost all cases. Only in the case of boring in wood, was
This contribution adds to these studies by providing: (1) there often an absence of evidence for consumption of
documentation of the major plant lineages through time live tissue. However, since many holometabolous insect
and identification of particular, ecologically dominant wood-borers live in live cambial tissue but also bore
host-plant clades; (2) a record of major arthropod herbi- through dead secondary xylem, borings were included
vore lineages through time (myriapods and especially whose diameters indicate production by beetles or other
mites and hexapods), including their intervals of domi- holometabolous larvae. Excluded were borings attributed
nance; (3) determination of the temporal distribution of to detritivorous arthropods, such as termite galleries and
the basic functional feeding groups based principally on all small diameter oribatid mite borings (Hueber and
Galtier, 2002), unless there was separate evidence for harbored five or more functional feeding groups, a cutoff
plant response tissues. Some groups of termites and mites of four was used per cell for graphical clarity in figures 3-
are herbivorous (Krantz and Lindquist, 1979; Waller and 6. Collectively, most of the data qualitatively occurred
La Fage, 1987), although their activities are more appro- between the two end-members of single occurrences and
priately registered in the fossil record from the evidence descriptions of multitudinous associations from a single,
of plant damage attributed to external foliage feeders or diverse biota.
as identifiable contents of coprolites.
The data (Appendix; www.geologica-acta.com) repre-
After herbivory was established by scrutiny of illus- sent 60 major clades of tracheophyte plants that have
trated examples from the paleoecological literature, as been considered typically at an ordinal taxonomic rank,
well as personally examined material, these data were with the exception of higher-ranked Devonian plant lin-
included in the Appendix (www.geologica-acta.com). The eages (Rothwell and Serbet, 1994; Hilton and Bateman,
Appendix was partitioned into four subsets, each defined 2006). For reasons of completeness, particularly as it
by the temporal ranges of major host-plant and plant- relates to the origin of the borer functional feeding group,
feeding arthropod clades. In particular, these separations a “woody” Devonian clade of massive, lignified fungi
into four principal intervals were based on a combination (Prototaxites) was added to the host list. For arthropod
of: (1) temporal ranges of major dominant plant-host and herbivores, which are dominantly insects, 18 major clades
arthropod-herbivore clades in the fossil record, (2) extinc- were included, which traditionally have been allocated to
tion or other major turnover events, whether of many ranks from order to superorder. These clades of plant
individual clades or entire biotas, and (3) occasional inter- hosts and insect herbivores interacted with each other
vals of ecologically distinctive associations not present in through 7 plant-arthropod functional feeding groups. The
earlier or later biotas (Knoll and Niklas, 1987; Labandeira quality of assignments for these three elements was best
and Sepkoski, 1993; Stewart and Rothwell, 1993; Laban- for functional feeding groups, whose assignments were
deira, 2002a). obvious; of high but somewhat lesser reliability was iden-
tification of the plant hosts; and the least reliable overall
The relevant literature on fossil plant-insect associa- were the inferred insect herbivores, which occasionally
tions was extensively but not exhaustively surveyed. In lacked diagnostic plant-damage evidence that would con-
those instances where there were multiple duplicative lit- fidently assign the potential culprit to a particular myria-
erature citations documenting the same association or set pod, mite, or insect clade.
of associations, only the most comprehensive or recent
source, or both, were used. Occurrence data from this
search (Appendix; www.geologica-acta.com) represents a TEMPORAL PATTERNS OF HERBIVORY
420 m.y. interval from the latest Silurian to the latest
Pleistocene, and resulted collectively in 181 biotas pos- The temporal distribution of plant hosts, associations,
sessing 424 separate associations. These biotas were allo- and arthropod herbivores resulted in four distinctive her-
cated to 54 distinct, five-million-year bins of time based bivore expansion phases, paralleling in timing the previ-
on age dates or stage-level correlations of each biota. ously identified “evolutionary assemblages” for paly-
These biotas range from a single association between a nivory and nectarivory (Labandeira, 2000; Fig. 1).
plant host and its arthropod herbivore (Karpiński, 1962; Defining features of these assemblages are: (1) a tempo-
Amerom, 1966; Opler, 1982) to a few extensively studied rally constrained and taxonomically distinctive suite of
biotas that contain numerous clades of damaged plant plant host-clades typically at the ordinal rank, (2) a simi-
hosts, several types of associations, and a variety of larly juxtaposed assemblage of arthropod herbivore clades
inferred insect herbivores (Grauvogel-Stamm and Kelber, that consumed or otherwise interacted with plant-host
1996; Ash, 1997; Wilf and Labandeira, 1999; Scott et al., clades, and (3) the presence of a diverse, abundant,
2004). However, for those specific host-plant taxa that preservationally exceptional and associationally rich biota
FIGURE 1 A matrix of vascular plants, insects, and their associations for palynivory and pollination, placed in geochronological context. This matrix
details the four evolutionary assemblages of palynivores, nectarivores, pollinators, and their host plants that are distinct at the highest taxonomic lev-
els, discussed in Labandeira (2000). Fossil plant and animal taxa at top are those that display evidence for plant-insect associations, based on direct
or indirect evidence from dispersed coprolites, gut contents, plant reproductive biology, insect mouthpart structure, plant damage, and modern taxo-
nomic affiliation. Thin horizontal connectors are well-supported associations from identified insects with known plant palynomorphs as gut contents.
Plant taxa with asterisks (*) are spores or pollen; a few Paleozoic taxa refer to form-genera of plant organs with known whole-plant taxonomic attribu-
tions. Taxa persisting to the present are indicated by bold vertical links, solid for Assemblages 3 and 4, respectively. The arthropod culprits of Assem-
blage 1 remain unknown and probably include arthropods other than insects, such as myriapods. This compilation includes the most prominent stud-
ies and is not complete. Modified from Labandeira (2000) to conform to recent changes of the geologic time scale (Gradstein et al., 2004); the two,
topmost intervals of time of the Neogene are, from oldest to youngest, Pliocene and Pleistocene. Abbreviations: Miss.: Mississippian (or Early Car-
boniferous); Penn.: Pennsylvanian (or Late Carboniferous).
early within the development of each of the four expansion for digestion (Shear and Selden, 2001). However, vegeta-
phases. These biotas provide excellent examples of associ- tive parts of a plant also represent continua between less
ational richness within each expansion phase, and can be digestible tissues (e.g. vascular strands) to tissues with
considered as a “flagship” biota to which others within the greater nutritional value (e.g. mesophyll). Additionally,
same expansion phase can be compared (Table 1). pollen and spore protoplasts are accessible by a variety of
insects that extract their contents by a range of processes
Herbivory in the Earliest Terrestrial Ecosystems without subjecting the exterior grain walls to physical
breakage (Scott et al., 1985; Oliveira et al., 2002). Thus,
A controversial issue regarding the trophic structuring consumption of all live plant tissues is considered as her-
of early land-based ecosystems is the role of detritivores bivory, whereas the feeding on dead plant, fungal, and
versus herbivores in channeling the primary production of animal tissues is considered detritivory. It is this latter
land plants into the food web. An early view was that ear- definition, proposed by ecologists, which provides the
ly land ecosystems were scaled-down or otherwise simple context for the initial herbivory on land (Figs. 1 to 3), as
versions of extant ecosystems, containing the full compli- represented by Herbivore Expansions 1 and 2.
ment of major trophic groups, including herbivores. An
alternative view is that the earliest ecosystems either Temporal Lags between the Origin of Plant
lacked herbivores or the consumption of live plant tissues Tissues and their Herbivory
was insignificant (Shear and Selden, 2001; Habgood et
al., 2004). Given the evidence, principally from coprolites The Paleozoic fossil record of arthropod use of vascu-
and plant damage from several major Late Silurian–Mid- lar plant tissues is one of plant-host histological diversifi-
dle Devonian biotas, the truth is somewhere in between, cation up to the Pennsylvanian, with minor to major tem-
with detritivory and palynivory dominant and herbivory poral lags between the time of anatomical origin of
and carnivory subordinate. The earliest herbivory was a particular tissues and the time of origin of their herbivore
consequence of a few modes of feeding, but qualitatively consumption (Fig. 2). In some cases during the Late Sil-
much less than that of the subsequent Carboniferous urian to Early Devonian there was relatively sudden and
(Labandeira, 2002a). This conclusion is based on the early colonization by palynivorous herbivores of sporan-
sparse occurrence of insect-mediated plant lesions or bor- gia and their spores (Figure 1; Edwards et al., 1995; Hab-
ings during the Late Silurian and Devonian (Labandeira, good et al., 2004) by piercer-and-suckers of stems (Figure
1998a). The tracking of plant organs and the first instance 2; Kevan et al., 1975; Banks and Colthart, 1993), and
of herbivory on them has a bimodal distribution (Fig. 2), wood precursor tissues in the form of borings within the
suggesting a geologically earlier targeting of stems and hardened basidiomycete fungus Prototaxites DAWSON
spores but a delayed attack on roots, leaves, wood and 1859 (Hotton et al., 1996.). By contrast, subsequently
seeds that was launched during the Late Missi- derived tissues, occurring in organs such as roots, leaves,
ssippian–Middle Pennsylvanian (Iannuzzi and Laban- and seeds, originate from the later Early to Late Devon-
deira, unpubl. data). Although many recognize herbivory ian, yet do not exhibit evidence for earliest herbivory until
in the early land plant record, a dissenting view holds that the Late Mississippian, representing lags from ~ 50 to
some of these damaged tissues were either mediated by 100 m.y. This latter herbivory event initiates Herbivore
abiological agents or may not constitute herbivory (Shear Expansion Phase 2, which includes folivory (Iannuzzi and
and Selden, 2001). There have been a significantly greater Labandeira, unpubl. data), a second expansion of boring
number of relevant studies of Pennsylvanian-age, when into the true wood of lignophyte vascular plants (Dunn et
compared to Mississippian-age, floras (DiMichele and al., 2003), seed predation (Jennings, 1974), and roots
Hook, 1992), in addition to the fact that the Pennsylvan- (Labandeira, 2001). A similar lag also is evident at the
ian is about one-third the duration of the Mississippian. commencement of Herbivore Expansion 3, in which the
Both of these factors may account for an intrinsically earliest example occurs from the early Middle to early
deficient record of herbivory during the Mississippian. Late Triassic, postdating the prior presence of relevant
plant-host clades by several million years, especially true
Another issue is what constitutes ‘herbivory?’ The for the Karoo Basin of South Africa (Scott et al., 2004;
long-standing view of insect ecologists is that herbivory is Labandeira and Anderson, 2005.). Similarly, but perhaps
the ‘…consumption by animals of any plant parts, includ- more muted, was herbivore response that has been noted
ing foliage, stems, roots, flowers, fruits or seeds’ for the aftermath of the end-Cretaceous extinction within
(Schowalter, 2000). An alternative concept of herbivory is Herbivore Expansion 4, in which herbivore levels and
that only consumption of the vegetative parts of the plant richness during the latest Cretaceous were not reached
constitutes herbivory, and that seeds, spores and pollen until the Paleocene/Eocene boundary, some 10 m.y. after
and possibly roots are an exception because they are the extinction event (Wilf et al., 2001; Labandeira et al.,
either too nutritionally rewarding or too difficult to access 2002b). The existence of these lags between the initial
TABLE 1 Salient featuresof the four phases of herbivore expansion (excluding palynivory and nectarivory).
Herbivore Major Number

expansion Major arthropod Early Functional Number of biotas Number
phase, interval, host-plant- herbivore flagship feeding of asso- investi- of 5 m.y.
and duration clades1 clades1 biotas groups ciations gated intervals
1. 420–360 Ma protracheo- myriapods, Rhynie 3 10 5 5

