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JOURNAL OF CRUSTACEAN BIOLOGY, 34(6), 820-847, 2014
LOBSTER (DECAPODA) DIVERSITY AND EVOLUTIONARY PATTERNS
THROUGH TIME
Carrie E. Schweitzer 1and Rodney M. Feldmann 2,∗
1Department of Geology, Kent State University at Stark, 6000 Frank Ave. NW, North Canton, OH 44720, USA
2Department of Geology, Kent State University, Kent, OH 44242, USA
ABSTRACT
Analyses of diversity patterns at the infraorder, superfamily and family level demonstrate that the lobsters as a group were significantly
more diverse in the Mesozoic, especially the Triassic and Jurassic, than in the Cenozoic and Holocene. Peaks in numbers of genera
occurred in the Late Jurassic (Oxfordian and Tithonian), Upper Cretaceous (Cenomanian and Campanian), and Holocene. Smaller peaks
occurred in the Lower Triassic (Anisian) and Eocene (Lutetian). Analysis of substrate preferences for various families, based upon rock
type enclosing the fossils, suggests that Nephropoidea and Erymidae were quite variable in substrate preference. When examined by
energy-level preferences, most lobster groups exhibit preferences; some preferences changed through time. A long-standing preference
for low energy substrates appears to support the long held hypotheses that Nephropidae moved into deeper water either because of the
end-Cretaceous events, or due to competition with brachyurans. Inhabitation of many types of substrates seems to predict survival of mass-
extinction events in lobsters as does a shift in substrate preference through time. Lobsters can be arranged into three evolutionary faunas,
Triassic, Jurassic, and Modern, that are distinct and exhibit discrete body plan and faunal turnover patterns. Several biases in the dataset
have been identified. These include a strong bias toward European, Mesozoic occurrences at the expense of other locations and times; an
environmental bias toward epicontinental versus oceanic or active margin occurrences; and monographic biases.
KEY WORDS: Achelata, Astacidea, diversity, Glypheidea, habitats, Polychelida
DOI: 10.1163/1937240X-00002288
INTRODUCTION
Lobsters are one of the most recognizable denizens of the
marine environment and have a reasonably well-reported
fossil record (Garassino, 1996; Feldmann et al., 2013a, b;
Karasawa et al., 2013). They are economically important
in fisheries today; thus, their patterns of evolution and
extinction and the ecological influences on those patterns are
of importance to biologists and paleontologists (Patek et al.,
2006). Lobsters compose a group that is variously defined
depending on the country of origin of the investigator, but
in general, comprise the extant and extinct clawed lobsters,
spiny lobsters, and slipper lobsters as well as the mostly
extinct pseudochelate lobsters (Table 1). They are predators,
prey, and scavengers, and have been implicated as major
drivers in the Marine Mesozoic Revolution (Vermeij, 1977,
1987). Thus, they have been major components in the marine
food web since at least the middle of the Mesozoic.
The geologic history of fossil lobsters spans the Late De-
vonian through to the Holocene, but the Paleozoic occur-
rences are limited to two species, Palaeopalaemon new-
berryi (Whitfield, 1880) and Protoclytiopsis antiqua Bir-
shtein, 1958. Radiation of the group began in the Early Tri-
assic, and the record of taxa from that time to the present
is sufficiently robust to permit elucidation of patterns of
diversity within and among major clades. Similar to the
work of Tsang et al. (2008, Thalassinidea), Palero et al.
(2009, Palinuridae and Scyllaridae) and Bracken et al. (2013,
∗Corresponding author; e-mail: rfeldman@kent.edu
Anomura), Karasawa et al. (2013) analyzed a subset of De-
capoda, the lobsters, based upon adult morphologic char-
acters on 61 extinct and extant lobster genera; their study
built upon previous work and morphological character selec-
tion (Holthuis, 1974; Scholtz and Richter, 1995; Tshudy and
Babcock, 1997; Dixon et al., 2003; Ahyong and O’Meally,
2004; Amati et al., 2004; Schram and Dixon, 2004; Ahy-
ong, 2006, 2009). Their phylogenetic results were topologi-
cally similar to the results of Ahyong and O’Meally (2004)
and to Bracken et al. (2014) at higher taxonomic levels.
The results of the Karasawa et al. (2013) study permitted
arraying lobster taxa within five major clades, Palaeopalae-
monida, Polychelida, Achelata, Glypheidea, and Astacidea.
The Palaeopalaemonida is known only from the Paleozoic
from a single genus and is not useful in detailed diversity
analyses. The results of that study showed a relationship be-
tween Palinuridae and Scyllaridae congruent with that of
Palero et al. (2008), who used molecular data. Boisselier-
Dubayle et al. (2010) recovered a sister group relation-
ship between Glypheidae and Achelata +Polychelida using
molecular data, whereas Karasawa et al. (2013) recovered a
sister relationship between Glypheidea and Astacida. Shen
et al. (2013) showed that Achelata were most basal among
the lobsters based upon molecular data, whereas Karasawa
et al. (2013) showed that Polychelida was most basal and
Boisselier-Dubayle et al. (2010) recovered Astacidea as
most basal. Charbonnier et al. (2014) analyzed a subset of
© The Crustacean Society, 2014. Published by Brill NV, Leiden DOI:10.1163/1937240X-00002288
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 821
Table 1. Lobster classification used in this paper, based upon Karasawa et al. (2013), geologic range for each taxon, and common names for various lobster
groups. See Holthuis (1991) for complete listing of local common names. Polychelida was analyzed at the Infraorder level for energy level and environments
but included the listed families. Some Glypheidea families not analyzed for energy and environments due to very small numbers of taxa. † indicates extinct
taxa.
Infraorder Palaeopalaemonida † Schram and Dixon, 2004, Late Devonian–Early Mississippian
Superfamily Palaeopalaemonoidea † Brooks, 1962, Late Devonian–Early Mississippian
Family Palaeopalaemonidae † Brooks, 1962, Late Devonian–Early Mississippian
Infraorder Polychelida Scholtz and Richter, 1995, Late Triassic (Carnian)–Holocene
Superfamily Eryonoidea De Haan, 1841, Late Triassic (Carnian)–Holocene
Family Eryonidae † De Haan, 1841, Late Triassic (Norian)–Early Cretaceous (Berriasian–Hauterivian)
Family Coleiidae † van Straelen, 1925, Late Triassic (Carnian)–Late Cretaceous (Campanian)
Family Palaeopentachelidae † Ahyong, 2009, Late Jurassic (Kimmeridgian), ?Oligocene
Family Polychelidae Wood-Mason, 1875, Middle Jurassic (Callovian)–Holocene
Family Tetrachelidae † Beurlen, 1930, Late Triassic (Carnian)
Family Tricarinidae † Feldmann et al., 2007, Early Cretaceous (Barremian–Aptian)
Infraorder Achelata Scholtz and Richter, 1995, Middle Triassic (Anisian)–Holocene
Superfamily Palinuroidea Latreille, 1802, Middle Triassic (Anisian)–Holocene
Family Cancrinidae † Beurlen, 1930, ?Early Jurassic (Hettangian–Sinemurian), Late Jurassic (Tithonian)–Late Cretaceous
(Santonian)
Family Palinuridae Latreille, 1802, Middle Jurassic (Anisian)–Holocene, Common names: spiny lobsters, crayfish, crawfish, rock
lobster, langouste
Family Scyllaridae Latreille, 1825, Early Cretaceous (Albian)–Holocene, Common names: slipper lobsters, bugs
Infraorder Glypheidea Winkler, 1881, Permian–Recent
Superfamily Glaessnericarioidea † Karasawa et al., 2013, Upper Triassic (Norian-?Rhaetian)
Family Glaessnericariidae † Karasawa et al., 2013, Upper Triassic (Norian-?Rhaetian)
Superfamily Erymoidea † van Straelen, 1925, Middle Triassic (Anisian)–Eocene (Bartonian)
Family Erymidae † van Straelen, 1925, ?Middle Triassic (Anisian), Early Jurassic (Hettangian)–Eocene (Bartonian)
Family Pemphicidae † van Straelen, 1928, Middle Triassic (Anisian–Ladinian)
Superfamily Glypheoidea Winkler, 1881, Late Permian–Holocene
Family Platychelidae † Glaessner, 1969, Late Triassic (Carnian)
Family Chimaerastacidae † Amati, Feldmann, and Zonneveld, 2004, Middle Triassic (Ladinian-Carnian)
Family Clytiopsidae † Beurlen, 1927, Late Permian–Late Triassic (Carnian)
Family Litogastroidae † Karasawa et al., 2013, Early Triassic (Olenekian)–Late Jurassic (Oxfordian)
Family Glypheidae † Winkler, 1881, Jurassic (Hettangian)–Eocene
Family Neoglypheidae Karasawa et al., 2013, Holocene
Family Mecochiridae † van Straelen, 1925, Early Jurassic (Hettangian)–Late Cretaceous (Maastrichtian)
Infraorder Astacidea Latreille, 1802, Early Jurassic (Pliensbachian)–Holocene
Section Homarida Scholtz and Richter, 1995, Early Jurassic (Pliensbachian)–Holocene
Superfamily Enoplometopoidea de Saint Laurent, 1988, Early Jurassic (Pliensbachian)–Holocene
Family Enoplometopidae de Saint Laurent, 1988, Holocene, Common name: reef lobsters, Hawaiian red lobster, red reef lobster
Family Uncinidae † Beurlen, 1930, Early Jurassic (Pliensbachian)–Upper Jurassic (Tithonian)
Superfamily Stenochiroidea † Beurlen, 1928, Middle Jurassic (Bajocian)–Late Cretaceous (Cenomanian–Turonian)
Family Stenochiridae † Beurlen, 1928, Middle Jurassic (Bajocian)–Late Cretaceous (Cenomanian–Turonian)
Superfamily Nephropoidea Dana, 1852, Early Cretaceous (Berriasian)–Holocene, Common names: American lobster, Cape lobster,
European lobster, Maine lobster, common lobster
Family Nephropidae Dana, 1852, Early Cretaceous (Berriasian)–Holocene
lobsters and retrieved a tree with many unresolved clades but
that largely supported the results of Karasawa et al. (2013)
in the arrangement of Glypheoidea. Thus, each recent phy-
logeny generates a slightly different hypothesis for the rela-
tionships among the lobsters. Because the work of Karasawa
et al. (2013) is the only published phylogenetic hypothesis
to include a broad array of extinct and extant genera, as well
as representatives from each known group, extinct and ex-
tant, we have elected to use the classification generated by
that work. This work differs from that of Klompmaker et al.
