Penny 2021
Penny 2021
1–21
DOI: 10.1017/pab.2021.3
Article
Amelia M. Penny† and Björn Kröger. Finnish Museum of Natural History, University of Helsinki, Jyrängöntie 2,
00560 Helsinki, Finland. E-mail: bjorn.kroger@helsinki.fi. †Present address: Centre for Biological Diversity,
School of Biology, University of St. Andrews, St. Andrews, Fife KY16 9TF, United Kingdom. E‐mail: amp29@st‐
andrews.ac.uk
Olle Hints. Department of Geology, School of Science, Tallinn University of Technology, Ehitajate tee 5, 19086
Tallinn, Estonia. E-mail: [email protected]
© The Author(s), 2021. Published by Cambridge University Press on behalf of The Paleontological Society. This is an Open
Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/
licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original
work is properly cited. 0094-8373/21
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2 AMELIA M. PENNY ET AL.
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 3
FIGURE 2. Overview of brachiopod localities with lithologic information used in this study. Maps show extent of outcrop
and subcrop of strata of the respective time bins. Hatched lines show facies boundaries. Maps are adopted from Paškevičius
(1997: O1, Tremadoc, fig. 41; O2, Latorp, fig. 42; O4, Kunda, fig. 43; O5, Kukruse, fig. 44; O6, Oandu, fig. 45; O7, Ashgill, fig.
47; S1, Raikküla, fig. 59; S2, Adavere, fig. 60; S4, Ludlow, fig. 62; S5, Pridoli, fig. 63) and Raukas and Teedumäe (1997: O3,
Volkhov, fig. 142). 1, brachiopod locality with lithologic information; 2, reef occurrence; 3, boundary between our “basinal,”
“platform” areas; 4, facies boundary; 5, no time-bin sediments preserved; 6, extent of time-bin sediments.
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4 AMELIA M. PENNY ET AL.
FIGURE 3. North–south cross sections showing facies distribution through the northern flank of the Baltic Basin from west-
ern Estonia in the north to western Lithuania in the south. Vertical lines mark locations of drill-core sections and the dis-
tributions of different lithologies. A, Silurian strata; based on Nestor and Einasto (1982). B, Ordovician strata; based on
Dronov et al. (2011). Roman numerals identify depositional intervals sensu Nestor and Einasto (1997).
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 5
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6 AMELIA M. PENNY ET AL.
Diversity Partitioning.—We use the term diversity of the samples that have been pooled
“richness” to refer to the number of genera (Lande 1996). In hierarchical multiplicative
(which may be a raw count or an estimate diversity partitioning, beta diversity is the
based on some form of standardization) and pooled diversity at a given hierarchical level
“diversity” as an umbrella term when referring divided by the mean diversity of its component
to diversity measures in the Hill number fam- samples (Jost et al. 2010).
ily. These include richness, but also the Shan- Additive diversity partitioning is sometimes
non and Simpson diversities, which are the preferred over multiplicative diversity parti-
Shannon and Simpson indices converted to tioning, because it yields beta-diversity values
their corresponding Hill numbers (terminology in the same units as alpha and gamma diversity
after Hsieh et al. 2016). “Alpha diversity” refers (e.g., Patzkowsky and Holland 2007; Holland
to diversity within hierarchically partitioned 2010). Holland (2010) also advocated using
categories, “beta diversity” refers to diversity three indices of diversity in parallel—richness,
difference between categories, and “gamma the Shannon index (also called the Shannon
diversity” refers to the regional diversity; each entropy), and the Simpson index (also called
may be measured with any of the three diver- the Gini-Simpson index)—to evaluate diversity
sity measures used in this study. Where we patterns in common and rare taxa and to con-
refer to alpha, beta, or gamma diversity in trol for the fact that richness is highly sensitive
only one of the diversity measures, we use the to sample size (Magurran 2004). Richness,
Greek letter with a subscript (e.g., γS is regional being a simple count of the taxa in an assem-
diversity, measured with richness; γshan and blage, gives equal weight to all taxa, and conse-
γsim would give the corresponding Shannon quently may be strongly influenced by the
and Simpson diversities, respectively). All ana- rarest taxa in an assemblage, as most of the spe-
lyses were performed at the genus level. cies in a community tend to occur at low abun-
Total (gamma) diversity can be partitioned dance (Whittaker 1970; Reddin et al. 2015). The
into alpha and beta diversity using either addi- Simpson index is most strongly influenced by
tive or multiplicative approaches. While alpha the commonest taxa in an assemblage, which
and gamma diversity have the same meanings are arguably the most ecologically important
in both additive and multiplicative diversity (Gaston 2010), while the Shannon index has
partitioning, they differ in their concepts of an intermediate sensitivity (for a full explan-
beta diversity (which is also calculated using ation, see Jost 2007; Jost et al. 2010).