(~60 m.y.) phytes, apterygotes
trimerophytes,
rhyniopsids,
lycopsids,
zosterophylls,
pteridophytes,
prototaxales2
2. 335–251 Ma pteridophytes, myriapods, Calhoun 7 57 31 14

(~ 85 m.y.) stem mites,
spermatophytes, apterygotes,
(crown palaeo-
spermatophytes) dictyopteroids3
“prot-orthopteroids”
3. 245 Ma–present pteridophytes, mites, Molteno 8 95 56 33

(245 m.y.) crown orthopteroids,
spermatophytes, hemipteroids,
(stem early or
spermatophytes) plesiomorphic
holometabolan
clades
4. 115 Ma–present angiosperms, orthopteroids, Dakota 8 263 88 22

(115 m.y.) (pteridosperms), hemipteroids,
(conifers) late or
apomorphic
holometabolan
clades
1See text for listing of subclades within these groups; parentheses indicate a subdominant clade of plant hosts; groups in quotes are para-
phyletic.
2A columnar basidiomycete fungus (Hueber, 2001).
3A monophyletic clade united by unique mouthpart construction and consisting of the subclades Palaeodictyoptera, Megasecoptera,
Diaphanopterodea and Dicliptera (Grimaldi and Engel, 2005).
appearance of available tissues and their initial consump- vascular plants, namely protracheophytes, trimerophytes,
tion by arthropod herbivores may be a general feature of rhyniopsids, lycopsids, zosterophylls and pteridophytes,
the plant-arthropod associational record. but also the massive basidiomycete Prototaxites (Hueber,
2001). Herbivorous arthropods probably were myriapods
Herbivore Expansion 1: Late Silurian to Mid- (diplopods and arthropleurids), mites, and to a lesser
Carboniferous extent apterygote hexapods and possibly true insects.
Four plant-arthropod associations are represented. These
Records of Herbivore Expansion 1 (Fig. 3), which associations are external feeding (Trant and Gensel, 1985)
persisted for about 60 m.y., originate from the coastlines and piercing-and sucking (Kevan et al., 1975; Banks and
of Euramerica (Appendix; www.geologica-acta.com). Colthart, 1993), both on photosynthetic tissues of stems;
This phase consists of 5 examined biotas, each occurring boring in the cortical and medullary tissues of the indurat-
in a five-million-year interval, but clustered toward the ed axial organs of Prototaxites and piths of land plants
latest Silurian and Early Devonian. Three functional feed- (Kidston and Lang, 1921; Hueber, 2001); and palynivory
ing groups (excluding palynivory; see Labandeira, 2000) of spores and sporangia (Edwards et al., 1995; Habgood
and 10 discrete associations are present (Figs. 1 and 3; et al., 2004; Figs. 1 to 3). The dominantly targeted plant
Table 1). Dominant host plants consist of basal clades of host appears to be the Rhyniopsida. The dominant herbi-
vore group may have been subgroups of myriapods. The 2005a), and provide a reasonable spectrum of feeding
most pervasive association is external feeding on photo- types involved in the consumption of several available tis-
synthetic tissues. These sparse data document the relative- sue types from primitive vascular-plants. Tissues or parts
ly early origin of herbivory in terrestrial habitats within of organs that originated later during the Devonian, such
predominantly detritivore ecosystems (Labandeira, as leaves for folivorous external feeders, wood for borers
FIGURE 2 Occurrence data provided for the first appearance and initial herbivore colonization of six organ or tissue types. Colonization of Herbivore
Expansion 1 includes consumption of spores (Edwards et al., 1995; Edwards, 1996; Habgood et al., 2004); consumption of stem tissues by both
external feeders (Kevan et al., 1975; Trant and Gensel, 1985; Banks and Colthart, 1993) and by piercer-and-suckers (Kevan et al., 1975; Banks and
Colthart, 1993); and borers of “woody” tissues in the form of lignified cortical and medullary tissues, depicted in Fig. 7. Borings occurred initially in
vegetative portions of the massive basidiomycete fungus Prototaxites DAWSON 1859 (Hueber, 2001) that bear borings (Arnold, 1952; Hotton et al.,
1996; Hueber, 2001; Labandeira pers. observ.), shown by the finely stippled Devonian pattern, and later as Late Mississippian and Pennsylvanian
borings in plant hosts (Dunn et al., 2003), shown in the coarsely stippled pattern. These three latter functional feeding groups, and their plant hosts
and inferred insect herbivores, are shown in Fig. 3. Note that Prototaxites is a fungus (Hueber, 2001), but served as an initial host for borers of ligni-
fied tissues, a functional feeding group that later may have independently re-evolved on lignophyte plants by a different arthropod clade. Herbivore
Expansion 2 commences during the later Mississippian, and contains a second invasion of borings in indurated tissues (Dunn et al., 2003), this time
on lignophytic seed plants (Kenrick and Crane, 1997). Additionally, there is evidence for the initial herbivory on leaves, or external foliage feeding
(Iannuzzi and Labandeira, unpubl. data), initial seed predation (Jennings, 1974), and initial root feeding (Labandeira, 2001). Root feeding is very
poorly documented in the fossil record, and is not included in any figure as a distinct functional feeding group.
FIGURE 3 Herbivore Expansion 1. Illustrated is