(2013) who analyzed overall decapod diversity in the Meso-
zoic, grouping lobsters and shrimp and analyzing environ-
ments based upon occurrence in reefs, siliciclastics, or other
environments.
Using the phylogenetic analysis and generic systematic
placements recognized by Karasawa et al. (2013), diversity
patterns at infraorder, superfamily, and family levels can
be explored and temporal changes in diversity can be
analyzed. The basic taxonomic level used in this paper is the
genus following numerous recent examples (Alroy, 2010;
Aberhan and Kiessling, 2012; Smith et al., 2012; rationale
discussed therein). Most paleontological diversity studies
are conducted at the genus and family level (Smith, 2007).
Because the work is based upon data from all fossil species
of lobsters known to us through August, 2014, this study is
as robust as possible given the fossil record. Stratigraphic,
geographic, and lithologic data have been compiled in as
much detail as the published record permits, and many of
the species have been further verified by personal study of
822 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
specimens in museum collections. These data have permitted
an analysis of the ecological conditions under which species
lived so that broad patterns of ecological preference within
clades can be evaluated and comparisons of preferences
between clades can be established.
The strength of the resulting diversity patterns is based
upon the number of taxa available for analysis and the
quality of the data, published and observational, available
for each taxon. The results are, however, subject to a
variety of biases imposed by the vagaries of the geological
record. In order to present the results in as honest a manner
possible, the various biases we recognized are enumerated
and discussed.
MATERIALS AND METHODS
Database
The data upon which the diversity analyses are based is the list of fossil
species compiled in Schweitzer et al. (2010) to which newly named taxa
have been added. All known genera of lobsters in the fossil record were
included in the analysis. A total of 547 species within 102 genera represent
all of the species of fossil marine lobsters analyzed in this study through the
publication date of August, 2014. We used adult taxa with one exception,
Acanthophoenicides Audo and Charbonner, 2012, which is a subadult. We
did not include the palinurid larval form Phalangites Münster, 1839, as
adult palinurid forms are known from the same deposits. An unnamed
palinurid larval form has recently been published (Haug and Haug, 2013)
as has a named palinurid larval form (Haug et al., 2013). Phylogenetic
analyses, for example, have been shown to produce aberrant results if
not based upon semaphoronts (Wolfe and Hegna, 2014); thus, we restrict
our diversity analysis to only adult forms. In any event, if all palinurid
larval forms were included, an additional 4 genera would be added to the
Tithonian for that group; all are lithographic, Solnhofen-type limestone
occurrences.
A phylogenetic analysis based upon adult morphology, including repre-
sentatives from 44 extinct and 27 extant genera of lobsters, resulted in a new
classification for the group (Karasawa et al., 2013) and forms the basis for
the classification used herein. Every fossil genus with sufficiently preserved
members was included in that analysis. According to that analysis, the lob-
sters are grouped into clades defined upon adult morphology and include
the Palaeopalaemonida, Polychelida, Achelata, Glypheidea, and Astacidea
(Table 1). Each group is temporally as well as morphologically cohesive.
Modern lobster diversity was compiled from Holthuis (1991), De Grave
et al. (2009), and Chan (2010). Fossil diversity was based on Schweitzer
et al. (2010) as a starting point, augmented by subsequent modification
based upon field, museum, and literature analysis. The generic and family
approach involves the inevitable problem of whether decapod workers are
taxonomic lumpers or splitters: should taxa be grouped into many genera or
few? Similarly, decapod workers tend to work across time and across taxa,
not within one period or stage. They work with faunas, not with one family
or one infraorder. Thus, taxonomic names do not tend to be used for Creta-
ceous taxa only, with different names given to otherwise similar Paleogene
taxa (Ausich and Peters, 2005). This problem is obviated to some extent
within Decapoda as there have historically been relatively few workers on
the group and taxonomic decisions have been applied with some uniformity.
In addition, the authors have examined a large proportion of the material and
nearly all of the references on these species, and have been able to confirm
or reject and reassign taxa. Thus, taxonomic decisions herein (species as-
signed to a genus, genera assigned to a family) have been made uniformly,
if not to the agreement of all workers.
There are a few especially speciose genera that bear mentioning. Ho-
ploparia M’Coy, 1849, embraces 66 species, and one of these for example,
has eighteen synonyms, all named by the same author based on appendage
fragments. Tshudy and Sorhannus (2003) considered Hoploparia to be a
“wastebasket,” containing a broad array of morphologies, but did not pro-
pose a solution. Feldmann et al. (2007) examined all species of Hoploparia
and found a consistent morphological pattern among them, retaining all
species within it and maintaining the genus as now construed. Linuparus
White, 1847, contains 32 species, all of which were examined by Feld-
mann et al. (2007) and considered to be embraced within the same genus.
Eryma von Meyer, 1840, is very speciose, embracing 57 species. It has not
been recently revised but the concept of Eryma has been stable. Recently,
Charbonnier et al. (2013) resurrected three genera within Glypheidae that
had been named in the early Twentieth Century but later synonymized with
other genera, adding to the number of recognized genera within that group.
Thus, it is obviously possible that some of the very speciose genera may be
subdivided in the future as new material is recovered. Any diversity study
is a snapshot of the current knowledge at hand.
Geologic ranges and determination of the lithology enclosing the species
within the list have been determined by examination of published literature,
examination of museum labels, and in some cases communication with
specialists when questions arose. Geologic ranges were recorded and
plotted by stage, as this represents the most precise determination possible.
Diversity Curve
All taxa were counted, including singletons, because the latter comprise
such a large percentage of the decapod record. Occurrences of lobster
geologic ranges were assembled using the range through method (following
Aberhan and Kiessling, 2012), with some exceptions. Extant taxa with a
single occurrence prior to the Miocene were not automatically extended
into the Holocene. If an extant genus had scattered occurrences through the
record prior to the Oligocene and including the post-Oligocene (Eocene,
Oligocene, and Miocene), then its geologic range was extended into
the Holocene. If the authors had verified an identification personally
by examination of type material for long-ranging taxa, their range was
extended through the geologic time periods in which the taxon was absent.
Occurrences in which the first or last occurrence was separated from
the main cluster of occurrences by more than 50 million years were not
extended through the missing time slices, unless the identification was
verified by us as noted (for example, first occurrence as Middle Triassic and
then the main cluster of occurrences in the Late Jurassic and Cretaceous).
For taxa in 19th Century literature noted as “Cretaceous,” “Senonian,” or
other poorly constrained times, every attempt was made to more narrowly
pinpoint the time stage. In the few cases where this was not possible, the
taxon was assigned to the youngest time interval so as not to inflate the
range of the lineage, recognizing that this might inflate diversity in the
latest stage in an epoch. This does not seem to have happened, however,
as the Campanian is just as highly diverse as the Maastrichtian (both Late
Cretaceous), and every Tithonian (Late Jurassic) occurrence was actually
recorded in the literature as Tithonian, for example.
For taxa with ranges of occurrences or age ranges for rock units, we
assigned the taxon to each time unit. This may have artificially inflated the
diversity slightly for some intervals, notably Berriasian-Hauterivian (Early
Cretaceous), the units to which this relates most significantly.
A significant proportion of the identifications, taxonomy, and phylogeny
of the decapod crustaceans have been verified by the authors with
colleagues and students. Nearly all southern hemisphere specimens have
been examined by one (RMF) or both of us (Feldmann and Schweitzer,
2006). North Pacific and Central American decapods have been examined
and summarized (Schweitzer, 2001; Schweitzer et al., 2002). The lobster
phylogeny used herein (Karasawa et al., 2013) was prepared with our
colleague, H. Karasawa. Feldmann has a substantial record of fossil lobster
work (see references in Schweitzer et al., 2010). Thus, the problems of the
taxic approach (Aberhan and Kiessling, 2012) can be avoided because the
authors have personally verified as much of the database as possible.
Environments
Each species preserved as a fossil was examined for the rock type in which
it was enclosed. This was extracted from the primary literature, either from
the paper in which it was first named, from subsequent reports of the
species, from personal examination of specimens enclosed in rocks, or from
examination of other literature on the enclosing rock units (Roemer, 1846;
Fraas, 1851; Heer, 1876; Tate and Blake, 1876; Gignoux, 1955; Brookfield,
1973; Embry, 1984; Schweigert, 1999; McCann, 2008; Charbonnier et al.,
2010, 2012a, b; Powell, 2010). Species were tabulated by stage and rock
type.
We attempted to include all possible rock types encountered in the
literature, within the limitations of language constraints, in our raw data.