a range of other methods; reviewed in Koleff However, the Shannon and Simpson indices
et al. 2003). In multiplicative diversity parti- do not behave additively and can give mislead-
tioning, beta diversity is the ratio of gamma ing results (Jost et al. 2010). The solution, pro-
diversity to alpha diversity (Koleff et al. 2003; posed by Jost et al. (2010), is to convert each
Whittaker 1972), while in additive diversity index to its corresponding Hill number (Hill
partitioning, beta diversity is gamma diversity 1973), also called its “effective number of spe-
minus alpha diversity (Lande 1996). cies,” which is the number of species corre-
Both diversity partitioning approaches can sponding to the given diversity index when
be used in hierarchical sampling schemes, the species abundance distribution is perfectly
which assign samples to progressively larger even. These values can easily be compared
nested groups to assess how much diversity is with richness, because their units are the same.
contributed at each level. Hierarchical parti- After conversion to Hill numbers, the Shan-
tioning is readily applicable to fossil assem- non and Simpson indices yield the same
blages, because hierarchical sampling mirrors within-group diversities whether partitioned
the hierarchical nature of lithostratigraphic multiplicatively or additively (Ricotta 2005;
and chronologic units (Patzkowsky and Hol- Jost et al. 2010). In this study, we used hierarch-
land 2012). In hierarchical additive partition- ical multiplicative diversity partitioning with
ing, beta diversity is the difference between Hill numbers, because it yields the same
the pooled diversity of a group of samples at within-category diversities as additive diver-
a given hierarchical level and the mean alpha sity partitioning and is easier to implement
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 7
using the R package vegan (Oksanen et al. discussion on alpha and beta diversities with
2018). However, in places we have expressed mean p-values < 0.1.
the differences in mean alpha diversity between We used environmental partitioning to
hierarchical levels as a percentage of gamma evaluate how faunal differences between lithol-
diversity; this is not strictly equivalent to hier- ogies contributed to regional diversity patterns,
archical additive diversity partitioning, but we which relates to the influence of environmental
use it as an additional, intuitive indicator of preference and substrate heterogeneity on
faunal difference between sampling units. assemblage heterogeneity. The basic sample
We used the function multipart, within the R for alpha-diversity calculation was the lithos-
package vegan (Oksanen et al. 2018), to per- tratigraphic unit, approximately equivalent to
form multiplicative diversity partitioning a formation. Formations have been taken to
using the Tsallis entropy, with scale factors 0, represent metacommunities, and hence may
1, and 2 corresponding to taxonomic richness, be ecologically meaningful units (Hofmann
Shannon index, and Simpson index. The result- et al. 2019). We used a three-level hierarchy,
ing alpha and gamma diversities are given as with lithostratigraphic units nested within
their corresponding Hill numbers; we refer to carbonate-dominated or siliciclastic-dominated
these as the richness, Shannon diversity, and lithologies, with the highest hierarchical level
Simpson diversity, following Hsieh et al. being the whole study region. We also evalu-
(2016). ated beta diversity within siliciclastic- and
Data from 177 lithostratigraphic units were carbonate-dominated lithologies by perform-
analyzed, of which 38 contain only a single ing separate two-level partitions of siliciclastic-
collection. To avoid discarding these lithostrati- and carbonate-hosted diversity. Where we sus-
graphic units, we present the raw data for pected that time binning might be affecting the
all lithostratigraphic units containing five diversity curves produced, we performed second-
occurrences or more. We include coverage- ary analyses at stage-level temporal resolution.