the distribution of functional feeding groups
within specified biotas (at right), on accompa-
nying plant hosts within specified five-million-
year intervals (at left). Inferred insect herbi-
vores are provided in a middle panel that link
functional feeding groups with plant hosts with-
in each binned interval. Color symbols for func-
tional feeding groups (basic associations) are
at upper-left, and apply throughout the chart to
inferred insect herbivores and plant hosts for
each successive 5-million-year interval ranging
from the Late Silurian to Late Devonian. Arrow
indicates the “flagship” associational biota for
Expansion Phase 1, the silica permineralized
Rhynie and Windyfield Cherts of Scotland; Unit-
ed Kingdom. See Fig. 1 for the distribution of a
fourth functional feeding group, palynivory, not
displayed herein. Plant-host clades are from
Kenrick and Crane (1997); arthropod-herbivore
clades are from Wheeler et al. (2004). Geolog-
ic time scale is after Gradstein et al. (2004).
See Appendix at www.geologica-acta.com,
Table 1, and text for additional details. Abbrevi-
ation: C: Carboniferous; uppermost stage of the
Silurian is Přídolí.
on more massive plant trunks, or seeds for seed predators, boundary (Brauckmann et al., 1995). The other consists
were not consumed until the beginning of Herbivore of borings in plant axial tissues from slightly younger
Expansion 2. strata (Dunn et al., 2003). Notably, both occurrences are
on pteridosperm taxa. Documentation for Phase 2, which
The early “flagship” biota for Herbivore Expansion 1 lasted approximately 85 m.y., principally originates from
is the permineralized Rhynie and Windyfield Cherts from Euramerica during the Early Pennsylvanian to mid Early
southern Scotland, of Early Devonian (Pragian) age Permian, particularly the interior peat-bearing basins in
(Trewin and Rice, 2004). These deposits are part of the the eastern United States and western Europe and from
Dryden Flags Formation, and are interpreted as preserved Gondwanaland during the mid Early Permian to the latest
siliceous hot springs within a region of lacustrine and flu- Permian, especially the Karoo Basin of South Africa, the
vial sedimentary environments (Trewin, 1994; Habgood Sydney and Bowen Basins of Australia, and the Paraná
et al., 2004). The associated biota consists of early land Basin of east-central South America (Appendix;
plants, a relatively diverse assemblage of freshwater and www.geologica-acta.com). This phase of distinctive her-
especially terrestrial arthropods, and a modest number of bivory documents 6 functional feeding groups (only leaf
plant hosts and arthropod associations. mining is absent), representing 57 separate associations
from 31 biotas that occur among 14 five-million-year
Herbivore Expansion 2: Mid Carboniferous to intervals. Palynivory is not included in this tabulation
the Permo-Triassic Boundary (Labandeira 2000). The dominant host plants consist of
pteridophytes, mostly stem spermatophytes (seed plants),
The earliest expression of Herbivore Expansion 2 and to a lesser extent crown spermatophytes (sensu Hilton
(Fig. 4) is documented from two occurrences during the and Bateman, 2006) (Table 1; Figs. 1 and 4). Herbivorous
Late Mississippian. The older example is the earliest arthropods are significantly more varied than those of
occurrence of external foliage feeding (Iannuzzi and Phase 1, and consist principally of myriapods, mites, and
Labandeira, unpubl. data), about 6 million years prior to particularly paleopterous and neopterous insects, the lat-
the sudden and major increase in the taxonomic diversity ter two of which predominantly consist, respectively, of
of winged insects at the Mississippian-Pennsylvanian the monophyletic paleodictyopteroids and the paraphyletic
“protorthopterans.” The plant-host clades that were most non occurs both in compression floras, representing flu-
targeted overall by herbivorous arthropods were medul- vial subenvironments, and peat-substrate wetland envi-
losan pteridosperms during the Pennsylvanian and Early ronments (Labandeira, 2001). In a similar way glos-
Permian in Euramerica, and glossopterid pteridosperms sopterids were preferentially targeted during the Permian
throughout the Permian in Gondwana. Subordinately, in Gondwana (Plumstead, 1963; Holmes, 1995; Adami-
Euramerican lyginopterid pteridosperms were attacked Rodrigues et al., 2004a). An interesting pattern is the rela-
during the Pennsylvanian and cordaites in biogeographi- tively balanced distribution of all 6 associations (Fig. 4),
cally disparate sites during the Late Pennsylvanian and with the exception of overall dominance by external
Early Permian. One of the major trends of Herbivore foliage feeding.
Expansion 2 during the Pennsylvanian and Early Permian
is the launching of herbivory on medullosan pteri- The Calhoun Coal, from southeastern Illinois, U.S.A.,
dosperms in Euramerica, which exhibits by far the great- of early Late Pennsylvanian (Kasimovian) age, is a mem-
est amount of herbivore-mediated damage, especially ber of the Mattoon Formation, and is a calcite permineral-
external foliage feeding, of any contemporaneous plant ized coal-ball deposit representing a peat-swamp forest
clade, both in terms of herbivorized pinnule area and fre- dominated by marattialean tree-ferns and subordinately
quency of attack (Greenfest and Labandeira, 1997; Beck by medullosan seed ferns (Labandeira, 2001). No known
and Labandeira, 1998; Labandeira, 2001). This phenome- arthropod fauna has been found from this deposit, attri-
FIGURE 4 Herbivore Expansion 2. Illustrated is the distribution of functional feeding groups within specified biotas (at right), on accompanying plant
hosts within specified five-million-year intervals (at left). Inferred insect herbivores are provided in a middle panel that link functional feeding groups
(basic associations) with plant hosts within each interval. Color symbols for functional feeding groups are at upper-left, and apply throughout the
chart to inferred insect herbivores and plant hosts for successive 5-million-year intervals ranging from the Late Mississippian (Early Carboniferous)
to Late Permian. Arrow indicates the “flagship” associational biota for Expansion Phase 2, the carbonate permineralized Late Pennsylvanian (Late
Carboniferous) Calhoun Coal of Illinois, U.S.A. See Fig. 1 for the distribution of an additional functional feeding group, palynivory, not displayed here-
in. Plant-host clades are from Rothwell and Serbet (1994) and especially Hilton and Bateman (2006); arthropod-herbivore clades are from Codding-
ton et al. (2004) and especially Willmann (2004), except for the “protorthopteroid complex” which is either paraphyletic or monophyletic in the
sense that the group is used herein. The Palaeodictyopteroidea is considered to be monophyletic, united with distinctive mouthpart characters, and
consists of the subclades Palaeodictyoptera, Megasecoptera, Diaphanopterodea and Dicliptera (Grimaldi and Engel, 2005). Geologic time scale is
after Gradstein et al. (2004). See Appendix at www.geologica-acta.com, Table 1 and text for additional details. Abbreviation: T: Triassic.
butable to the apparent lack of chitin preservation. Never- 245 m.y. from the Middle Triassic to the Recent. It is like-
theless, the superb anatomical preservation of plant tis- ly that many of the Late Cretaceous to Neogene associa-
sues in the Calhoun Coal has resulted in the most diverse tions of Herbivore Expansion 4 were laterally transferred
assemblage of plant-insect associations for any known from angiosperm hosts onto the gymnospermous hosts of
deposit of Herbivore Expansion 2 (Labandeira and Herbivore Expansion 3, and thus are temporally distinct
Phillips, 1996a, 1996b, 2002). Evidence for insect-medi- from the more ancient, Mesozoic associations such as
ated herbivory is principally from varied types of anatom- those detailed by Farrell (1998).
ically preserved endophytic and exophytic plant damage
that reveal details such as mouthpart stylet tracks, callus The dominant host plants of Herbivore Expansion 3
and other types of reaction tissue such as those from are pteridophytes, crown spermatophytes, and, to a lesser
stylet tracks and galls, and diverse populations of copro- extent, stem spermatophytes, the latter of which were
lites whose contents preserve pollen, spores, and vegeta- experiencing replacement by more derived gymnosper-
tive tissues assignable to both plant-host tissues and taxa. mous clades. Herbivorous arthropods comprise principal-
This evidence allows identification at the species level of ly mites, orthopteroids (Orthoptera, Holophasmatodea),
almost all interacting host plants (Labandeira, 1998a, hemipteroids (Lophioneurida, Thysanoptera, Auchenor-
1998c). rhyncha, Sternorrhyncha) and early or otherwise ple-
siomorphic holometabolan clades (such as Archostemata,
Although this account is concerned principally with Nemonychidae, Belidae, Symphyta, Aneuretopsychina
plant-herbivore associations, the pattern of Paleozoic (sensu Rasnitsyn and Kozlov, 1991), Nemestrinidae,
detritivore associations also expresses a significant expan- Archaeolepidae, Micropterygidae and Agathiphagidae).
sion coincident with the second phase of herbivory. Com- There is a distinctive evolution of major host-plant use by
mencing during the Middle Mississippian (Visean), par- herbivores within Phase 3, beginning with a variety of
ticularly in Euramerican floras, there is a major increase ginkgoopsids, peltasperms, broadleaved voltzialean (e.g.
of oribatid mite detritivory in a variety of host-plant and Heidiphyllum) and podocarpalean (Podocarpaceae +
environmental settings (Rex, 1986; Scott, 1977; Scott et Araucariaceae) conifers, and perhaps pteridophytes that
al., 1992; Labandeira et al., 1997; Tomescu et al., 2001), were colonized during the Middle to early Late Triassic
mixed with limited evidence for herbivory (Dunn et al., by all functional feeding groups (Anderson and Anderson,
2003). This degradation of a wide variety of Middle Mis- 1985; Grauvogel-Stamm and Kelber, 1996; Ash, 1997).
sissippian plant tissues continued throughout the Paleo- This diversification was followed during the Jurassic to
zoic and was extensively supplemented during the later Early Cretaceous by a shift to a variety of cycads, non-
Mesozoic by clades such as termites and wood roaches voltzialean conifers, bennettitaleans and pentoxylaleans
(Labandeira et al., 1997; Nalepa et al., 2001). that were colonized by borers and subordinately by seed
predators. Probable culprits for some of these associations
Herbivore Expansion 3: Middle Triassic to Recent include principally beetles (Crepet, 1974), such as the
Obrieniidae (Klavins et al., 2005) and Nitidulidae (Nishi-
The lower boundary of Herbivore Expansion 3 (Fig. da and Hayashi, 1996). Late Cretaceous to Neogene
5) is probably the end-Permian event, coupled with major occurrences emphasize the conifer order Pinales (includ-
floral turnover during the Late Permian. The earliest evi- ing “cupressoid” families) in which boring and seed pre-
dence for Herbivore Expansion 3 is found in three princi- dation virtually are the only remaining major associations
pal regions during the Middle Triassic, but continues (Fig. 5). The dominant insect herbivore taxon throughout
through the early Late Triassic. They are the southwestern Herbivore Expansion 3 is the Coleoptera, early Triassic
United States (Walker, 1938; Ash, 1997, 2000, 2005), occurrences of which emphasize external leaf mining,
Western Europe (Linck, 1949; Roselt, 1954; Grauvogel- external foliage feeding, and especially boring on many
Stamm and Kelber, 1996), and the Karoo Basin of South broadleaved gymnospermous clades. Subsequently, there
Africa (Anderson and Anderson, 1989; Scott et al., 2004). was the re-emergence of seed predation, and especially
This worldwide pattern extends into the Mesozoic and is wood boring by saproxylic coleopterans during the Lower
supplemented with deposits from all continents. Herbi- Cretaceous on pinaceous conifers (Jarzembowski, 1990;
vore Expansion 3 contains the full modern compliment of Falder et al., 1998; Ratzel et al., 2001). These clades, par-
7 functional feeding groups (Figs. 1 and 5; Table 1) that ticularly the Curculionoidea, were involved in the recolo-
provide 95 distinct associations from 56 biotas that repre- nization of xylic substrates in pinaceous conifers and
sent 33 five-million-year intervals. (Palynivory and sur- included invasion of cambial tissues. During the Neogene
face fluid feeding are excluded from Fig. 5; Labandeira, the emergence of galling Diptera, principally Cecidomyi-
2000). Notably, unlike the other three major assemblages idae (Labandeira, 2005c), supplemented existing associa-
of plant-insect associations, Herbivore Expansion 3 has tions. Throughout Phase 3, odonatopterans evidently
the longest persistence in geologic time, lasting for about oviposited in pteridophytes and a restricted variety of
hosts within specified five-million-year intervals (at left). Inferred insect herbivores are provided in a middle panel that link functional feeding groups
(basic associations) with plant hosts within each interval. Color symbols for functional feeding groups are at upper-left, and apply throughout the chart to
inferred insect herbivores and plant hosts for successive 5-million-year intervals ranging from the early Middle Triassic to the Recent. Arrow indicates
the “flagship” associational biota for Expansion Phase 3, the compression Late Triassic Molteno Formation of South Africa. See Fig. 1 for the distribution
of additional feeding groups, palynivory and nectarivory, not displayed herein. Plant-host clades are from Rothwell and Serbet (1994), Anderson et al. (in
press) and especially Hilton and Bateman (2006); arthropod-herbivore clades are from Coddington et al. (2004) and especially Willmann (2004), with
“Orthopteroidea” synonymous with his “Polyneoptera”. Geologic time scale is after Gradstein et al. (2004); the two, topmost intervals of time of the Neo-
gene are, from oldest to youngest, Pliocene and Pleistocene. See Appendix at www.geologica-acta.com, Table 1 and text for additional details.
gymnospermous seed plants. Evidence for piercing-and- herbivory contains 7 functional feeding groups represent-
sucking is rare. ing 262 associations from 89 biotas among 22 five-mil-
lion-year intervals. Palynivory and surface fluid feeding
The Molteno Formation is of Late Triassic (Early are excluded from the listed functional feeding groups
Carnian) age that yields compression fossils of diverse (Labandeira, 2000). The primary data documenting Her-
plants and insects that geographically span much of the bivore Expansion 4 is worldwide in origin, although the
Karoo Basin of South Africa, and has been extensively greatest concentrations of localities are in Western Europe
examined since the late 1960’s (Anderson and Anderson, and North America, primarily because of greater paleon-
1983, 1985, 1989, 2003; Scott et al., 2004). This “flag- tological examination rather than a necessarily richer
ship” deposit for Herbivore Expansion 3 contains about record. The relatively compact but even distribution of
100 anthropogenically unbiased and censused localities taxa and associations during this phase resembles that of
representing an estimated 180,000 plant-organ specimens Herbivore Expansion 2 of the late Paleozoic rather than
(overwhelmingly leaves and seeds), many of which pro- that of the earlier and contemporary phase of Herbivore
vide clear and abundant evidence for external foliage Expansion 3, the latter of which is twice as long-lived and
feeding, piercing-and-sucking, leaf mining (earliest displays a more open matrix of colonized plant-hosts, their
known occurrence), galling, seed predation, and oviposi- herbivores and associations (Figs. 5 to 7). Interpretations of
tion (Anderson and Anderson, 1989, 2003; Scott et al., this latter pattern are that it either reflects a real biological
2004; Labandeira and Anderson, 2005). The major plant- signal, or alternatively represents an absence of investiga-
host clades include sphenophytes, filicalean ferns, tion, particularly for the Jurassic and Early Cretaceous.
voltzialean conifers, peltasperms, cycads, subclades of
ginkgoopsids that include ginkgoaleans, as well as pen- Twenty-eight orders of angiosperms, including a sig-
toxylaleans, and gnetopsids (Anderson and Anderson, nificant number of unassignable taxa, constitute the plant
2003; Anderson et al., in press; Hilton and Bateman, hosts for Herbivore Expansion 4. Nine identifiable plant-
2006). Several major habitats have been reconstructed host orders have the most occurrences (Fig. 6), all of
from the floristic data: Dicroidium riparian forest (two which bear the full or almost full spectrum of functional
types), Dicroidium woodland, Sphenobaiera woodland, feeding groups and colonization by the major groups of
Heidiphyllum thicket, Equisetum marsh, and a fern–Kan- arthropod herbivores. However, there is a threefold, suc-
naskoppifolia meadow (Anderson et al., 1998; Anderson cessive pattern of insect colonization of families within
and Anderson, 2003). These taxonomic and community- each of these orders. First, the Laurales (Lauraceae, Chlo-
level data indicate a diverse, ecologically heterogeneous ranthaceae) and Proteales (Platanaceae, Trochoden-
ecosystem of several million years duration with exten- draceae) have among the earliest occurrences but also
sive associations among insect herbivores (Scott et al., have the most persistent record of colonization, well into
2004; Labandeira and Anderson, 2005). The Molteno For- the late Neogene. Second, the Saxifragales (Cercidiphyl-
mation overwhelmingly contains the most diverse associ- laceae, Hamamelidaceae) represents a subsequent, later
ations of any deposit from Herbivore Expansion 3, and Cretaceous to mid-Eocene colonization. Third, the
was present within 20 m.y. of commencement of the remaining six orders—Malpighiales (Malpighiaceae,
phase. The absence of any subsequent Jurassic or earliest Euphorbiaceae, Salicaceae), Fabales (Fabaceae sensu
Cretaceous biota with equivalent levels of associations as lato), Rosales (Rosaceae, Ulmaceae, Celtidaceae,
the Molteno is anomalous, and may be attributable to a Moraceae), Fagales (Fagaceae, Betulaceae, Juglandaceae,
poor fossil record or an intrinsic decrease in associational Myricaceae), Sapindales (Sapindaceae, Anacardiaceae,
complexity in most ecosystems. Rutaceae), and Ericales (Ericaceae, Theaceae,
Sapotaceae)—have late Paleocene to Recent patterns of
Herbivore Expansion 4: Mid Lower Cretaceous exploitation by arthropod herbivores that are represented
to Recent by all functional feeding groups. The principal arthropod
clades of interacting herbivores are orthopteroids
The most recent and intensively studied phase of (Orthoptera, Phasmatodea), crown-group hemipteroids
arthropod herbivory on vascular plants is Herbivore (especially Sternorrhyncha and Heteroptera), and princi-
Expansion 4 (Fig. 6). This phase is synonymous with the pally crown-groups of holometabolous insects (Polypha-
angiosperm radiation, and their herbivore associations are ga, Formicidae, Apoidea, Ditrysia, Cyclorrhapha).
predominantly with more apomorphic clades of hemi- Notably, the density of functional feeding groups is more
pteroid and holometabolous insect orders. The 115 m.y. intense for Herbivore Expansion 4 than any of the three
duration of this phase parallels the last half of Herbivore preceding or contemporaneous phases, and no particular
Expansion 3, although their temporal overlap suggests dietary mode is dominant. Additionally, there is more par-
evolutionary host switching of herbivores to and from titioning of individual plant clades by multiple functional
plant-host clades of both major expansions. This phase of feeding groups and arthropod herbivores. For individual
hosts within specified five-million-year intervals (at left). Inferred insect herbivores are provided in a middle panel that link functional feeding
groups (basic associations) with plant hosts within each interval. Color symbols for functional feeding groups are at upper-left, and apply throughout
the chart to inferred insect herbivores and plant hosts for successive 5-million-year intervals ranging from the mid Early Cretaceous to the Recent.
Arrow indicates the “flagship” associational biota for Expansion Phase 4, the compression latest Early Cretaceous Dakota Formation of Kansas,
U.S.A. See Fig. 1 for the distribution of additional functional feeding groups, palynivory and nectarivory, not displayed herein. Plant-host clades are
from Soltis and Soltis (2004); arthropod herbivore clades are from Coddington et al. (2004) and especially Willmann (2004), with “Orthopteroidea”
synonymous with his “Polyneoptera”. Geologic time scale is after Gradstein et al. (2004); the two, topmost intervals of time of the Neogene are, from
oldest to youngest, Pliocene and Pleistocene. See Appendix at www.geologica-acta.com, Table 1 and text for additional details.
functional feeding groups, external foliage feeding CO2 to recent CO2, presented in Berner and Kothavala
(Straus, 1977), leaf mining (Kozlov, 1988; Labandeira et (2001). These data and their relationship to the four herbi-
al., 1994), galling (Scott et al., 1994) and oviposition vore expansion phases are plotted in Fig. 8.
(Hellmund and Hellmund, 2002b) have the greatest
throughput and are present from the beginning of the Herbivore Expansion 1 occurs from the Late Silurian
angiosperm radiation. Curiously, piercing-and-sucking, to the Early Mississippian and is linked to a distinct com-
boring and seed predation are relatively delayed, collec- bination of primitive, terrestrial vascular plant and arthro-
tively appearing from the later Late Cretaceous to the pod clades. The earlier part of Phase 1 is associated with
middle Eocene. These patterns also occur on the unas- a greenhouse world that was gradually transformed mid-
signed plant hosts, which could represent a random sam- way to an icehouse world. Additionally, there are initially
ple of available plant hosts because of their uncertain and stable, high O2 levels (22.5% to 25%) during the Late Sil-
variable taxonomic affinities. urian and Early Devonian that subsequently plummet to
13% by the early Late Devonian, compared to present
The Dakota Fm, the “flagship” biota for the Herbivore value of 21%. Very elevated but highly fluctuating CO2
Expansion 4, spans the Early to Late Cretaceous bound- levels characterize Phase 1 during the early to middle
ary (late Albian to mid Cenomanian), and encompasses Devonian, but are followed by a sharp decline to modern
fluvial, deltaic and other terrestrial deposits bordering the levels during the Early Mississippian.
east and west sides of the midcontinental seaway that
extended from the Arctic Ocean to the Gulf of Mexico After an Early Mississippian hiatus occurring prior to
(Brenner et al., 2000; Wang, 2002). Sites from Kansas Herbivore Expansion 2 (Ward et al., in press), there is a
and Nebraska, U.S.A., of late Albian age (~103 Ma) rep- resurgence of herbivory during the Late Mississippian to the
resent coastal swamp, flood plain lake, and ox-bow chan- Permo Triassic boundary, similar to the previous phase by
nel deposits, and contain the earliest, highly diverse, occupying a strong greenhouse world. Moreover, Phase 2
abundant, and well preserved associations with occurs during a broad, unimodal distribution of elevated
angiosperms during their initial ecological radiation atmospheric O2 levels, starting with relatively depressed
across a variety of lowland environments (Doyle and concentrations (~ 17 %), reaching an unprecedented peak of
Hickey, 1976). The Dakota Formation probably has in 31% during the Early Permian (but see Lenton, 2001), and
excess of 300 species of vascular plants (Wang 2002) and decreasing to approximately current levels at the Permian-
many remain undescribed. Plant-insect associations from Triassic boundary. This decrease may explain major
this important deposit have only recently been investigat- turnover of Permian plant clades. Atmospheric CO2 was
ed (Stephenson, 1991; Scott et al., 1994; Labandeira et consistently low, comparable to modern levels throughout
al., 1994; Labandeira, 1998b), but contain the earliest the Carboniferous and Permian, but experienced a sharp rise
occurrences of leaf mines and among the earliest galls during the Triassic, in which there may have been a reorga-
and external foliage feeding of any early angiosperm flo- nization of the global carbon cycle, and thus resetting the
ra. There are virtually no insect body fossils. CO2 level at higher values (Berner, 2002).
Synchronicity of Herbivore Expansion Phases Herbivore Expansion 3, from the Middle Triassic to
with Environmental Change? the Recent, originates during an icehouse world and
evolves into the anomalous greenhouse world of the
An obvious issue is whether the temporal distribution of Jurassic to mid Cretaceous, in which there evidently is
the four herbivore expansion phases (Fig. 7) matches major latitudinal cooling but no polar icecaps. Atmospheric O2
cycles or otherwise aperiodic intervals of environmental content during Phase 3 reaches its lowest levels of the
change, such as paleoclimatologic shifts (Fig. 8). Three evi- past 420 m.y., fluctuating from 12.5 to 19% for Early Tri-
dent phenomena—greenhouse/icehouse cycles and associat- assic to Middle Jurassic, only to remain flat (17% to
ed global temperature and fluctuations in both atmospheric 19%) for about a 100 m.y. interval from the Late Jurassic
O2 and CO2 content—may have possible links with the four to mid Cretaceous, and rising during the past 50 m.y. to
phases of herbivore expansion. These possible relationships somewhat higher than present levels (23% to 21%). By
could be mediated by an effect on underlying vegetation as a contrast, atmospheric CO2 levels are elevated at ~ 5%
resource for arthropod herbivores. Data for establishing a throughout the Triassic (Berner, 2002), followed by a
greenhouse/icehouse world are from Frakes et al. (1992) and modest increase to higher levels of ~ 9% during the Mid-
global temperatures are from Scotese (2005). Atmospheric dle Jurassic, and thenceforth steadily declining to approx-
O2 levels are based on the sediment abundance model (Ber- imately the 0.035% preindustrial value of today.
ner et al., 2000, 2003; Falkowski et al., 2005; see also
Lenton, 2001). Atmospheric CO2 levels are inferred from the Herbivore Expansion 4 occurs from the mid Early
data of Cornette et al. (2002), based on the ratio of historical Cretaceous to the Recent. Unlike the initial part of Phase
3, Phase 4 commences during a greenhouse world, and DISCUSSION