For example, in German, Kalk or Kalkstein means limestone, so we
presumed it to mean any type of limestone, not the more specific meaning
that chalk has in English. Once the enclosing rock type was determined
for each lobster species, it became readily evident that there were many
environments that suggested similar habitat conditions in terms of energy
level. In addition, analysis based upon rock type only was too cumbersome
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 823
Table 2. Rock types and their assigned energy level.
Rock type Assigned energy/
Environment
Shale, clay, ironstone Low
Mixed siliciclastic (sandy siltstone,
sandy mud) Medium
Sandstone, greensand, ferruginous
oolite High
Coralline limestone, coral, limestone
with corals Coral Reef
Sponge reefs, limestone with sponges Sponge-rich
environments
Limestone, chalk, calcarenite,
dolomite, phosphates, marls,
oolitic limestone
Carbonates
Lithographic limestone Lithographic limestone
(variable
environmental
interpretations)
to yield interpretable results. As an example, the genus Glyphea alone is
preserved in fourteen different rock types. Thus, a means of simplifying the
characterization of environment became necessary.
For the environmental analysis, we grouped environments according to
rock type and energy level (Table 2). We grouped those rocks described as
limestones of any kind, including marls, as carbonates. Coral-bearing lime-
stones and sponge-bearing limestones were retained as separate environ-
ments recognizing that they could be reefs or bioherms, specific environ-
ments. Lithographic limestones were also retained as a separate environ-
ment because there is considerable variation in the possible environments
that the animals may have inhabited. The Solnhofen-type lithographic lime-
stone may include animals from sponge-algal mounds, corals, the open
ocean, or lagoons (Barthel et al., 1994). The Lebanese plattenkalks are de-
scribed as having been deposited in shallow basins, sometimes containing
rudists (Audo and Charbonnier, 2012). We used the terms low, medium, and
high to represent low, medium, and high energy siliciclastic environments
based upon grain size. If the specimen was enclosed in sediment described
as sandy mud, silty mud, etc., we recorded it as mixed siliciclastic.
The issue of autochthony and allochthony of specimens was dealt with
on a case by case basis. Rarely was there evidence of allochthony for the
specimens of lobsters, but in each case interpreted to be allochthonous,
we recorded the lithology and age of the original enclosing sediment, not
the sediment in which the specimen was ultimately deposited. For those
specimens which might have been washed into a different area than that in
which they lived, we have used the interpretations of the authors in the most
recent work on the relevant species and rocks.
Abbreviations
CM, Carnegie Museum of Natural History, Pittsburgh, PA, USA; GSC, Ge-
ological Survey of Canada, Eastern Paleontology, Division, Ottawa, ON,
Canada; KSU D, Kent State University Decapod Comparative Collection,
Kent, OH, USA; SM Sedgwick Museum, Cambridge University, Cam-
bridge, UK; USNM, United States National Museum of Natural History,
Smithsonian Institution, Washington, DC, USA.
Software
Excel 2010; PAST version 2.00 (April 2010), Paleontological Statistics
software package for education and data analysis (Hammer et al., 2001).
BIASES
Overview
Numerous biases have been recognized as impacting diver-
sity curves. We examined several. In general, two different
effects might be anticipated as a result of intrinsic and ex-
trinsic biases.
Extrinsic biases, including rock volume, geographic ef-
fects, and monographic issues, among others have a pro-
found effect on the total number of taxa that have been de-
scribed and that are incorporated within the study. Thus,
when examining plots of total numbers of taxa against time,
these factors clearly bias the results. However, the extrin-
sic biases exert less effect on relative abundances of lobster
taxa because the extrinsic biases affect the lobster taxa more
or less equally through time. Although some monographic
studies are focused on subgroups within Decapoda, such as
Brachyura (Wright and Collins, 1972, for example), mono-
graphic studies dealing with lobsters (Woods, 1925-1931,
for example) would cover the entire spectrum of groups con-
sidered within this study. As a result, comparisons of diver-
sity patterns between various lobster groups should not be
significantly affected.
Intrinsic biases on lobster diversity include habitat dif-
ferences and preservational potential. These factors poten-
tially could have an effect on both total diversity of lob-
ster taxa and on relative diversity of taxa compared to one
another. If, for example, a group of organisms is confined
to a particular habitat that is generally underrepresented in
the fossil record, the frequency of occurrence of that taxon
will be minimized. Similarly, if the habitat preference of a
group changes through time, as seen in the families within
Glypheoidea discussed herein, diversity records will be bi-
ased by those changes. These habitat issues can affect both
total and relative diversity measures. Similarly, preserva-
tional potential can affect total and relative diversity deter-
minations. Preservation of decapods in general is controlled
by durability of the cuticle comprising the skeletal elements
of the animal and on taphonomic factors. Considering that
taphonomic factors are extrinsic factors affecting diversity
measures as described above, durability of cuticle remains
as an intrinsic factor (Waugh et al., 2009; Feldmann et al.,
2012). Preservational potential of cuticle is governed largely
by the degree of calcification of the integument. Although
this varies considerably across the entire spectrum of de-
capods, lobsters in general have well calcified cuticle and,
therefore, cuticle structure is likely to be a minimal factor in
diversity studies.
Rock Volume
Raup (1976a, b) first called attention to the issue of differ-
ential rock volume for various time periods. Later workers
have explored the correlation between rock volume and sea
level and the impact that these have for the number of fos-
sils, species, and genera per time period (Smith and Mc-
Gowan, 2007; Wall et al., 2009). We explored two aspects
of rock outcrop with respect to impact on diversity. Raw
data of number of rock outcrops of Mesozoic and Ceno-
zoic stages from England, Wales, France, and Spain (Smith
and McGowan, 2007: supplementary data, raw data, avail-
able online at http://www.palass.org) were plotted versus to-
tal number of lobster genera per stage. This was done to de-
termine if lobster diversity was driven by the prevalence of
north-western European rock outcrop (Fig. 1A); there are a
large number of lobster occurrences from the UK and France
for example, that we hypothesized could overprint the lob-
ster diversity curve. There seems to be some relationship be-
tween the two curves; at times as numbers of outcrops in-
824 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 1. Lobster genera plotted versus rock volume. A, Rock volume measured as number of maps with outcrops of a given age in England, Wales, France
and Spain (dashed line) plotted with number of lobster genera in the same ages (solid line). Ages in millions of years on x-axis; y-axis log scale. Data from
Smith and McGowan (2007: supplementary data, raw data, available online at www.palass.org). B, Number of North American formations at selected time
intervals (circles) plotted with number of lobster genera in the same ages (solid line). Ages in millions of years on x-axis; y-axis log scale. Data from Peters
and Foote (2001: Appendix 2).
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 825
crease, the number of genera also increases (Pliensbachian,
Early Jurassic; Albian, Early Cretaceous). At other times,
as the rock volume increased (Danian, Paleocene; middle
Miocene), number of genera decreased or stayed flat. We
also plotted data from Peters and Foote (2001: Appendix 2)
on number of North American named marine formations of
various time intervals versus the total number of lobster ge-
nera (Fig. 1B). Patterns between the two plotted curves are
not really consistent, albeit note that data points for number
of North American formations are few. Data for the Ceno-
zoic, similar to that for England, Wales, France, and Spain,
are not consistent with the number of lobster genera. Thus,
the lobster record as a whole does not seem to be driven by
the rock record of a particular continent, or geographic area
or of a particular time period based upon these examinations.
Geographic Bias
The lobster record, especially in the Triassic and Juras-
sic, appeared to be heavily skewed toward European occur-
rences (Europe here defined as all countries on the Euro-
pean continent west of the Ural Mountains excluding for-
mer USSR countries) based on our qualitative examina-
tion. We supposed that may be because of collector bias
or longer history of collecting in that region (biases sum-
marized in Aberhan and Kiesling, 2012). We plotted the
total number of genera within various higher level lobster
taxa against region and found that European occurrences
had varying biasing effects. Nephropoidea seems to be dom-
inated by non-European occurrences (Fig. 2A), whereas
Glypheidea appears to be heavily biased by the European
record (Fig. 2B). Non-European Achelata and Polychelidae
records were heavily dominated by European occurrences
in the Triassic and Jurassic, but non-European records more
closely paralleled European records from the Cretaceous on-
ward (Fig. 3).
A part of the apparent European bias is based upon the
amount of research and the length of time research has
been conducted on that continent. Serious study of fossil
decapods in Europe began in the late 18th Century and
was actively being pursued long before similar studies be-
gan in North America in the 19th Century (Smith, 2007;
Schweitzer and Feldmann, in press). Work in other parts of
the world followed much later. Additionally, the vast major-
ity of paleontologists have been concentrated in Europe and
North America historically. Thus, it is not surprising that the
majority of records of fossil decapods are known from these
areas.
A final geographic bias is imposed by the observations
that the largest proportion of continental landmass is in
the Northern Hemisphere and that a vast amount of that
area has received epicontinental seaway and continental
shelf sediments. These are the areas in which most marine
invertebrates live and are preserved in the fossil record, so it
is not surprising that the patterns of distribution and diversity
are dominated by taxa from North America and Europe.
Paleontological research on decapods in Africa and Asia
has been slower to develop, and the fauna there is vastly
underrepresented for that reason.
Nomenclatorial Inconsistencies
Systematic studies by different workers in different regions
and at different times historically can result in proliferation
of names that do not reflect precise biological relationships.