standardized results, which show a substan- For temporal diversity partitioning, we used
tially similar pattern, in Supplementary Mater- a three-level hierarchy to evaluate how faunal
ial 2. Coverage was measured using Good’s u turnover between stages contributed to the
(Good 1953), using a method from Alroy diversity curve. Again, we used lithostrati-
(2014) as implemented by Brocklehurst et al. graphic units as the basic units for alpha-
(2018). We coverage-standardized each lithos- diversity calculation, followed by within-stage
tratigraphic unit to a Good’s u of 0.5, a value and within–time bin diversity. Brachiopod
chosen to maximize the number of time bins occurrences lacking stage level temporal reso-
that would run. Lithostratigraphic units with lution were excluded from this analysis.
lower coverage were discarded, and those We also performed diversity partitioning
whose coverage was higher were subsampled within shelf and basin facies. Brachiopod
until the standardization coverage was re- occurrences were assigned to shelf or basin
ached. Subsampling was repeated 100 times facies based on their geographic locations; the
for each time bin, and the mean diversities R package icosa (Kocsis 2017) was used to gen-
and associated p-values from null modeling erate hexagonal equal-area grids of ∼37 km per
recorded. Increasing the number of subsamp- side, and occurrences were assigned to grids
ling runs had a minimal effect on the results. using the locate() function. The resulting grids
We used the null modeling function in multi- were used as the sampling unit and were over-
part to perform individual-based randomiza- lain on a facies map of the Baltic region to
tion, which randomly assigns occurrences to assign them to the shelf or basin facies for ana-
samples at the lowest level of the hierarchy lysis. Comparison of shelf and basin diversity
and assesses whether the diversities in the sam- patterns covers the time interval from bin O6
ple are significantly different from those of the onward, as this is when the platform differenti-
randomized data. Null models were run with ation phase began (Fig. 4).
99 iterations for each subsampling run, and The meanings of different diversities within
when reporting results, we focus our the partitioning schemes are summarized in
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8 AMELIA M. PENNY ET AL.
Alpha 1 (α1) Beta 1 (β1) Alpha 2 (α2) Beta 2 (β2) Gamma (γ)
Temporal Mean diversity The mean value of alpha 2/ Mean Gamma/mean Regional
partitions within mean alpha, across all diversity alpha 2. Reflects diversity
lithostratigraphic stages in a time bin. within faunal difference within a
units Reflects faunal difference stages in a between stages time bin
between lithostratigraphic time bin in a time bin.
units within stages.
Lithologic/ As above The mean value of alpha2/ Mean Gamma/mean As above
environmental mean alpha, across both diversity alpha 2. Reflects
partitions lithological categories. within faunal difference
Reflects faunal difference lithologic between
between lithostratigraphic categories in lithological
units within lithological a time bin categories in a
categories. time bin.
Table 1. Alpha, beta, and gamma diversity are Occupancy as an Abundance Proxy.—In our
referred to by their Greek characters, with sub- study, direct brachiopod abundance data were
scripts denoting richness, Shannon diversity, or not available. We therefore use the number of
Simpson diversity. localities with occurrences as a rough proxy of
Independent Gamma-Diversity Estimates.—In abundance with the a priori assumptions that
the hierarchical diversity partitioning analyses, detection in the fossil record is, at least in
coverage may vary between time bins. To esti- part, dependent on abundance and that
mate the richness trajectory through time, we more-abundant species have higher occupan-
used independently calculated richness curves cies (see e.g., Gaston et al. 2000).
produced using two methods. First, we used
the estimateD function in the R package
Results
iNEXT (Hsieh et al. 2016), standardizing to a
coverage of 0.7, which is the maximum cover- Regional Richness (γS) over the Ordovician–
age at which time bins O3–S5 will run. Second, Silurian.—The GOBE and LOME are expressed
we used the capture–recapture (CR) modeling in the regional richness patterns derived from
approach (Nichols and Pollock 1983; Liow CR (Fig. 5). With stage-level resolution, γS
and Nichols 2010) by fitting the Jolly-Seber increases in two pulses, one in the Floian–
model following the POPAN formulation early Dapingian (Billingen–Kunda regional
(Schwarz and Arnason 1996; see also Kröger stages), with a second, larger pulse in the Sand-
et al. [2019] for details of the method). bian. The γS decline during time bin O7 is an
We also produced coverage-standardized effect of the time-binning strategy used, and
and CR modeling approach richness curves with stage-level resolution, γS continues to
at stage-level temporal resolution to visualize increase up until the Porkuni (Fig. 5) and then
how the time binning might influence abruptly declines into the Silurian, with only
richness trajectories, excluding stages without a limited recovery. The LOME is not preceded
occurrences or where the estimateD function by a diversity loss and instead represents a rela-
suggested large prediction bias. Because tively abrupt diversity decline between the Por-
different methods can be expected to produce kuni and Juuru regional stages.