similarly evolves into a subsequent icehouse world of the
Paleogene and Neogene. Unlike Phase 3, however, earlier This compilation and the resulting matrices depicting
Phase 4 associations occurred during somewhat depressed the ecological associations of host plants and their insect
but stable O2 levels (18% to 17%), peaking at 23% at 30 herbivores in the fossil record have provided some basic
Ma (early Oligocene), and then declining but remaining macroevolutionary patterns. One is the presence of sever-
high, reaching levels of 21% O2 today. Atmospheric CO2 al fixed and persistent functional feeding groups of
levels begin with levels of ~ 7% during the Early Creta- arthropod herbivores since they originated during the Late
ceous, followed by a steady decline to the preindustrial Paleozoic, or Triassic in the case of leaf mining. By con-
values of 0.035%. trast, the specific plant and arthropod occupants of these
associations are ephemeral in time and space. Member-
There is limited evidence that these three paleocli- ship volatility is attributable to clade turnover of plants
matologic indicators have a rational, predictive combi- and their herbivore arthropods, as well as associated fun-
nation of a greenhouse/icehouse switch that is linked to gi, and perhaps other interacting organisms. This turnover
values of atmospheric O2 and CO2 (Veizer et al., 2000). provides a dynamic context to the evolution of not only
However, it is not clear what the proximal cause is associations among specific biotas within each of the four
between the apparent synchrony of these three paleocli- herbivore expansions, but also among each of the four
matologic variables and each of the four herbivore major phases. Additionally, comparisons of the biogeog-
expansion phases (Fig. 8). For example, the role of pale- raphy of these associations within and among penecon-
ogeography remains unknown. It is known, though, that temporaneous floras of each phase may record the emer-
all phases except for Phase 3 began during periods of gence and evolutionary radiation of arthropod herbivore
globally warm temperatures. One possibility is that clades. A potential example of hemispherical extent
these atmospheric variables have direct physiologic con- would be Gondwanan floras occurring in the same paleo-
sequences on the selection and turnover of particular continental and similar basinal settings from Southern
plant clades globally, which in turn elicit an association- Africa, South America, Australia, India and Antarctica
al response from selected clades of insect herbivores. during the end of pteridophyte- and glossopterid dominated
The Early Cenozoic Thermal Maximum may be more a Herbivore Expansion 2, and the beginning of more diverse
temporally restricted example of this phenomenon (Wilf floras that were dominated by ginkgoopsid, voltzialean
et al., 2001). It remains untested whether a unique, com- conifer, cycadophyte and gnetopsid taxa of Herbivore
bined signal of these three (or perhaps other) paleocli- Expansion 3 (Fig. 7). It remains unknown whether similar
matologic variables is explainable by plant or insect plant hosts across the broad supercontinent of Gondwana
physiology (Lincoln et al., 1993; Graham et al., 1995), shared similar herbivores in distant basins within each of
and if there is a trigger which resulted in phases of these two phases, or if they evolved geographically distinc-
unique or augmented plant-insect associations. Modern tive suites of herbivores. A related issue is whether there is
studies from the experimental physiology of plants (Gra- persistence of taxonomically conservative lineages of plant
ham et al., 1995; Whittaker, 2001) and insects hosts and their insect herbivores through time, particularly
(Bartholomew and Casey, 1977; Loudon, 1988; Nicolas among the preangiospermous portion of Herbivore Expan-
and Sillans, 1989; Hagner-Holler et al., 2004) that are sion 3 to the Recent. Such associational longevity may
subjected to fluctuating levels of O 2 and CO 2 may be address whether some associations of today are truly
relevant. ancient in origin (Farrell, 1998; Powell et al., 1998).
FIGURE 7 Summary of herbivore expansion phases in Phanerozoic continental environments. This figure is a condensation of, from left to right,
plant host, plant-insect associational, and inferred insect herbivore patterns provided in figures 3 to 6. At left are major plant-host clades, based on
classifications from Kenrick and Crane (1997), Rothwell and Serbet (1994), Judd et al. (1999), Soltis and Soltis (2004), Pryer et al. (2004) and
Anderson and Anderson (2003). At right are the dominant arthropod herbivore clades that are based on classifications from Wheeler et al. (2004),
Coddington et al. (2004), and Willmann (2004). An exception is the major fungal clade, Prototaxales, at far left. At center are the four, major
expansion phases, based on associations from the literature, each of which has a “flagship” biota early in its development (Table 1). They are, from
bottom to top: the first is Herbivore Expansion 1, with an axis of the rhyniopsid Rhynia gwynne-vaughanii KIDSTON and LANG 1917 containing a stem
boring, from the Rhynie Chert (Early Devonian, Early Pragian) of Scotland (Kevan et al., 1975). Second is Herbivore Expansion 2, with a gall in the
rhachis tissues of the marattialean fern Psaronius chasei MORGAN 1959, from the Calhoun Coal (Late Pennsylvanian, Kasimovian) of the Illinois
Basin, Illinois, U.S.A. (Labandeira and Phillips, 2002). Third is Herbivore Expansion 3, with external foliage feeding on the matatiellalean ginkgoop-
sid Dejerseya lunensis (JOHNSTON) ANDERSON and ANDERSON 1989, from the Molteno Formation (Late Triassic, Carnian) of the Karoo Basin of South Africa
(Scott et al., 2004). Last and fourth is Herbivore Expansion 4, with a leaf mine on the protealean (platanoid) Sapindopsis beekeria WANG 2002 from
the Dakota Formation (Early Cretaceous, late Albian) of Kansas, U.S.A. (Labandeira et al., 1994; Wang, 2002). The absolute widths of each bubble
do not necessarily conform to the absolute diversity of associations, but the relative distributions of widths among the four phases indicate relative
importances. At right are major groups of interacting arthropod herbivores, namely clusters of clades that consist of myriapods, mites, and several
insect lineages (Labandeira, 1994; Rasnitsyn and Quicke, 2002). Geologic time scale after Gradstein et al. (2004); the two, topmost intervals of
time of the Neogene are, from oldest to youngest, Pliocene and Pleistocene.
One observation that contextualizes this study involves Those that may have survived from Herbivore Expansion 3
the overwhelming focus by biologists on associations occur on conifers, cycads and possibly gnetaleans (Burdick,
between angiosperms and holometabolous insects. This sin- 1961; Norstog, 1987; Kato and Inoue, 1994) that have geo-
gular emphasis involves documentation of numerous associa- graphically unique or relictual distributions. Apparently, there
tions and co-radiations between these currently hyperdiverse are no extant associations from herbivore Expansions 1 and 2.
clades (Grimaldi, 1999; Grimaldi and Engel, 2005). This This “angiocentrism” has clouded our understanding of the
predilection is reasonable, as the overwhelming majority of deeper history of plant-insect associations by limiting our under-
extant associations available for study involve angiosperms standing of how extinct host-plants and their insect herbivores
and holometabolous insects from Herbivore Expansion 4. have associated at deeper macroevolutionary timescales.
FIGURE 8 The relationship of Late Silurian to Recent physical variables—atmospheric O2 and CO2 content, and average global temperatures—with
that of the four herbivore expansion phases, as determined by the fossil history of plant-insect associations. Atmospheric CO2 ratios are from Berner
and Kothavala (2001); atmospheric O2 percentages from Falkowski et al. (2005) and Ward et al. (in press); average global temperature from Scotese
(2005); and greenhouse/icehouse cycles are from Frakes et al. (1992).
Current ecological studies of plant-arthropod associa- biota of exceptional associational richness that establishes
tions thus focus overwhelmingly on plant-host taxa that are most of the broad limits for subsequent associations and
crown spermatophytes (angiosperms) and on insect herbi- their modifications. This early expression, represented by a
vore taxa that are clades of crown-group Holometabola “flagship” biota, can be considered ecologically either as a
(“phytophagan” beetles, cyclorrhaphan flies, ditrysian rebound from an earlier event characterized by the demise of
moths, and aculeate hymenopterans) and to a lesser extent the previous herbivore expansion phase, or alternatively the
Hemipteroidea (especially sternorrhynchans and phy- origination of a fundamentally new major clade of host
tophagous bugs such as Lygaeidae, Coreidae, and Miridae; plants and insect herbivores without linkage to a causative
Fig. 7). Paleobiologically, this is a rather atypical situation, environmental perturbation.
historically confined to the last fourth of the history of
plant-arthropod associations. A retreat to Herbivore Expan- 3. There are qualitative links between each of the her-
sion 3 would reveal taxa in which the crown spermato- bivore expansion phases and three paleoclimatologic vari-
phytes of the time were various gymnospermous clades ables (greenhouse or icehouse world and atmospheric O2
(several lineages of ginkoopsids, voltzialean and cheirolep- and CO2 levels). However, no consistent or predictive pat-
idaceous conifers, peltaspermaleans, pentoxylaleans, ben- tern emerges that can explain the specific relationship
nettitaleans) and their arthropod herbivores were more between these paleoclimatologic variables and a particu-
basally positioned clades of holometabolous insects lar herbivore expansion phase.
(polyphagan coleopterans, brachyceran flies, aneuretopsy-
chid mecopterans, micropterygid to monotrysian lepi- 4. Associations persist through geologic time whereas
dopterans, symphytan hymenopterans) and hemipteroids the occupants of those associations are spatiotemporally
(thysanopterans, lophioneurids, sternorrhynchans, auchen- constrained. This suggests that the several modes of
orrhynchans). Would the partitioning and use of plant-host arthropod feeding on vascular plant tissues are fixed, con-
taxa be the same as those for Herbivore Expansion 4? strained possibly by plant tissue architecture and arthro-
Retreat yet one more phase to that of Herbivore Expansion pod mouthpart structure, and thus displays extensive taxo-
2, and the landscape of host plants then consisted of pteri- nomic and functional convergence.
dophytes (especially marattialean ferns) and stem sper-
matophytes (dominantly medullosan and glossopterid 5. An “angiocentric” focus on Herbivore Expansion 4
pteridosperms and cordaitalean conifers), and the dominant (angiosperms vs. crown holometabolans and hemipteroids)
phytophagous clades of arthropod herbivores were mites, has had the effect of de-emphasizing patterns and processes
apterygotes, paleodictyopteroids and a diverse assemblage of plant-arthropod associations during the preceding 300
of primitive “protorthopteroid” insects. How would the her- million years. In particular, more understanding of herbivore
bivore component community on a host plant dominant within expansions 1 through 3 is needed to comprehend the full
Phase 2 compare ecologically with a component community breadth of how vascular plants and their arthropod herbi-
of today’s Phase 4? More specifically, could the varied associ- vores historically have monopolized life on land.
ations seen among the Herbivore Expansion 4 be an ecologi-
cally similar or repackaged version of those from Herbivore
expansions 2 or 3, but with different players? ACKNOWLEDGEMENTS
I thank José Luis Sanz, Xavier Delclòs, and Carmen Soriano