There seems to be a tendency to apply new names to
taxa discovered in different countries or regions, despite
their being quite similar morphologically. It is also possible
that systematists working primarily on material from a
limited time interval, for example the Maastrichtian, might
generate unique names based upon the notion that the taxa
are different from those in bounding time units despite
their being similar morphologically (Ausich and Peters,
2005). The judgments about assignment of species to genera
(Schweitzer et al., 2010) and part of the development of
the phylogeny of the lobsters (Karasawa et al., 2013) were
performed by the authors. The problems of nomenclature
were therefore minimized to the best of our abilities. It is
possible that names of extant taxa, particularly at the generic
level, might have been extended too far into the geological
record, but examination of the diversity patterns does not
suggest a significant effect. As with other nomenclatorial
biases, the judgment about generic placement was that of
the authors and was uniformly applied.
Ecological Biases
The occurrences of fossil decapods in littoral, sublittoral,
and deep water (bathyal and abyssal) are not uniformly rep-
resented. Without question, the vast preponderance of deca-
pod fossils, as with most marine organisms, have been col-
lected from rocks deposited in sublittoral environments. Lit-
toral environments tend to be zones of relatively high en-
ergy and ones in which long term sediment accumulation
is minimal. The preservation potential for decapods occu-
pying these settings is minimized Although sediment accu-
mulation in regions beyond the continental shelves does oc-
cur actively, the sediments are only infrequently exposed for
study. Thus, the possibility of finding fossils of organisms
occupying these regions is limited. These are issues that are
difficult to assess quantitatively as the evidence is negative.
Absence of taxa in the fossil record cannot simply be at-
tributed to the animals living in an environment that is un-
likely to be represented. For example, the majority of occur-
rences of Nephropoidea in modern oceans are from deep wa-
ter (Holthuis, 1991), whereas many of those from the fossil
record are from rock units interpreted to have been deposited
in shallow water (Rathbun, 1935; Stenzel, 1945; Feldmann
et al., 2013c). The sparse record of nephropoids in the Ceno-
zoic as compared to the Mesozoic could either be attributed
to a real decline in diversity or, more probably, to a shift in
habitat preference.
Overprint of Lagerstätte
Diversity patterns can be affected by occurrences of remark-
ably preserved faunas, or Konservat-Lagerstätte. The Titho-
nian Solnhofen and related rocks in southern Germany are
one such example. Another is the Cenomanian (Late Cre-
taceous) of Lebanon. These occurrences are remarkable be-
cause a wide range of organisms are preserved, and the state
of preservation is excellent. This permits detailed discrim-
ination between specimens that has resulted in description
826 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 2. Effect of geographic bias on diversity. A, All Nephropoidea, here including Nephropidae, Stenochiridae and Uncinidae (dashed line), plotted with
only non-European occurrences (solid line). B, All Glypheidea (dashed line), plotted with only non-European occurrences (solid line). Numbers of genera
may add up to more than the total number of genera for the families on the diversity curve (Fig. 4) for a given time interval because a genus can occur in
non-European areas and in Europe during the same time interval (Europe defined as all countries on the European continent west of the Ural Mountains
excluding former USSR countries).
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 827
Fig. 3. Effect of geographic bias on diversity. All Achelata and Polychelida, here (dashed line), plotted with only non-European occurrences (solid line).
Numbers of genera may add up to more than the total number of genera for the families on the diversity curve (Fig. 4) for a given time interval because a
genus can occur in non-European areas and in Europe during the same time interval (Europe defined as all countries on the European continent west of the
Ural Mountains excluding former USSR countries).
of a large number of species. As a result, the total diver-
sity of decapods within those time intervals may be inflated
as a result of inclusion of exceptionally preserved assem-
blages, but relative frequencies of lobster taxa should not be
affected. The overall record of lobsters does not seem to be
as dependent on Lagerstätte as had been previously thought
(Sepkoski, 2000).
Monographic Issues
Publication of monographic works can exert a strong effect
on data employed in diversity studies (Raup, 1976a, b).
For example, the work of Rathbun (1935) substantially
increased the records of decapods, including lobsters, from
North America. Perhaps even more striking, Garassino and
Teruzzi (1993) and Garassino et al. (1996) described a
large assemblage of macrurans from the Triassic of Italy.
As with other extrinsic factors, although these works had
a considerable effect on the total number of American and
European decapods, respectively, they did not substantially
perturb diversity considerations within groups. There is no
reason to conclude that one group of decapods would have
been sampled at the expense of others. Rather, they provide
information to more accurately reflect the actual differences
in diversity between groups. As suggested by Raup (1976a,
b), if studies are conducted on a large group over broad
geographic areas, these effects should be minimal.
Preservational Biases
Resistance of cuticle material to taphonomic processes is an
intrinsic factor that can potentially affect conclusions about
overall diversity as well as diversity between taxa. Cuticle is
comprised of an organic matrix that is variously thickened
and hardened by precipitation of calcium carbonate, either
as calcite or as amorphous calcium carbonate (Amato et al.,
2008; Mutel et al., 2008). Although these studies focused
primarily on cuticle of brachyurans, both demonstrated a
strong relationship between calcification and preservation
potential. Work under way on cuticle thickness and dura-
bility of lobster cuticle (Waugh, 2013) indicates that lobsters
in general have well calcified, and therefore durable, cuti-
cle. The general resistance to taphonomic effects would lead
to the conclusion that lobsters, like the crabs, would have a
reasonably high probability of being preserved, contrary to
previous hypotheses (Sepkoski, 2000). Further, work with
extant lobsters in the spirit collection of the Smithsonian
Institution confirms that, at least qualitatively, lobsters are
828 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
more or less equally well calcified. Thus, the condition of
calcification and strength of cuticle would not seem to be a
significant bias in terms of diversity studies between taxo-
nomic groups.
Another preservational bias relates to the nature of preser-
vation in different sedimentary environments. Dodd and
Stanton (1981: 128) indicated that carbonate fossil remains
preserved in siliciclastic sediments have a higher probabil-
ity of dissolution than those preserved in carbonate rocks.
This observation would seem to suggest that decapods would
have a higher preservation potential in carbonate terraines
than in siliciclastic terraines. Because carbonate rock abun-
dance is highest in low latitude regions (Schopf, 1980: 195),
it would seem to follow that low latitude decapods would
be better represented than those from higher latitude regions
where siliciclastic sedimentation predominates. The combi-
nation of these observations would suggest that decapods
preserved in low latitude, carbonate regimes would tend to
retain their carbonate composition and to be more likely pre-
served, whereas those in higher latitude, siliciclastic rocks
would be altered chemically or lost. There is some anec-
dotal evidence, currently being investigated in more detail,
to suggest that this is true. Cuticle preserved in carbonate
rocks more often seems to preserve calcite than does cuti-
cle from siliciclastic rocks. Preservation in siliciclastic rocks
yields cuticle that is often altered to apatite as recently docu-
mented in the Late Cretaceous Bearpaw Formation in Mon-
tana (Feldmann et al., 2012). The effect of preservational
style on diversity has not yet been explored in detail.
RESULTS
Diversity
The overall diversity curve for all lobster families indi-
cates two clear patterns: family level diversity was high-
est in the Mesozoic with a very clear decline by the end
of the Cretaceous with another, smaller peak in the Eocene
(Fig. 4). Generic level diversity is highest in the Holocene.
Family-level diversity reached two peaks, one in the Titho-
nian (Jurassic), when 16 families were present, with 29 ge-
nera, and another in the Cenomanian (Cretaceous), when 11
families were present with 26 genera. Following the Titho-
nian high, faunal turnover occurred in the Early Cretaceous,
resulting in the appearance of new families (Nephropidae,
Scyllaridae) and the disappearance of some Jurassic fami-
lies (Litogastroidae, Eryonidae), building up to the middle
Cretaceous peak. Some of these families became extinct in
the middle Late Cretaceous, but only one became extinct at
the end of the Cretaceous, Mecochiridae.
The Tithonian contained a high of 29 genera, followed
by the Cenomanian at 26 genera. Extant lobsters number
about 62 genera. This could be interpreted as being more
than twice as diverse at the generic level than the Mesozoic,
but this seems unlikely as that would imply that the complete
record of Mesozoic lobsters is known. Thus, it is almost
certainly the case that lobsters overall were more diverse at
the family level in the Mesozoic than in the Holocene.
Examination of the Maastrichtian (end Cretaceous)
through Chattian (Oligocene) portions of the curve show a
decline, with a spike in the Lutetian (Eocene), followed by
a plateau, then a slight climb, followed by a giant spike at
the Holocene. The classic (legacy) hypothesis for this has
been that lobsters moved offshore into deeper water and thus
were not as likely to have been discoveredas fossils (Glaess-
ner, 1969; Tshudy, 2003). As we will show, this is supported
for two modern families (Nephropidae, Polychelidae) and
rejected for two others (Palinuridae, Scyllaridae). Indeed,
all extant lobster families inhabit environments unlikely to
be fossilized, and it is possible that they could have inhab-
ited such environments since the beginning of their geologic
range but lack a fossil record for those environments.
Polychelida.—The polychelids (Fig. 5C) embraces those
lobsters with 4 pairs of claws and a dorsoventrally flattened
body. Extant forms inhabit bathyal habitats, with some rang-
ing to abyssal or outer sublittoral (Poore, 2004; Ahyong,
2012). The families within Polychelida are small, composed
of seven or fewer genera. Eryonidae reached a peak generic
diversity in the Tithonian, whereas Coleiidae plateaued in
diversity from the Norian (Late Triassic) through the Callo-
vian (Middle Jurassic). The remaining families within the
Infraorder had scattered occurrences through the Mesozoic
and Cenozoic, and the Polychelidae is represented today by
six genera.
Achelata.—The achelates (Fig. 5D, E) consists of those
lobsters without claws. Their body form varies from dorso-
ventrally flattened to rectangular in cross-section. Although
appearing early in the record in the Triassic, they began to
radiate in the Late Jurassic and continued the radiation from
that time forward. Achelata achieved their highest diversity
in the Holocene. Both of the major families within the group,
Palinuridae and Scyllaridae, began their major radiation in
the Late Cretaceous.