different richness estimates, we focus our Temporal Taxonomic Turnover.—Taxonomic
discussion on trends in the data rather than turnover between regional stages, β2 in our
absolute values. To determine whether temporal partitioning scheme, is a major con-
carbonate-dominated and siliciclastic-dominated tributor to within-bin diversity, though its
lithologies showed the same diversity impact is highly dependent on the length of
patterns, we calculated independent coverage- time bins and the diversity index used
standardized richness curves for each lithologic (Fig. 6). The Sandbian–Katian (time bin O6) is
category. particularly strongly influenced by this effect
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 9
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10 AMELIA M. PENNY ET AL.
TABLE 2. Taxonomic turnover between stages (β2) measured as a percentage of γ diversity and the associated p-values. β2
was measured using richness, the Shannon diversity, and the Simpson diversity. The values used to produce this table are
given in Supplementary Material 3A. Data where p-values were below 0.1 are highlighted in gray. NAs denote time bins
with insufficient data for analysis, or where bins were not subdivided into multiple stages.
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 11
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12 AMELIA M. PENNY ET AL.
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 13
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14 AMELIA M. PENNY ET AL.
TABLE 4. Beta diversity within carbonate- and differences between the regional Keila–Vormsi
siliciclastic-dominated lithostratigraphic units, measured
using richness. Full values used as a basis for this table and stages, which are unconformity-bounded units
the Shannon and Simpson diversities are given in on the Estonian and Lithuanian shelves. The
Supplementary Material 3C and 3D. Beta diversities with base Oandu, base Rakvere, and base Nabala
p-values lower than 0.1 are highlighted in gray. “NA”
denotes time bins with insufficient data for analysis, or stages are karstic over large areas of the North
containing only a single lithostratigraphic unit in the Estonian shelf (Calner et al. 2010), contrasting
relevant lithological category. with the more complete Livonian basin sec-
Carbonate-dominated Siliciclastic-dominated tions. Although exceptionally high turnover
rates from this interval are not known from glo-
β1S β1S p-value β1S β1S p-value
bal (Kröger et al. 2019) or
C NA NA NA NA
O1 NA NA 1.17 1.00
paleocontinental-scale (Franeck and Liow
O2 NA NA 2.00 0.01 2019) analyses, brachiopod faunas of this inter-
O3 1.85 0.99 NA NA val on the Estonian shelf cluster into highly dis-
O4 3.79 0.01 NA NA
O5 5.03 0.01 1.36 0.59
tinctive time groups, while Livonian basin
O6 6.31 0.01 4.19 0.01 brachiopod assemblages are not as distinct
O7 5.68 0.01 1.80 0.03 (Hints et al. 2018).
S1 2.94 0.01 NA NA
S2 NA NA 1.76 0.95
Immigrations from Avalonia (increasingly
S3 3.19 0.01 2.75 0.85 from the early Sandbian onward) and Lauren-
S4 1.94 0.15 3.54 1.00 tia (increasingly from the late Sandbian
S5 1.64 0.97 1.43 0.01
onward) also contributed to temporal turnover
during O6 (Hints and Harper 2003; Hansen
a Late Ordovician diversity peak and a diver- and Harper 2008; Rasmussen and Harper
sity crisis across the Ordovician/Silurian 2011b; Rasmussen et al. 2012). They occurred
boundary. These patterns are roughly consist- alongside changes to sediment-transporting
ent with global and regional richness curves sea currents within the Baltic paleobasin (Kiipli
for the GOBE (e.g., Trubovitz and Stigall et al. 2008, 2009) and reflect the regional “mid-
2016; Colmenar and Rasmussen 2017; Hints dle Caradoc facies and faunal turnover” (Mei-
et al. 2018; Rasmussen et al. 2019) and the dla et al. 1999; Ainsaar et al. 2004). Temporal
LOME (e.g., Sheehan 2001; Harper et al. 2013; turnover also makes a large contribution to
Rasmussen 2014), including in the Baltic paleo- apparent diversity in time bin S3, which
basin (e.g., Nestor et al. 1991; Rasmussen et al. includes two major regional and global biotic
2007, 2009; Kaljo et al. 2011; Hints et al. 2018). turnover events (Ireviken and Mulde events;
The hierarchical partitioning schemes and mul- Calner 2008).