CONCLUSIONS for organizing and inviting me to the Eighth Course of Paleontol-
ogy in Cuenca, Spain. This contribution is an extension of a presen-
Although this study is only a first attempt to under- tation given at the Cuenca symposium. I am grateful to Finnegan
stand the broad patterns of plant-arthropod associations in Marsh for rendering the figures. I am also indebted to the Smith-
deep time, five conclusions are made. These conclusions sonian Institution Libraries for assistance in procuring primary liter-
can be considered as hypotheses that should engender ature that were difficult to obtain. William DiMichele and two
additional analyses and testing though the gathering of anonymous reviewers provided valuable reviews and feedback.
additional data at finer spatiotemporal scales. This is contribution no. 126 of the Evolution of Terrestrial Ecosys-
tems Consortium at the National Museum of Natural History.
1. The fossil record of plant-insect associations can be
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Wesenberg-Lund, C., 1913. Fortpflanzungsverhältnisse: Paarung etatis Scientarum Torunensis, 4, 43-48.
un Eiablage der Süßwasserinsekten. Fortschritte der natur- Zherikhin, V.V., 2002a. Insect trace fossils. In: Rasnitsyn, A.P., Quicke,
wassenschaftenlichen Forschung, 8, 161-286. D.L.J. (eds.). History of Insects. Dordrecht, Kluwer, 303-324.
Wheeler, W.C., Giribet, G., Edgecombe, G.D., 2004. Arthropod Zherikhin, V.V., 2002b. Ecological history of the terrestrial
systematics. In: Cracraft, J., Donoghue, M.J. (eds.). Assembling insects. In: Rasnitsyn, A.P., Quicke, D.L.J. (eds.). History of
the Tree of Life. New York, Oxford University Press, 281-295. Insects. Dordrecht, Kluwer, 331-388.
Whittaker, J.B., 2001. Insects and plants in a changing atmos- Zhou, Z., Zhang, B., 1989. A sideritic Protocupressinoxylon with
phere. Journal of Ecology, 89, 507-518. insect borings and frass from the Middle Jurassic, Henan,
Wieland, G.R., 1906. American Fossil Cycads. Washington, China. Review of Palaeobotany and Palynology, 59, 133-143.
Manuscript received June 2005;