Glypheoidea.—The glypheoids (Fig. 5B) are part of the In-
fraorder Glypheidea and is composed of several families
that are united in possessing either pseudochelate or weakly
chelate first pereiopods, cylindrical carapaces, and small
pleons for their size as compared to other lobsters. Kara-
sawa et al. (2013) recently proposed a phylogeny separating
the group into six families. The group exhibited a Triassic
diversity pulse, with Clytiopsidae, Litogastroidae, and Chi-
maerastacidae appearing early in its history only to be re-
placed by Glypheidae and Mecochiridae, which were abun-
dant and important lobster groups in the Jurassic and into the
Cretaceous. Glypheoidea is represented in the modern fauna
only by Neoglypheidae.
Erymoidea.—The erymoids also are part of the Infraorder
Glypheidea and includes the clawed lobsters with cylindrical
cephalothoraxes and well-developed groove patterns. They
flourished primarily in the Jurassic although extending
into the Cretaceous and Paleogene. One small family,
Pemphicidae, is known only from the Triassic.
Homarida.—The homarids include nephropoids (Fig. 5A),
what are commonly known to the general public as the
American and European lobsters, and comprise those lob-
sters with large claws, cylindrical cephalothoraxes, and well-
developed pleons. Their diversity increased dramatically by
the Late Cretaceous and they continue to be one of the ma-
jor lobster faunal components in modern oceans. Smaller
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 829
Fig. 4. Diversity curve at genus level for all lobster families, extinct and extant. Note x-axis not scaled to length of time intervals in years. This figure is
published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.
groups of clawed lobsters, Stenochiridae and Enoplome-
topoidea, exhibited their highest diversity in the Jurassic.
Overall Diversity.—From the above summaries of each
lobster group, it is clear that the tempo and mode of radiation
of individual families, and even infraorders, was unique.
Lobster suprageneric groups replaced one another through
time until about the Turonian (Late Cretaceous), when this
turnover began to wane and seems to have stopped in the late
Paleogene.
Habitat Preferences
Keeping in mind the biases discussed above, we inferred
habitat preferences for every species for which the informa-
tion was available within each lobster family, by examining
the rock type and inferring energy level and environment
in which they were deposited. These results were plotted
graphically (Figs. 8-12, see below) and tested using Princi-
ple Coordinates Analysis (PCO, using PAST, Euclidean dis-
tance), in which the number of species in an environment
per stage in each family was compared (Fig. 6). We tested
low energy, high energy, coral-bearing, lithographic lime-
stone, and carbonate environments as these were those that
were most widespread across taxa. In each case, coordinates
1 and 2 accounted for at least 60% of the variance in the
dataset. PCO is described as particularly powerful for eco-
logical datasets (Hammer and Harper, 2006) and is useful
for finding similarities (or differences) among datasets. Un-
like Principal Components Analysis, which is described as
most useful for data with normal distributions, PCO can be
used on any type of data distribution (Hammer and Harper,
2006).
Examination of the PCO plots of lobster families by
environment (except Homarida), plotted as a group with
all three fossil marine families together, and Clytiopsidae,
Fig. 5. Representative lobsters and body plan. A, Nephropidae, Hoploparia longimana (Sowerby, 1826), KSU D 1488, cast of SM B 62690, with cylindrical
cephalothorax and chelate first pereiopods body plan; B, Glypheidea, Glyphea robusta Feldmann and McPherson, 1980, latex mold of holotype GSC 61398,
with cylindrical cephalothorax and pseudochelate first pereiopods body plan; C, Polychelida, Cycleryon propinquus (Schlotheim, 1822), CM 34359, with
flattened cephalothorax and chelate pereiopods body plan; D, Linuparus grimmeri Stenzel, 1945, USNM 2009820, cylindrical/rectangular cephalothorax,
achelate pereiopods body plan; E, Scyllarides nodifer (Stimpson, 1866), USNM 574950, dorsoventrally flattened cephalothorax, achelate pereiopods body
plan, ventral view. Scale bars =1cm.
830 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 831
Fig. 6. Principle Coordinates Analysis of number of lobster species occurring in various environments during time intervals plotted in Figs. 6-10, analyzed
in PAST, Euclidean distance, coordinates 1 and 2 plotted in all cases. Note low energy is only environment which includes all lobster groups. A, lobster
occurrences in low energy environments; B, lobster occurrences in high energy environments; C, lobster occurrences in carbonate environments; D, lobster
occurrences in coral-bearing environments; E, lobster occurrences in lithographic environments. C/C/P =Chimaerastacidae, Clytiopsidae and Platychelidae,
analyzed together because of small numbers of genera and species and because all belong to Glypheoidea.
Chimaerasticidae, and Platychelidae (plotted together due
to small numbers of genera within each) suggests that
each lobster taxon had a distinct environmental distribution
through time. There is some indication of niche partitioning.
Litogastroidae is preferentially preserved in carbonates and
coral bearing rocks as is Erymidae. Homarida have never
yet been recovered from coral-bearing rocks although they
are found in low and high energy siliciclastics and carbonate
rocks. Lithographic limestone appears to show separation
of all groups, but almost all occurrences in this type
of environment are in the Tithonian (Jurassic). Note the
small distances of separation among taxa in lithographic
limestones (Fig. 6E), which suggests that the groups were
largely separated by numbers of taxa within the Tithonian.
Homarida plotted distinctly from the other groups in low,
high, and carbonate environments (Fig. 6A-C) and was
excluded from the coral analysis as it never occurred in
that environment. Erymidae also plotted distinctly in those
three environments although not in the coral environment.
The achelate families, Palinuridae and Scyllaridae, gener-
ally plotted distinctly from the other families and generally
were preferentially recovered from low energy rocks when
other families occurred in high energy or carbonate environ-
ments in addition to low energy. Of the glypheoid families,
the most distinct were Litogastroidae and Mecochiridae. In-
terestingly, none of the Polychelida families ever achieved
good separation from the other groups, except the Poly-
chelidae in low energy. Eryonidae and Coleidae were dis-
tinct from one another, perhaps indicating niche partitioning
within the Infraorder.
To test our assumption that habitats could be general-
ized at the family or superfamily level for extinct lob-
sters, recognizing that there will be exceptions, we ana-
lyzed nearly all extant lobster genera, for which we coded
environmental characters available from literature searches
(Gore, 1984; Holthuis, 1984, 1991; Galil, 2000; Ahyong
and Brown, 2002; Poupin, 2003; Ahyong and Chan, 2004;
Poore, 2004; Chan, 2010). These characters most often in-
cluded depth (depth and environment assignment based on
NOAA.gov) and substrate. For some taxa, data was available
for environmentally-related behavior (infaunal, epifaunal,
nektonic, burying, migratory, nocturnal, gregarious, scav-
enger, predator). The resulting data matrix has 49 genera and
24 characters. Cluster analysis demonstrates that lobster ge-
nera group largely by family (Fig. 7). Thus, our data indi-
cate that both extant and extinct lobsters cluster by preferred
habitat.
Energy and Environments
Polychelida.—Eryonidae and Coleiidae inhabited very dif-
ferent environments. Coleiidae primarily inhabited car-
bonates, including sponge-bearing carbonates (Fig. 8A),
whereas Eryonidae inhabited primarily lithographic lime-
stone depositional environments, dominated by the Soln-
hofen-type limestones (Fig. 8B). Thus, niche partitioning ap-
pears to have occurred within Polychelida during the early
832 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 7. Cluster analysis of extant lobster genera, Euclidean distance measure, paired group, bootstrapping N=100 (numbers on branches), generated in
PAST, based upon 23 characters. Families indicated.
history of the group. Overall, the group preferred carbonate-
dominated environments in its early history, whereas the
modern members of the group prefer low energy environ-
ments. The spike of sponge-rich environment forms in the
Sinemurian (Early Jurassic) is largely a monographic effect
(Garassino and Teruzzi, 1993).
Glypheoidea.—Within Glypheoidea, some families exhib-
ited environmental preferences distinct from other con-
stituent families. Mecochiridae inhabited low energy sili-
ciclastic environments, primarily shallow ones, during the
Late Jurassic and Early Cretaceous while also exhibiting
a preference for non-coral bearing carbonates (Fig. 9A).
Many mecochirids are found in the Solnhofen-type lime-
stones. Litogastroids are almost exclusively found in non-
coral bearing carbonates with some also occurring in low-
energy siliclastics (Fig. 10B). Other glypheoid families have
more variable occurrences. The small families, grouped to-
gether, Clytiopsidae, Chimaerastacidae, and Platychelidae
are primarily found in non-coral bearing carbonates and
high-energy deposits (Fig. 9B). Glypheidae occur in nearly
all examined rock types but show a trend over time from car-
bonates to high energy siliciclastics (Figs. 6, 11B).
Erymidae.—Erymidae, part of the glypheideans, exhibits the
most diverse array of environments inhabited, based upon
rock type, of any lobster group. PCO plots show Erymidae
as distinctive for low energy and carbonate environments
and similar to Glypheoidea for high energy environments
(Figs. 6, 10A). Thus, they apparently occupied a distinctive
niche among the lobsters for much of their existence.
Erymids, like the glypheids, shifted their environmental
preferences. Erymids always had a strong presence in
carbonate environments, but they also occurred in low
energy as well as other environments early in their history.