tiple diversity indices used here allow us to dis- The amplitude and frequency of sea-level
sect the roles of environmental heterogeneity change can influence evolutionary rates at glo-
and temporal turnover in generating these bal and regional scales. Globally and region-
regional patterns. ally, high origination and extinction rates
Impact of Temporal Turnover on Ordovician– cluster at sequence boundaries in shallow-
Silurian Diversity Patterns.—Temporal turnover water environments, and contrastingly at max-
within time bins has a major impact on the imum flooding intervals in deeper-water envir-
regional richness curve for the eastern Baltic onments (Holland 1995, 2020; Holland and
paleobasin (Fig. 6A) (an effect also noted by, Patzkowsky 2002). Sea-level changes can act
e.g., Kröger and Lintulaakso [2017]). The as a global diversification driver in marine eco-
impact is largest in time bin O6 (late Sand- systems, because sea-level falls can isolate mar-
bian–early Katian) where the total richness ine communities for long periods of time until
strongly exceeds the richness estimates of the sea-level increases allow them to disperse (Sti-
respective regional stage bins (Fig. 5). The prin- gall et al. 2017). In the Baltic paleobasin,
cipal source of temporal substrate changes in repeated cycles of sea-level rise and fall during
the Baltic paleobasin is sea-level change (e.g., the Middle Ordovician have also been sug-
Lazauskiene et al. 2003; Harris et al. 2004), gested as a contributor to brachiopod diversifi-
and bin O6 coincides with high faunal cation through the same process (Pedersen and
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 15
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16 AMELIA M. PENNY ET AL.
carbonate hardgrounds has been suggested as processes generated the conditions for the
a diversification driver in hard substrate taxa development of tropical carbonate platforms
(e.g., Taylor and Wilson 2003; but see also and facies mosaics in the Baltic paleobasin as
Franeck and Liow 2020). In the Paleozoic, the well as in Laurentia and South China (see earl-
global balance between the extent of carbonate ier discussion). Hence, as well as releasing
and siliciclastic substrates may also influence environmental constraints, global cooling dur-
the global balance between extinction and ing the Ordovician combined with other
origination (Foote 2006), and the substrate regional-scale environmental changes to
affinities of benthic clades (Miller and Con- enhance biodiversity through intensified bio-
nolly 2001). While regional in scale, our logically mediated carbonate production and
results support the hypothesis that the the development of heterogeneous carbonate
expansion of shallow-marine carbonate environments.
environments in the Ordovician enhanced Regional Response to the LOME.—A global
regional diversity by enhancing substrate signature of the Ordovician–Silurian transition
heterogeneity in the forms of both differenti- is a general shift from highly differentiated to
ation between shelf and basin and finer-scale cosmopolitan faunas, including in brachiopods
differentiation within carbonate (e.g., Sheehan and Coorough 1990; Darroch
environments. and Wagner 2015). The end-Ordovician extinc-
Impacts of Broadscale Environmental Changes: tion led to a global decline in provincialism,
Climate and Paleogeography.—Simultaneous with genera lost during the extinction replaced
diversification in both siliciclastic- and by those dispersing from elsewhere (Sheehan
carbonate-dominated environments suggests 1975; Krug and Patzkowsky 2007; Finnegan
broadscale, generalized diversification driver et al. 2016; Congreve et al. 2019; Penny and Krö-
(s) affecting both types of lithologies. One of ger 2019; Rasmussen et al. 2019). Sheehan
these drivers was repeated sea-level oscilla- (1975), discussing North America, and Ras-
tions, which promoted allopatric speciation in mussen and Harper (2011b), discussing the glo-
brachiopods (Lam et al. 2018; Stigall 2018; bal record, both suggest that the LOME led to
Pedersen and Rasmussen 2019); the long-term, the loss of shallow-marine specialist brachio-
global climatic cooling trend was another pod genera, which were replaced by assem-
(Rasmussen et al. 2019). blages of presumably more eurytopic genera
At a global scale, Ordovician climatic cooling with broader geographic and depth
probably released environmental controls on distributions.