revision accepted May 2006.
APPENDIX
Register of biotas for Figures 3 to 6
Herbivore Expansion 1
Biota Locality Stratigraphic placement Time interval Major references
1 Ludford Lane, Perton Downton Castle Limestone Silurian, Přídolí Edwards et al.
Lane, Welsh Borderland 1995; Edwards
U.K. 1996
2 Brown Clee Hill, Welsh Old Red Sandstone, micr- Devonian, Edwards et al.
Borderland, Wales, U.K. ornatus-newportensis Spore Lochkovian 1995; Edwards
Biozone 1996
3 Rhynie Chert, Aberdeen- Dryden Flags Formation Devonian, Kidston and Lang,
shore, Scotland, U.K. Pragian 1921; Kevan et al.
1975; Shear and Selden
2001; Habgood et al. 2004
4 Gaspé, Quebec, Canada Battery Point Formation Devonian, Trant and Gensel 1985;
Emsian Banks and Colthart
1993; Hotton et al.
1996; Hueber 2001;
Labandeira, pers. observ
5 Kettle Point, Ontario, Huron Shale Devonian, Arnold 1952; Hueber
Canada Famennian 2001
1 Sydney Basin, New McInnes Formation Mississippian, Iannuzzi and Laban-

South Wales, Australia earliest deira, unpubli. data
Serpukhovian
2 White River, Washing- Fayetteville Formation Mississippian, Dunn et al. 2003
ton Co., Arkansas, U.S.A. Serpukhovian
3 Jackson Co., Illinois, Caseyville Formation Pennsylvanian, Jennings 1974
U.S.A. Bashkirian
4 Northumberland, Coal Measures, below the Pennsylvanian, Thomas 1969;
England, U.K. Ashlington Marine Band late Bashkirian to Amerom 1973
early Moscovian
5 North-central England, Coal Measures, below and Pennsylvanian, Holden 1910, 1930;
U.K. above the Catharina Seam Moscovian Chaloner et al. 1991
6 Méricourt, Arras, France “Assise de Bruay” unit Pennsylvanian, Amerom and Boersma
middle Moscovian 1971
7 Zwickau, Germany “Hellgrauer Schieferton” Pennsylvanian, Müller 1982
late Moscovian
8 Saline and Gallatin Cos., Carbondale Formation, Pennsylvanian, Labandeira and Phillips
Illinois, U.S.A. (Herrin Coal) late Moscovian 2002
9 Mazon Creek, north- Carbondale Formation, Pennsylvanian, Scott and Taylor 1983;
central Illinois, U.S.A. (Francis Creek Shale) late Moscovian Labandeira and Beall
1990; Scott et al. 1992
10 Berryville, Lawrence Mattoon Formation Pennsylvanian, Lesnikowska 1990;

Co., Illinois, U.S.A. (Calhoun Coal) Kasimovian Labandeira and Phillips
1996a, 1996b, 2002;
Roßler 2000; Laban-
deira, pers. observ.
11 La Magdalena Coalfield, [formation not reported] Pennsylvanian, Amerom 1966; Castro
León, Spain Gzehlian 1997
12 Graissessac, Hérault, “…between the Grand Pas Pennsylvanian Béthoux et al. 2004
France and Burelle Coal Seams…” late Gzhelian
13 Saar-Nahe Basin, Lower Rotliegende Carboniferous- Florin 1945; Meyen
Germany Permian boundary 1984; Kerp 1988;
Zherikhin, 2002a
14 Chunya, Siberia Russia [formation not reported] latest Carboni- Sharov 1973; Zherikhin
ferous 2002a
15 Pingquan, Hebei Taiyuan Formation Permian, Hilton et al. 2001, 2002
Province, China Asselian
16 Faxinal and Morro do Rio Bonito Formation Permian, Artin- Guerra-Sommer 1995;
Papaléo, Rio Grande do skian/Kungurian Adami-Rodrigues et al.
Sul, Brazil 2004a, 2004b
17 Coprolite Bone Bed, Nocona Formation Permian, Greenfest and Laban-
Archer Co., Texas, Sakmarian deira 1997
U.S.A.
18 Chemnitz, Erzgebirge Leukersdorf Formation, Lower Permian Roßler1996
Basin, Sachsen, Germany “Zeisgwald-Tuff-Horizontes”
19 Gaines Co., Texas, Clear Fork Group Permian, late DiMichele et al. 2000
U.S.A. Artinskian
20 Ranigang Coalfield, Barakar Formation Permian, Artins- Srivastava 1987
West Bengal, India kian/Kungurian
21 Taint, Baylor Co., Waggoner Ranch Formation Permian, Beck and Labandeira
Texas, U.S.A. Artinskian 1998
22 Chubut, Argentina Río Genoa Formation Lower Permian Cúneo 1987
23 Minas do Leão, Rio Irati and Serra Alta Permian, Kun- Adami-Rodrigues and
Grande do Sul, Brazil Formations gurian/Roadian Iannuzzi 2001; Adami-
Rodrigues et al. 2004a
24 Taiyuan City, Shanxi, Shihhotse Formation Permian, Kungurian Glasspool et al. 2003
Province, China /Roadian
Boundary
25 King Co., Texas, U.S.A. Blaine Formation Permian, DiMichele et al. 2004
?Roadian
26 Vereeniging, Gauteng, Vryheid Formation Permian, Plumstead 1963;
South Africa ?Roadian Amerom 1966
27 Cooyal, Sydney Basin, Upper Illawarra Coal Late Permian Holmes 1995
Australia Measures
28 Ranigang Coalfield, Ranigang Formation Late Permian Srivastava 1996;
Banerjee and Bera 1998
29 Northern Prince Charles Bainmedart Coal Measures Late Permian Weaver et al. 1997
Mountains, East
Antarctica
30 Bowen Basin, Queens- Rangal Coal Measures, Permian, McLoughlin 1994a,
land, Australia Bandanna Formation Wuchiapingian 1994b
31 Clouston Farm, Kwa- Estcourt Formation Permian, Labandeira and Prevec,
Zulu-Natal, South Africa Changhsingian unpubl. data
1 Grès-à-Voltzia, northern Bundsandstein Formation Triassic, Grauvogel-Stamm and

Vosges Mtns., France Lower Anisian Kelber 1996
2 Wasselonne, Bas-Rhine, Lettenkohle and Lower Triassic, Linck 1949; Roselt
France; Schleerieth and Keuper Formations Upper Ladinian 1954; Geyer and Kelber
Ochsenfurt, Franconia, 1987; Kelber 1988;
Germany Grauvogel-Stamm and
Kelber 1996
3 Karoo Basin: KwaZulu- Molteno Formation Triassic, Anderson and Anderson
Natal, Eastern Cape and Carnian 1989; Scott et al. 2004;
Northern Cape Provinces, Labandeira and Ander-
South Africa son 2005
4 Sydney Basin, New Blackstone Formation Triassic, Tillyard 1922; Roze-
South Wales, Australia Carnian felds and Sobbe 1987
5 Petrified Forest National Chinle Formation Triassic, Walker 1938; Ash 1997,
Park, Arizona, U.S.A. Norian 1999, 2000, 2005; Ash
Savidge 2004; Creber
and Ash 2004
6 Xiniguá, Rio Grande Santa Maria Formation Triassic, Minello 1994
do Sul, Brazil Norian
7 Küfner, Bavaria, “Pflanzensandstein” Jurassic, Weber 1968; Van Kon-
Germany Hettangian ijnenburg-Van
Cittert and Schmeißner 1999
8 Yima, Henan Province, Yima Formation Jurassic, Toarcian Zhou and Zhang 1989
China to Aalenian
9 Murlipahar, Bihar, Dubrajpur Formation Jurassic, Vishnu-Mittre 1957;
India Aalenian Sharma and
Harsh 1989
10 Yorkshire, northern Scarborough Formation Jurassic, Harris 1942; Alvin et al.
England, U.K. Bajocian 1967; Scott et al. 1992
11 Jaramillo, Santa Cruz La Matilde Formation Jurassic, Callovian Genise and Hazeldine
Province, Argentina to Oxfordian 1995
12 Mikhailovka, Chayan Karabatsu Formation Jurassic, Rasnitsyn and Krassilov
District, Kazakhstan Kimmeridgian 2000
13 Steiner, Mt. Ellen, and Morrison Formation Jurassic Tidwell and Ash 1990;
East McElmo Creek Tithonian Tidwell and Medlyn
localities, Western 1992; Dayvault and
Interior, U.S.A. Hatch 2003
14 Clack Island, northern Battle Camp Formation Latest Jurassic to Rozefelds 1988a
Queensland, Australia earliest Cretaceous
15 Crowborough, East Waldhurst Clay or Cretaceous, late Jarzembowski 1990
Sussex, England, U.K. Ashdown Formation Berriasian to mid-
dle Valanginian
16 Przenosza, Limanowa ?Verovice Shale Cretaceous, Reymanówna 1960,
District, Poland ?Barremian 1991
17 Makhtesh Ramon, Hatira Formation Lower Aptian Krassilov et al. 2004b
Negev, Israel
18 Chunakhal and Hiran- Rajmahal Formation Cretaceous, Banerji 2004
duba, Jharkhand State, (Intertrappean Beds) Aptian/Albian
India boundary
19 Amarjola, Bihar State, [formation not reported] Lower Cretaceous Bose 1968
India
20 Glen Rose, Somervell Glen Rose Formation Aptian/Albian Watson 1977