By the early Late Cretaceous, their occurrence in low energy
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 833
Fig. 8. Lobster groups (Table 1) by energy and environments as defined in Table 2 through time. A, Coleiidae; B, Eryonidae. Numbers of species may add up
to more than the total number of species for the genera recorded during a given time interval because a species can occur (rarely) in multiple environments dur-
ing the same time interval. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.
com/content/journals/1937240x.
834 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 9. Lobster groups (Table 1) by energy and environments as defined in Table 2 through time. A, Mecochiridae; B, Chimaerastacidae, Clytiopsidae and
Platychelidae, plotted together because of small numbers of genera and species and because all belong to Glypheoidea. Numbers of species may add up to
more than the total number of species for the genera recorded during a given time interval because a species can occur (rarely) in multiple environments during
the same time interval. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.
com/content/journals/1937240x.
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 835
Fig. 10. Lobster groups (Table 1) by energy and environments as defined in Table 2 through time. A, Erymidae; B, Litogastroidae. Numbers of species may
add up to more than the total number of species for the genera recorded during a given time interval because a species can occur (rarely) in multiple environ-
ments during the same time interval. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.
brillonline.com/content/journals/1937240x.
836 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 11. Lobster groups (Table 1) by energy and environments as defined in Table 2 through time. A, Homarida (includes Nephropidae, Stenochiridae,
Uncinidae, Enoplometopidae); B, Glypheidae. Numbers of species may add up to more than the total number of species for the genera recorded during a
given time interval because a species can occur (rarely) in multiple environments during the same time interval. This figure is published in colour in the
online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 837
siliciclastic environments disappeared entirely. Erymidae
barely lingered into the Paleogene (Garassino et al., 2009d).
Homarida.—Homarida includes the modern nephropid lob-
sters as well as some closely related extinct families, and
they plot distinctly in PCO plots (Fig. 6). The group ex-
panded its occupation of carbonate environments in the Cre-
taceous, always maintaining presence in all siliciclastic envi-
ronments, but disappeared from carbonates after the Danian
(Fig. 11A). Modern occurrences are almost all in low energy
environments although Homarus Weber, 1795, was histori-
cally noted in shallow, intertidal to sublittoral habitats until
fisheries truncated its range (Herrick, 1911; Thomas, 1968).
Palinuridae.—Palinuridae have a modest fossil record, and
their fossil occurrences are almost all in siliclastic environ-
ments in a mix of low, medium, and high energy (Fig. 12A).
Their habitat preferences seem to be quite different than that
of most lobsters (Fig. 6), and they shifted preferences toward
coral reefs and into more medium and high energy environ-
ments in modern oceans. The shift may have been relatively
recent, as Eocene, Oligocene, and Miocene records are from
low energy settings, and there are certainly abundant coral
reefs known from those time periods from which specimens
might have been collected were they present. This hypothe-
sis will be tested as more and more reef rocks are collected
from these time periods.
Scyllaridae.—Scyllaridae have a very poor fossil record,
and where known, they are mostly from low energy or
lithographic deposits. Like Palinuridae, modern forms live
in completely different environments from the fossil forms,
including coral reefs, sponge reefs, and high and medium
energy environments (Fig. 12B). The sparse fossil record of
this group makes it difficult to speculate on when the habitat
shift occurred, but as for palinurids, coral occurrences in the
Eocene through Miocene might be expected if they indeed
where living there. This hypothesis will be tested as more
and more reef rocks are collected from these time periods.
Environments and Extinction
Most notable is the very low number of occurrences of
lobster fossils from coral bearing rocks. To be sure they
are found associated with corals, but no group of lobsters
is found predominantly in coral-bearing rocks, or sponge-
bearing rocks, until the Holocene. Klompmaker et al. (2013)
hypothesized that the collapse of reef environments may
have played a role in extinctions of decapods at the end of
the Jurassic, but data on lobsters indicates that other factors
must also play a role in lobster extinction. Based upon our
data, it appears that those lobster families surviving the end-
Cretaceous extinction event were those that inhabited the
broadest array of environments prior to and leading up the
extinction event. Of the K/T boundary crossers with ample
data, Erymidae and Glypheidae inhabited seven total of our
defined environments (Table 2), were found in numerous
rock types, and five (Erymidae) and four (Glypheidae) of
these in the Late Cretaceous (Figs. 10A, 11B). Homarida
and Palinuridae inhabited five total environments and four
of these during the Late Cretaceous (Figs. 11A, 12A).
Mecochiridae, which became extinct at the end of the
Cretaceous, inhabited five environments but only two of
these in the Late Cretaceous (Fig. 9A). Of those lobster
groups that became extinct by the end of the Jurassic,
Litogastroidae inhabited four total environments but only
one during the Late Jurassic (Fig. 10B). Clytiopsidae,
Chimaerastacidae, and Platychelidae each are known only
from one or two environments each and are only Triassic in
age (Fig. 9B). Eryonidae is known from four environments,
one of which is lithographic limestone (Fig. 8B). It exhibits
most of its occurrences in the latter. Thus, it appears
clear that adaptation to a wide range of substrate types
or environments was much more important for lobster
survival of extinction events than was inhabitation of coral
or other reefs. Also important for survival of lobsters was
the ability to shift environmental preferences (Fig. 13).
Every lobster group that has persisted from the Mesozoic
to the Holocene has shifted environmental preferences
over that time. Homarida have shifted from predominantly
high energy, low energy, and carbonates to low energy
environments. Palinuridae and Scyllaridae have shifted their
environmental preferences as discussed above. These three
are the predominant lobster groups in modern oceans.
Lobster Evolution
The complex pattern of family-level diversity as well as the
perceived similarity of the lobster body plan among and be-
tween infraorders originally suggested to us that individuals
within lobster groups interacted with one another, leading to
interspecific competition for resources and perhaps the even-
tual extinction of some lineages or the movement of groups
from one habitat to another to reduce such competition.
Erymidae and Nephropidae both possess fully chelate
first pereiopods but they do not seem to have interacted.
PCO (Fig. 6) shows them as distinct from one another in
their environmental preferences; thus, they probably were
not competing for environmental space. Erymidae peaked
in diversity in the Jurassic and was already declining in the
Cretaceous. Nephropoidea peaked in the Late Cretaceous,
and they still exist. Of potential importance here is the
shift of erymids into limestone environments near the end
of their range as detailed above. We hypothesize that they
may have competed with crabs or other decapods such as
anomurans for space in limestone environments; analysis
of environmental parameters of those decapod groups is
underway. Alternatively, limestone environments in general
are nearshore and subtropical to tropical in nature, all
environments preferentially impacted by extinction events
(Harries et al., 1996; Kauffman and Harries, 1996). Thus,
we hypothesize that erymids might have been affected
preferentially by the end-Cretaceous events due to their
habitat shift or due to competition from other decapods or
other animals.
Pseudochelae are unusual structures within decapods that
have evolved more than once in lobsters (Karasawa et al.,
2013) and exists today in several decapod groups including
mud and ghost shrimp. Because the movable finger occludes
with the distal end of the manus and not with a fixed finger,
the strength, utility, and efficiency of such a structure would
appear to be more limited than that of a true chela and more
than that of a simple terminal dactyl (Fig. 5B). Pseudochelae
may be useful for foraging for infaunal food resources in
soft substrata, whereas true chelae (Fig. 5A) may be better
838 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Fig. 12. Lobster groups (Table 1) by energy and environments as defined in Table 2 through time. A, Palinuridae; B, Scyllaridae. Numbers of species may
add up to more than the total number of species for the genera recorded during a given time interval because a species can occur (rarely) in multiple environ-
ments during the same time interval. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.
brillonline.com/content/journals/1937240x.
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 839
Middle
Triassic Late
Triassic Early Jurassic Middle
Jurassic Late Jurassic Early
Cretaceous Late
Cretaceous Cenozoic Holocene
Eryonidae Low Low,
medium,
carbonates
Lithographic
Coleiidae Sponge-
bearing rocks
(deep), high,
carbonates
Deep water
carbonates Lithographic,
low Low
Palinuridae Low Lithographic,
coral Low High, low Low,
carbonates Medium,
high,
carbonates
Scyllaridae Low Lithographic Low,
lithographic High, car-
bonates,
coral
Mecochiridae Carbonates Low,
carbonates Low,
carbonates Low,
carbonates
Litogastroidae Carbonates Carbonates Carbonates,
low Low Carbonates
Erymidae Carbonates,
low Coral,
carbonates,
high
Coral,
carbonates,
lithographic
Carbonates,
low Carbonates,
high
CCP Carbonates,
high Carbonates,
high
Glypheidea Carbonates Carbonates,
high Carbonates,
high,
lithographic
High, low High
Homarida High, low Carbonates,
high Low Low
Fig. 13. Predominant environments for each lobster group (comprising over 50% of occurrences in all cases). Environments as defined in Table 2. CCP, Chimaerastacidae, Clytiopsidae and Platychelidae,
analyzed together because of small numbers of genera and species and because all belong to Glypheoidea.
840 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
suited for capture of epifaunal prey. Thus, pseudochelate and
chelate forms may have been able to coexist without coming
into direct competition.
Within lobsters, the pseudochela must have been a suc-
cessful adaptation early in their history, as supported by its
presence in four families within Glypheoidea, one of which
was Triassic (Litogastroidae), two of which were primar-
ily Jurassic but ranged into the Cretaceous (Glypheidae and
Mecochiridae), and one that is extant (Neoglypheidae, two
genera). Perhaps this is because fully chelate lobsters ap-
peared somewhat later. Some extant Achelata (Palinuridae)
exhibit pseudochelae as well. Examination of the environ-
mental preferences of these families indicates little strong
overlap with Erymidae or Nephropidae, the clawed lobsters.
Rather, the pseudochelate lobster families cluster together.