the growth of skeletal organisms, allowing This is also evident in the eastern Baltic
increased diversification (e.g., see Trotter et al. paleobasin. Notably, mean alpha diversity
2008; Kröger 2017; Edwards 2019; Rasmussen (i.e., diversity within lithostratigraphic units)
et al. 2019). The shift of Baltica toward the tro- is largely unaffected by the extinction, and
pics brought regional warming but kept the instead the regional diversity decline princi-
Baltic paleobasin within this optimal tempera- pally consists of a reduction in faunal differen-
ture range (Cocks and Torsvik 2005; Dronov tiation between lithostratigraphic units and
and Rozhnov 2007). carbonate- and siliciclastic-dominated litholo-
The development of a carbonate platform in gies after the Hirnantian, alongside a decline
the Baltic paleobasin and the impacts on in richness at platform localities (Figs. 6A and
regional diversity were ultimately a result of 9). In this context, the particular role of Baltica’s
Baltica’s plate-tectonic drift toward the tropics. late Katian–Hirnantian deeper-water brachio-
But time-specific climatic constraints within the pods, which became important during Silurian
tropical realm, namely the intensity of ocean recovery in shallow-water habitats and on
currents and levels of global SST, oxygenation, Laurentia, needs to be emphasized (Rasmussen
and pH, controlled the details of how this plat- et al. 2010, 2012; Rasmussen and Harper 2011a,b;
form accommodated, which organisms pro- Harper et al. 2013). This study corroborates
duced the carbonate sediments, and the the finding of these previous works that
characteristics of their facies mosaic. These the evolutionary postextinction shift in
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EARLY PALEOZOIC BENTHIC DIVERSITY, BALTIC BASIN 17
brachiopods had simultaneous paleogeo- modulated by sea level, niche breadth, and
graphic and paleoecological dimensions, with environmental stressors. Although the Baltic
shifting habitats among clades. paleobasin retained its heterogeneous car-
Globally, the biotic recovery from the LOME bonate shelf environment through the Silur-
was protracted, and Silurian global richness ian, an increase in the dominance of
never returned to Ordovician levels (Rasmus- brachiopods with broad environmental and
sen et al. 2019). This global pattern is also geographic distributions may have reduced
reflected in the Baltic paleobasin (Figs. 5, 6A). the impact of habitat heterogeneity during
As noted in previous studies (e.g., Harper the Silurian recovery from the LOME.
et al. 2015; Harper and Hints 2016; Hints et al.
2018), the regional faunal homogenization
Acknowledgments
was long-lived, and while regional diversity
recovered during the early Silurian, it never We thank L. Hints and L. Ainsaar for con-
returned to Ordovician values. In the Silurian, structive discussion during development of
while the carbonate platform persisted in the this study, U. Toom for help with locating rele-
Baltic paleobasin, faunal differentiation vant literature, S. Scholze for data entry into the
between lithostratigraphic units had a persist- Paleobiology Database over the course of this
ently reduced impact on regional diversity, study, and D. Matthews for assistance with
with generally more similar brachiopod assem- compiling lithologic data. This paper is part
blages occupying siliciclastic and carbonate of the project “Ecological Engineering as a Bio-
environments. The Silurian decline in faunal diversity Driver in Deep Time,” funded by the
differentiation between lithologies, combined Academy of Finland, and is a contribution to
with the rise of more cosmopolitan brachiopod the IGCP program 653 “The Onset of the
assemblages, suggests that habitat heterogen- Great Ordovician Biodiversification Event.”
eity became a less important driver of regional O.H. acknowledges support from the Estonian
diversity than it had been in the Ordovician. Research Council grant PRG836.
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