Co., Texas boundary
21 Yuleba, Queensland, Mooga Sandstone Lower Cretaceous Tidwell and Rozefelds
Australia 1991
22 Hokodz, northwestern Diadochoceras nodosocos- Cretaceous, upper Falder et al. 1998;
Caucasus Region, Russia tatum and Acanthoplites Albian Ratzel et al. 2001
bigoureti Zone
23 Blackhawk, Black Hills “Sandstones” Lower Cretaceous Wieland 1906; Dele-
South Dakota, U.S.A. voryas 1968; Crepet
1974; Crowson 1976
24 Valcheta, Rio Negro, [formation not reported] Late Cretaceous Andreis et al. 1991;
Argentina Genise 1995
25 Nammoura, Mont-Liban “Namoura Plattenkalk” Cretaceous, Krassilov and Bacchia
District, Lebanon Cenomanian 2000
26 Shuparo Lake, Hokkaido Yezo Group Cretaceous, Ceno- Saiki and Yoshida 1999
Japan manian to early
Santonian
27 Ohyubari, Hokkaido Yezo Group Cretaceous, Late Nishida and Hayashi
Japan Turonian to Early 1996
Campanian
28 Brannen Lake, Vancou- Haslam Formation Cretaceous, lower Stockey and Rothwell
ver Island, British Campanian 2003
Columbia, Canada
29 Big Cedar Ridge, Meeteetsee Formation Cretaceous, early Wing et al. 1993;
Wyoming, U.S.A. Maastrichtian Labandeira et al. 1995
30 Williston Basin, Hell Creek Formation Cretaceous, late Labandeira et al. 2002a,
southwestern North Maastrichtian 2002b
Dakota, U.S.A.
31 Republic, Ferry Co., Klondike Mountain Paleogene, Labandeira, pers.
Washington, U.S.A. Formation Ypresian observ.
32 Baltic amber; Baltic Prussian Formation Paleogene, Conwenz 1890
Region, northern Europe Lutetian
33 Geodetic Hills, Axel Buchanan Lake Formation Paleogene, Labandeira et al. 2001
Heiberg Island, Nunavut, Lutetian
Canada
34 Dynów, Skole Nappe, Kliwa Sandstone Paleogene, Rajchel and Uchman
Outer Carpathians, Rupelian 1998
Poland
35 Klepzig, Germany “lignite” Paleogene, Linstow 1906
middle Oligocene
36 Rott, Schwaben, “porcellanite” Paleogene, Hellmund and Hell-
Germany Chattian mund 1991
37 Freilendorf, Hessen, “lignite” Paleogene Roselt and Feustel
Germany Chattian 1960
38 Southern Limburg, [formation not reported] Neogene, late Süss 1979, 1980
Netherlands Miocene
39 Szentgal, Hungary “lignite” Neogene, Dudich 1961
Tortonian
40 Konin, Poland “lignite” Neogene, Madziara-Borusiewicz
Tortonian 1970
41 Düren, Nordrhein- “lignite” Neogene, ?early Möhn 1960
Westfalien, Germany Miocene
42 Bohemia, Czech [formation not reported] Neogene, ?early Engelhardt 1876
Republic Miocene
43 Wieliczka, Poland Salzstocke Formation Neogene, ?early Zablocki 1960

Miocene
44 Clarkia, Latah Co., Latah Formation Neogene, middle Lewis 1985, Lewis et al.
Idaho, U.S.A. Miocene 1990b
45 Emerald Creek 1, Bene- Latah Formation Neogene, middle Lewis et al. 1990b
wah Co., Idaho, U.S.A. Miocene
46 Emerald Creek 2, Sho- Latah Formation Neogene, middle Lewis et al. 1990b
shone Co., Idaho, U.S.A. Miocene
47 Oviatt Creek, Clearwater Latah Formation Neogene, middle Lewis et al. 1990b
Co., Idaho, U.S.A. Miocene
48 Juliaetta, Nez Perce Latah Formation Neogene, middle Lewis et al. 1990a
Co., Idaho, U.S.A. Miocene
49 Jasper Creek, Yellow- [formation not reported] Neogene, Brues 1936
stone National Park, Messinian
Wyoming, U.S.A.
50 Guntramsdorf, Austria “Congeriensand” Neogene, lower Abel 1933
Pliocene
51 Königsbruck, Alsace, [formation not reported] Neogene, middle Geissert et al. 1981
France to upper Pliocene
52 Peary Land, northern Kap København Formation Neogene, Plio- Böcher 1995
Greenland, Denmark cene/Pleistocene
Boundary
53 Scarborough, Ontario, interglacial clays Neogene, Scudder 1900
Canada Pleistocene
54 Washington, District unconsolidated sediment Neogene, Berry 1924; Gagné
of Columbia, U.S.A. Pleistocene 1968
55 Santa Cruz Island, Santa Cruz Island Formation Neogene Chaney and Mason
California, U.S.A. (alluvial deposits) Pleistocene 1934
56 Near Fairbanks, Alaska, Goldstream Formation Neogene, late Péwé et al. 1997
U.S.A. Pleistocene
1 Kiowa Co., Kansas, Cheyenne Formation Cretaceous, lower Stephenson 1991

U.S.A. to middle Albian
2 Stump Neck, Maryland, Patapsco Formation Cretaceous, Hickey and Doyle 1977;
U.S.A. middle to upper Larew, 1992
Albian
3 Quantico, Virginia, Potomac Group, unassigned Cretaceous, Upchurch et al. 1994
U.S.A. middle to upper
Albian
4 Rose Creek, Hoisington, Dakota Formation Cretaceous, late Stephenson 1991;
Linnenberger Ranch, and Albian Labandeira et al. 1994;
Acme localities, Kansas Labandeira 1998b;
and Nebraska, U.S.A. unpubl. data
5 Cumberland, Lincoln Frontier Formation (lower) Cretaceous, early Stephenson 1991
Co., Wyoming, U.S.A. Cenomanian
6 Tumbler Ridge, east- Dunvegan Formation Cretaceous, early Crabtree 1987
central British Cenomanian
Columbia, Canada
7 Vyserovice, Lidice, Perucer Formation Cretaceous, Frič 1882, 1901;

Kounice and other middle Stephenson 1991
localities, Bohemia, Cenomanian
Czech Republic
8 Karatau, Kzyl Dzhar Beleutinskaya Formation Cretaceous, Kozlov 1988
Region, Kazakhstan Turonian
9 Gerofit, Negev, Israel Gerofit Formation Cretaceous, Krassilov et al., 2004a
middle Turonian
10 Cumberland, Lincoln Frontier Formation (upper) Cretaceous, Knowlton 1917;
Co., Wyoming, U.S.A. late Turonian Crabtree 1987
11 “Oilfield Coulee”, Madi- Two Medicine Formation Cretaceous, lower- Crabtree 1987
son, Co., Montana, most Campanian
U.S.A.
12 Gaillard, Crawford Gaillard Formation Cretaceous, late Lupia et al. 2002
Co., Georgia, U.S.A. Santonian
13 Big Cedar Ridge, north- Meeteetsee Formation Cretaceous, early Wing et al. 1993;
west Wyoming, U.S.A. Maastrichtian Labandeira, pers.
observ.
14 Bisti Badlands, New Fruitland Formation Cretaceous, late Tidwell et al. 1981
Mexico, U.S.A. Campanian
15 Valcheta, Rio Negro, [formation not reported] Late Cretaceous Andreis et al. 1991;
Argentina Genise 1995
16 Williston Basin, south- Hell Creek Formation Cretaceous, late Wilf et al. 2000; Laban-
western North Dakota, Maastrichtian deira et al. 2002a, 2002b
U.S.A.
17 Williston Basin, south- Fort Union Formation Paleogene, Labandeira et al. 2002a,
western North Dakota, (lower) Danian 2002b
U.S.A.
18 Various localities, north- Fort Union Formation Paleogene, Wilf and Labandeira
ern Western Interior, (upper) Selandrian to 1999; Wilf et al. 2001
Wyoming, Montana, Thanetian
Utah, U.S.A.
19 Cold Ash, Berkshire, Woolwich and Reading Paleogene, Crane and Jarzem-
U.K. Beds Thanetian bowski 1980
20 Damalgiri, Meghalaya Tura Formation Paleogene, Srivastava et al. 2000
State, India Thanetian
21 Almont, North Dakota, Sentinel Butte Formation Paleogene, Crane et al. 1990
U.S.A. Thanetian
22 Foster Gulch, Carbon Fort Union Formation Paleogene, early Lang 1996
Co., Montana, U.S.A. (upper) Ypresian
23 Puryear, Henry Co., Wilcox Formation Paleogene, Berry 1923; Brooks
Tennessee, U.S.A. Ypresian 1955; Wittlake 1969
24 Sourdough Flora, Sweet- Wasatch Formation Paleogene Wilf et al. 2000, 2001,
water Co., Wyoming, Ypresian 2005
U.S.A.
25 Laguna del Hunco, La Huitrera Formation Paleogene, Wilf et al. 2005
Chubut Province, Ypresian
Argentina
26 Dubois, Park Co., Wind River Formation Paleogene, late Hickey and Hodges
Wyoming, U.S.A. Ypresian 1975
27 Bonanza, Uintah Co., Green River Formation Paleogene, Wilf and Labandeira
Utah, U.S.A. Lutetian 1999; Wilf et al. 2001,
2005
28 Darmstadt, Hessen, Messel Formation Paleogene, Kinzelbach 1970; Straus