This phenomenon, especially in comparison with Nephrop-
idae, appears to be because Nephropidae appear later in the
record. Thus, even if pseudochelate lobsters occupied the
same environments as chelate lobsters, they were not really
speciose at the same time. Furthermore, as indicated above,
their feeding habits may have been sufficiently different that
they were not ecological competitors. Looking at the records
of Mecochiridae and Glypheidae, the most speciose fami-
lies within Glypheoidea, and records of the chelate Nephro-
poidea and Erymidae, it seems as if the glypheoids yielded
to the two clawed lobster families in about the Cenomanian.
Thus, direct competition seems not to have been a factor.
Perhaps the escalation of fish and sharks, during which a
well-developed claw would be useful defense mechanism,
was a factor. Indeed, a major radiation of osteichthyan fish
appears to have occurred at that time (Friedman and Salan,
2012). Pseudochelate lobsters may have been victims of the
Mesozoic Marine Revolution.
It has long been hypothesized that nephropid lobsters
moved from shallower water, nearshore environments in
which they have an excellent fossil record during the Cre-
taceous, into offshore, deeper water environments, result-
ing in their relatively poor Cenozoic fossil record (Glaess-
ner, 1969; Tshudy, 2003). This pattern was documented by
Jablonski et al. (1983) in many marine invertebrate groups.
This is supported in the lobsters by the modern occurrences
of nephropid and polychelid lobsters which are largely
bathyal in depth, with some ranging into the sublittoral zone.
However, note that historical records show that the Ameri-
can lobster, for example, was littoral and sublittoral as men-
tioned above. Certainly nephropids were exapted to such
a move in terms of substrate preference, as they have in-
habited low energy siliciclastic environments for much of
their existence (generally at least 20% of occurrences). Post-
Cretaceous occurrences are mostly low energy, so the hy-
pothesis is supported by these data. Certainly we found noth-
ing to reject it. Alternatively, the nephropids may have oc-
cupied deep water habitats throughout their existence and
subsequently become extinct in the shallower parts of their
range. The same may be true for Polychelida which, al-
though with a much more sparse record, follow the same
pattern as Nephropoidea.
Lobsters radiated early and rapidly at the family level.
This resulted in explosive radiations followed by stasis. This
is punctuated equilibrium, of course, at a higher taxonomic
level. The first member of Clytiopsidae is known from the
latest Permian; by the Anisian (Middle Triassic), only 8
million or so years, six lobster families had appeared. By
the Norian (Late Triassic), 29 million years later, seven
families were present, but only two extended from Anisian
time. By the Pliensbachian, eight families were present of
which five had extended from the Norian and one carried
over from the Anisian, another 27 million years later. By
the Tithonian, 11 families were present of which five carried
over from the Pliensbachian. Radiation thus occurred until
the Tithonian, for nearly 100 million years, after which
it slowed dramatically. During the Cretaceous, only two
new families appeared, and five became extinct. Two more
became extinct during the Cenozoic, and two more appeared.
Evolution in lobsters embracing the first 100 million years
of the Mesozoic was rife with rapid evolution and turnover.
Far too little is known of evolutionary patterns of Decapoda
in the Paleozoic to speculate. By the Early Cretaceous,
stasis at the family level was achieved, with much slower
extinction and radiation rates.
Lobsters do not appear to have been strongly affected
by the major mass extinction events, and instead their
evolution and extinction was governed by other, intrinsic and
extrinsic events. The diversity curve shows a clear decline
in generic diversity at the end of the Cretaceous (Fig. 4).
The decline began in the Campanian and reached its lowest
in the Selandian (Paleocene), indicating that many forms
survived the end-Cretaceous events. Thus, the decapods
were affected by the end-Cretaceous events but in what
appears to have been a prolonged extinction event, not
a catastrophic event. Nephropidae were affected by this
decline but notably high latitude areas show little to no affect
(Schweitzer and Feldmann, 2005). Glypheidae appears to
be unaffected at the family level by the extinction event,
whereas the closely related Mecochiridae did become extinct
at the end of the Maastrichtian. Other Cretaceous groups had
already become extinct by the Coniacian (Late Cretaceous).
Thus, catastrophic hypotheses for the end-Cretaceous event
cannot explain our data.
Perhaps more puzzling is the sharp decline at the end
of the Jurassic (Tithonian). There are some reports of a
minor regional extinction event at that time in Europe of
primarily pelecypods (Hallam, 1986). This could in part
explain our data, as much of the Tithonian record of lobsters
is in fact European. In addition, much of the lobster record
of this time is overprinted by the Solnhofen-type limestone
deposits, which are well-known and well-published. Thus,
the Tithonian drop in diversity may in part be an artifact of
this diversity spike in the Tithonian caused by the Solnhofen.
As mentioned, lobsters did not particularly inhabit reefal
rocks, so reefal collapse cannot explain a drop in diversity
at the end of the Jurassic for these animals (Klompmaker et
al., 2013).
Our data show a slight decline in the Turonian, another
proposed time of extinction (Harries and Little, 1999).
However, this extinction is at the Cenomanian/Turonian
boundary, and the lobster record is impacted by another
biasing agent, the Cenomanian lithographic limestones of
Lebanon. These may inflate the Cenomanian record similar
to the Solnhofen-Tithonian relationship.
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 841
Evolutionary Faunas
Lobsters can be separated into distinctive evolutionary fau-
nas based upon their appearance, radiation, and decline over
time. Other invertebrate groups have been demonstrated to
have been arranged into evolutionary faunas. Using exten-
sive datasets, Adrain et al. (1998) demonstrated that trilo-
bites could be arranged into component faunas during the
Ordovician and Silurian. Ausich et al. (1994) have suggested
that crinoids may be divided into component faunas in the
Paleozoic. The orthid brachiopods displayed a biphasic di-
versity pattern during the Paleozoic (Harper and Drachen,
2010). Our qualitative and quantitative observations of the
Decapoda strongly suggest that there are differential domi-
nant faunal components during different time periods, com-
prising evolutionary faunas (Fig. 14).
Triassic Fauna.—The lobsters exhibit a clear Triassic radi-
ation; all of the Triassic families later became extinct. The
Triassic Fauna is distinctive, and during the Triassic Period,
families comprising the Triassic Fauna make up at least half
of occurrences in every stage. Triassic Fauna lobsters in-
clude Chimaerastacidae, Clytiopsidae, Glaessnericarididae,
Litogastroidae, Pemphicidae, Platychelidae, and Tetracheli-
dae. Of these, only Litogastroidae survived beyond the Trias-
sic, represented by one genus, Pseudoglyphea Oppel, 1861.
The Triassic fauna was numerically lower in abundance of
genera than the other two faunas, reaching a peak of 11 ge-
nera in the Anisian (Middle Triassic). The two later faunas
would be much more numerically abundant and more di-
verse.
Within the Triassic Fauna, three lobster body forms are
represented, cylindrical cephalothoraxes with chelae; cylin-
drical cephalothoraxes with pseudochelae, and dorsoven-
trally flattened cephalothoraxes and pleons with chelae
(Fig. 5A-C).
Jurassic Fauna.—Turnover into the Jurassic Fauna began in
the Late Triassic, and by the Hettangian, two-thirds or more
of the lobsters were part of this fauna until the Oxfordian,
where the percentage began to diminish to about half for
the remainder of the Jurassic. The Jurassic Fauna was com-
prised of the Coleiidae, Erymidae, Eryonidae, Glypheidae,
and Mecochiridae, representing the same three body plans as
seen in the Triassic with the addition of cylindrical or rect-
angular cephalothoraxes and achelate pereiopods (Fig. 5A-
D). This fauna decreased in abundance and diversity in step-
wise fashion, showing declines in the Turonian (Late Cre-
Fig. 14. Three lobster evolutionary faunas. Note: x-axis not scaled to length of time intervals in years. This figure is published in colour in the online
edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.
842 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
Table 3. Three lobster evolutionary faunas, their assigned families (row 2) and their assigned
body plan. Number of body plans increases with time (arrow).
taceous), Maastrichtian-Danian (Cretaceous-Paleocene), fi-
nally becoming extinct later in the Paleogene.
Modern Fauna.—The Modern Fauna appeared early, during
the Triassic, but radiated strongly during the Cretaceous,
and four out of the seven families are extant. Taxa very
similar to modern ones in the Nephropidae and Palinuridae,
even extant genera, were abundant and diverse by the Late
Cretaceous. This fauna includes the same body plans as the
Jurassic with one additional (Fig. 5). The Modern Fauna
has a particularly long tail early, demonstrating that some
members within this fauna evolved early on, only to radiate
much later.
DISCUSSION
Overall Evolutionary Patterns
The presence of three evolutionary faunas demonstrates that
as in other invertebrates and the marine fauna in general,
some lobster forms evolved early and yielded to more
numerically abundant groups. Numerical abundance and
generic abundance seem to have increased through time
(probably at least in part subject to biases in rock record
preservation). Diversity at the family level seems to have
remained high throughout the Triassic and Jurassic faunas
and diminished in the Late Cretaceous.
The fact that each evolutionary fauna yields an additional
body form is suggestive of an increase in morphological
disparity in lobsters over time, at least in the large scale,
overall morphology (Table 3). All of the lobster body plans
seen in the Modern Fauna are still extant. This may seem
contradictory: early evolution within the group yielded high
diversity at the family level, yet new body plans continued
to arise at least into the Cretaceous.
The explanation appears to be at least two-fold. One is
that early radiation and high diversity do not preclude later
evolutionary innovations. Another is that lobster body plan
differences can be examined at several scales. The body
plan differences within Glypheoidea which yield multiple
families, for example, tend to involve the presence or
absence of chelae, pseudochelae, or no chelae on pereiopods
1-4, and in which combination they occur. These patterns
in most glypheoid families are quite stereotyped. As an
example, Glypheidae always have pereiopods 1 and 2
pseudochelate, pereiopod 3 may be pseudo- or achelate, and
pereiopods 4 and 5 are achelate, as an example (Karasawa et
al., 2013). Thus, these appear to be small scale body plan
differences, but they broadly are applicable. Family level
differences are often defined on these types of differences.
Larger scale body plan differences (Table 3), include
general shape of the lobster body and pleon and the nature
of the first pereiopod or cheliped. These major body plans
seem to have been added over time. Thus, has disparity
increased in lobsters through time? Large scale body plan
changes seem to have increased over time, whereas smaller
scale variations, such as family level variations, seem to have
SCHWEITZER AND FELDMANN: LOBSTERS IN TIME 843
decreased due to extinction. The variation in claw formula as
described above is certainly smaller now.
We suggest that this indicates that lobsters have become
more specialized. Variations in the number of chelae or pseu-
dochelae are variations on a single theme. Thus, the early
body plan differences either at the large or small scale,
were based largely on differences in numbers of clawed
appendages. Moving into the Jurassic and Modern faunas,
dorsoventral flattening and a shift to achelate appendages
appears. Dorsoventral flattening of the carapace can provide
the animal with cryptic options not open to a more rectangu-
lar or cylindrical carapace. Changes to completely achelate
claws, but the ability to become cryptic or burrowing, may
have opened up more habitats for the animals. Thus, it may
be that even though the family level diversity decreased, the
level of specialization increased over time due to the addi-
tion of new body plans.
Lobsters and the Mesozoic Marine Revolution
Examination of the body plan (Table 3; Fig. 5) and their
distribution shows that lobsters evolved from mostly clawed
forms into an array of forms, clawed and achelate, flattened,
and cylindrical (Fig. 15). Lobsters may have played a role in
the Mesozoic Marine Revolution, but that role would have
been through a variety of predatory means, not just crushing
claws, as the claws of the Triassic and many Jurassic forms
were small and slender or were pseudochelate (Schweitzer
and Feldmann, 2010). The trend toward flattened forms, with
no claws, in the Cretaceous and into the Cenozoic, suggests
even more strongly than crushing claws were neither the
preferred nor the only method of durophagous predation
in the MMR of lobsters. The other methods outlined by
Schweitzer and Feldmann (2010) should be investigated.
The variety of environmental preferences exhibited by
lobsters in the Modern Fauna supports the hypothesis that
lobsters were using means other than and in addition to
crushing claws during the MMR and into the Cenozoic.
Certainly nephropid lobsters and the erymids, part of the
Jurassic fauna that had a strong presence in the Cretaceous,
had claws that could be used for crushing or capturing prey.
However, other elements of the modern fauna present in the
Cretaceous, viz., Palinuridae and Scyllaridae, do not have
claws, and Polychelidae have very slender claws. Evidence
indicates that these groups survived in other habitats, such
as deeper water, characterized by reduced predation pressure
from bony fish and sharks, the former of which radiated in
the Late Cretaceous (Friedman and Sallan, 2012). Modern
scyllarids and palinurids often live in reefs and rocky
environments, hiding in crevices (Holthuis, 1991). Modern
polychelids most often live in bathyal environments, away
from the crowded, predator dense shallow environments they
seem to have preferred in the Mesozoic. These groups may
have been victims as well as perpetrators of the MMR.
Conclusions
Based upon the phylogenetic analysis of 44 extinct and 27
extant genera and a compilation of all known species of fos-
sil lobsters, a total of 547 species arrayed in 102 genera,
patterns of diversity yield numerous conclusions. Lobster
diversity measured at the family level increased rapidly in
the early Mesozoic, reaching peaks of 16 families in the
Tithonian and 11 families in the Cenomanian; six fami-
lies are known from the Holocene. Although the generic
maxima of 29 in the Tithonian and 26 in the Cenoma-
nian are well below the 62 genera in the Holocene, at least
some of this difference can be attributed to incompleteness
Fig. 15. Lobsters grouped by nature of chelae and arrayed by time. Note that all four chela types are still extant, but chelate and achelate types dominate
the Holocene fauna. Note: x-axis not scaled to length of time intervals in years. This figure is published in colour in the online edition of this journal, which
can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.
844 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 6, 2014
of the fossil record, particularly because the fossils record
of bathyal and abyssal sediments is almost totally lacking.
Changes in diversity over time are also reflected in over-
all family-level composition that reveals three major lob-
ster faunas. The Triassic Fauna comprises Chimaerastaci-
dae, Clytiopsidae, Glaessnericarididae, Litogastroidae, Pem-
phicidae, Platychelidae, and Tetrachelidae; all but the Lito-
gastroidae became extinct by the end of the Triassic. The
Jurassic Fauna includes Coleiidae, Erymidae, Eryonidae,
Glypheidae and Mecochiridae; it arose in the Triassic, flour-
ished in the Jurassic and Cretaceous, and the last members
became extinct in the Eocene. The Modern Fauna, Neo-
glypheidae, Nephropoidea, Tricarinidae, Palinuridae, Poly-
chelidae, and Scyllaridae, was sparsely represented in the
Triassic and Jurassic rising to numerical dominance in the
Cretaceous. Each evolutionary fauna exhibits an increase in
body plan, suggesting an overall increase in faunal disparity
with time.
Diversity patterns are biased by extrinsic and intrinsic fac-
tors. Extrinsic factors, rock volume and monographic effects
among others, are suggested to affect overall lobster diver-
sity but exert little effect on relative patterns of diversity. In-
trinsic factors, including preservation potential and ecolog-
ical niche preferences, potentially affect both absolute and
relative diversity patterns.
At the level of family, lobster taxa exhibit preferences for
specific environmental energy levels interpreted from rock
type in which the specimens are preserved. Most lobster
families retained the same, or similar, ecological preferences
through time. However, Mecochiridae, Palinuridae, and
Scyllaridae inhabited low energy settings early in their
history and exploited higher energy and reefal environments
subsequently. Erymidae exhibited a broad ecological range
throughout its history.
The timing of dominance of individual family-level taxa
coupled with ecological preferences suggests that changes in
faunal composition did not occur as a result of competition
between lobster taxa nor was it a result of extinction events.
For example, differences in feeding styles resulting from dif-
ferent pereiopod architecture could have sufficiently sepa-
rated otherwise similar groups from coming into competi-
tion. Rather, predation pressure resulting from an increase
in bony fish near the end of the Cretaceous, coupled with
the apparent, but not real, appearance of extinction result-
ing from onshore-offshore habitat preference may explain at
least some of the change. Adaptation to a broad range of
environments is a predictor for survival of mass extinction
events in lobsters.
ACKNOWLEDGEMENTS
NSF grants DEB-EF-0531674 and EAR-1223206 funded this research.
L. Baltzly and S. Yost, Kent State University undergraduates, performed
data analysis for the project. G. Schweigert, Staatliches Museum für
Naturkunde, Stuttgart, Germany; M. Nose, Bayerische Staatsammlung für
Paläontologie, München, Germany; staff at the Museum für Naturkunde
Berlin, Paläontologisches Museum; O. Schultz and A. Kroh, Geological and
Palaeontological Department of the Naturhistorisches Museum Wien, Vi-
enna, Austria; M. Munt and C. Mellish, The Natural History Museum, Lon-
don, UK; P. Müller Természettudományi Múzeum, Föld-és ˝
oslénytar, Bu-
dapest, Hungary; S. Shelton and J. Martin, South Dakota School of Mines
and Technology, Rapid City, SD, USA; H. Kato and K. Komai, Natural
History Museum and Institute of Chiba, Japan; R. Lemaitre and K. Reed
(Crustacea) and M. Florence and J. Thompson (now retired) (Paleobiology),
Smithsonian Institution, United States National Museum of Natural History,
Washington, DC, USA; A. Lord and C. Franz (Paleontology), Senckenberg
Museum, Frankfurt, Germany; the late A. Dhondt, Institut Royal des Sci-
ences Naturelles de Belgique, Brussels, Belgium; J. Sklenaˇ
r, National Mu-
seum, Prague, Czech Republic; J. Sprinkle and A. Molineux, University
of Texas, Austin, TX, USA; A. Kollar, Invertebrate Paleontology, Carnegie
Museum of Natural History, Pittsburgh, PA, USA; J. Cundiff, Museum of
Comparative Zoology, Harvard University, Cambridge, MA, USA; S. Char-
bonnier, Muséum National d’Histoire Naturelle, Paris, France; M. Coyne,
Geological Survey of Canada, Ottawa, ON, Canada; and the Urweltmu-
seum Hauff, Holzmaden, Germany, all facilitated access to paleontological
or biological collections or loans from their respective institutions. T. Sny-
der and all of the staff at the Inter-Library Loan service at the KSU li-
brary assisted in finding old and obscure literature. Kent State University
at Stark funded an Undergraduate Research Assistant, E. Johnson, during
Spring 2012, academic year 2012-2013, and summer 2013. A. J. Smith, De-
partment of Geology, Kent State University, provided advice on statistical
analyses. Very helpful and constructive reviews were provided by S. Char-
bonnier, Museum national d’Histoire naturelle, Paris, France; F. Schram,
Whidbey Island, WA, USA; and J. Haug, Associate Editor, Journal of Pale-
ontology. Our sincere thanks to these gentlemen.
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RECEIVED: 11 September 2014.
ACCEPTED: 20 September 2014.