Germany Lutetian 1976; Barthel and
Rüffle 1976; Collinson
1990; Schaarschmidt
1992; Hellmund and
Hellmund 1998
29 Geiseltal, Halle, Mittelkohle Formation Paleogene, Mai 1976
Sachsen-Anhalt, Lutetian
Germany
30 Republic, Ferry Co., Klondike Mountain Paleogene, Wolfe and Wehr 1987;
Washington, U.S.A. Formation Lutetian Lewis 1992; Lewis and
Carroll 1991; Pigg et al.
2001; Labandeira 2002b
31 Bonanza, Uintah Co., Green River Formation Paleogene, Wilf et al. 2001, 2005
Utah, U.S.A. Lutetian
32 Eckfeld, Manderscheid, Maar Formation Paleogene, Wilde and Franken-
Rheinland-Pfalz, Lutetian häuser 1998
Germany
33 Baltic amber; Baltic Prussian Formation Paleogene, Wappler and Engel
Region, northern Europe Lutetian 2003
34 Prineville, Crook Co., Clarno Formation Paleogene, Chaney 1927; Gregory
Oregon, U.S.A. Rupelian 1969
35 Holley, Crook Co., Eugene Formation Paleogene, Gregory 1968
Oregon, U.S.A. Bartonian
36 Bournemouth, East Branksome Sand Formation Paleogene, Stephenson 1991;
Dorset, England, U.K. (including Bournemouth Bartonian Lang et al. 1995
Fresh Water Beds)
37 White Lake Basin, [unreported formation] Paleogene, Freeman 1965
British Columbia, Canada Bartonian
38 Kennedy Range, West Merlinleigh Sandstone Paleogene, McNamara and
Australia, Australia ?Priabonian Scott 1983
39 Amethyst Mountain, [formation not reported] Paleogene, Süss and Müller-
Park Co., Wyoming, ?Priabonian Stoll 1980
U.S.A.
40 Comstock, Douglas Co., Comstock Formation Paleogene, Sanborn 1935
Oregon, U.S.A. Priabonian
41 Florissant, Park Co., Florissant Formation Paleogene, Cockerell 1908a, 1908b,
Colorado, U.S.A. Priabonian 1910; Opler 1982;
Meyer 2003
42 Anglesea, Victoria, [formation not reported] Paleogene, Rozefelds 1988b
Australia Priabonian
43 Vargem, Grande do Sul, Tremembé Formation Paleogene, Martins-Neto 1989,
São Paulo, Brazil Rupelian 1998
44 La Porte, Plumas Co., ?Ione Formation Paleogene, Potbury 1935
California, U.S.A. Rupelian
45 Seifhennersdorf, “Kuclin” diatomite Paleogene, middle Hellmund and Hell-
Sachsen, Germany Oligocene mund 1996
46 Wind brickyard, [formation not reported] Paleogene, Ambrus and Hably 1979
Baromállás, Hungary Chattian
47 Ruby River, Madison Renova Formation Paleogene, Becker 1965, 1969;
Co., Montana, U.S.A. Chattian Lewis 1976
48 Rott, Schwaben, Bayern, Köln Formation Paleogene, Heyden 1862; Sittig
Germany Chattian 1927; Hellmund and
Hellmund 1991
49 Simojovel, Chiapas, Baluntun Formation Neogene, Poinar and Brown 2002

Mexico Aquitanian
50 Most, Czech Republic “Main Brown Coal Seam” Neogene, Mikuláš et al. 1998
Aquitanian
51 Düren, Nordrhein- “Flöz Garzweiler” Paleogene, Schmidt et al. 1958
Westfalien, Germany Aquitanian
52 Mainz and Petersburg, Hydrobia Limestone Neogene, Schmidtgen 1938
Rheinland-Pfalz, Aquitanian
Germany
53 Osieczow, Poland “Quadersandstein formation” Neogene, Karpiński 1962;
Aquitanian Radwanski 1977
54 Oeningen, Baden, “Stinkschiefer” Neogene, ?early Heer 1853; Mädler 1936
Switzerland Burdigalian
55 Berzdorf (Sachsen), Toneisensteinblock Neogene Hellmund and Hell-
Germany Formation Burdigalian mund 2002a, 2002b
56 Ribesalbes, Castellón, [formation not reported] Neogene, Peñalver and Martínez-

Spain Lower Miocene Delclòs 1997; Peñalver
and Martínez-Delclòs
2004
56a Bellver de Cerdaña [formation not reported] Neogene, Martínez-Delclòs 1996
Upper Miocene
57 Rubielos de Mora, [formation not reported] Neogene, Peñalver and Martínez-
Teruel, Spain Lower Miocene Delclòs 1997
58 Seed site 64, Sioux Co., Sheep Creek Formation Neogene, Thomasson 1982
Nebraska, U.S.A. Burdigalian
59 Salhausen, Hessen, Braunkohle Formation Neogene, late Hellmund and Hell-
Germany Burdigalian mund 2002a, 2002b
60 Randecker Maar, Stutt- Dysodil Formation Neogene, late Hering 1930; Hellmund
gart, Baden-Württemburg, Burdigalian and Hellmund 2002b
Germany
61 Stinking Water Harney Astoria Formation Neogene, Opler 1973; Waggoner
Co., Oregon, U.S.A. Langhian 1999
62 Gilliam Springs, Washoe [formation not reported] Neogene, early Waggoner and Poteet
Co., Nevada, U.S.A. Langhian 1996
63 Linqu, Shandong, Shanwang Formation Neogene, Guo 1991
China Langhian or
Serravallian
64 Upper Goldyke, Cedar [formation not reported] Neogene, Opler 1973
Mtns., Nevada, U.S.A. Langhian
65 Trout Creek, Oregon, Buffalo Canyon Formation Neogene, Opler 1973
Stewart Valley and and others of similar age Langhian
Buffalo Canyon,
Nevada, U.S.A.
66 Clarkia, Shoshone Co., Latah Formation Neogene, Knowlton, 1926; Opler
Idaho, U.S.A. Tortonian 1973; Lewis 1985
67 Albertine Rift Valley, Nkondo Formation Neogene, Nel 1994
Uganda Tortonian
68 Aldrich Station, [formation not reported] Neogene, late Opler 1973
Nevada, U.S.A. Serravallian
69 Southern Idaho, U.S.A. Trapper Creek Formation Neogene, Liebhold et al. 1982
Tortonian
70 Wüstensachsen, Hesse, Lettengraben Formation Neogene Müller-Stoll 1989
Germany Tortonian
71 Table Mountain, Tuo- Mehrten Formation Neogene, Condit 1944

lumne Co., California, Tortonian
U.S.A.
72 Paris Basin, France [formation not reported] Neogene, late Brues 1936
Miocene
73 La Cerdaña, Lérida, “Unit C of lower Neogene Neogene, late Diéguez et al. 1996;
Spain unit” Miocene Barrón et al. 1999
74 Nagano and Gumma “Kabutiowa Plant Bed” Neogene, late Kuroko 1987
Prefectures, Japan Miocene
75 Douglas Co., Washing- [formation not reported] Neogene, late Hoffman 1932
ton, U.S.A. Miocene
76 Egweil, Württemburg, “Sußwassermolasse” Neogene, late Selmeier 1984
Germany Miocene
77 Mikófalva, Hungary [formation not reported] Neogene, Süss and Müller-
Messinian Stoll 1975
78 Willershausen, Thuringia, “lacustrine clay sediments” Neogene, early Kernbach 1967; Heie
Germany Pliocene 1967; Steinbach 1967;
Straus 1967, 1977
79 Chuizbaia, Maramure,s Diatomit-Schiefer Neogene, early Givulescu 1984
northwestern Romania Pliocene
80 Brun-Vösendorf and [formation not reported] Neogene, early Berger 1949, 1953
Laaer Berg, Austria Pliocene Straus 1967, 1977
81 Neu-Isenburg, Hessen, “Tonscholle” Neogene, Rietschel 1983
Germany Pliocene
82 Wetterau, Hesse, “lignite” Neogene, Greiling and Schneider
Germany Pliocene 1973
83 Santa Rosa, Sonoma Sonoma Formation Neogene, middle Axelrod 1944
Co., California, U.S.A. Pleistocene
84 Vitošov, Czech Republic “karst breccia” Neogene, Mikuláš et al. 1998
Pleistocene
85 Beceite, Teruel, Spain travertine deposits Neogene, Peñalver et al. 2002
Pleistocene
86 Königsburg, Alsace, “lignite” Neogene, middle Geissert et al. 1981
France to late Pliocene
87 Birtley, Durham, unconsolidated peat Neogene, late Heslop-Harrison 1926
England, U.K. Pleistocene
88 Rancho La Brea, Los asphaltum Neogene, late Larew 1987
Angeles Co., California, Pleistocene
U.S.A.

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The Four Phases of Plant-Arthropod Associations in Deep Time

1 Smithsonian Institution, National Museum of Natural History

2 University of Maryland, Department of Entomology

KEYWORDS Plant-insect-associations. Herbivory Fossil-record. Land plants. Insects. Paleoclimate.

INTRODUCTION vided the basic evolutionary history and ecological struc-

Herbivore Major Number

1. 420–360 Ma protracheo- myriapods, Rhynie 3 10 5 5

2. 335–251 Ma pteridophytes, myriapods, Calhoun 7 57 31 14

3. 245 Ma–present pteridophytes, mites, Molteno 8 95 56 33

4. 115 Ma–present angiosperms, orthopteroids, Dakota 8 263 88 22

Diaphanopterodea and Dicliptera (Grimaldi and Engel, 2005).

FIGURE 3 Herbivore Expansion 1. Illustrated is

3, Phase 4 commences during a greenhouse world, and DISCUSSION

I thank José Luis Sanz, Xavier Delclòs, and Carmen Soriano

Manuscript received June 2005;

Biota Locality Stratigraphic placement Time interval Major references

Biota Locality Stratigraphic placement Time interval Major references

1 Sydney Basin, New McInnes Formation Mississippian, Iannuzzi and Laban-

10 Berryville, Lawrence Mattoon Formation Pennsylvanian, Lesnikowska 1990;

Biota Locality Stratigraphic placement Time interval Major references

1 Grès-à-Voltzia, northern Bundsandstein Formation Triassic, Grauvogel-Stamm and

20 Glen Rose, Somervell Glen Rose Formation Aptian/Albian Watson 1977

43 Wieliczka, Poland Salzstocke Formation Neogene, ?early Zablocki 1960

Biota Locality Stratigraphic placement Time interval Major references

1 Kiowa Co., Kansas, Cheyenne Formation Cretaceous, lower Stephenson 1991

7 Vyserovice, Lidice, Perucer Formation Cretaceous, Frič 1882, 1901;

28 Darmstadt, Hessen, Messel Formation Paleogene, Kinzelbach 1970; Straus

49 Simojovel, Chiapas, Baluntun Formation Neogene, Poinar and Brown 2002

56 Ribesalbes, Castellón, [formation not reported] Neogene, Peñalver and Martínez-

71 Table Mountain, Tuo- Mehrten Formation Neogene, Condit 1944

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