John C.

Barry a, Morgan a, Lawrence J

Louis L. Jacobs b, Lindsay ‘, S. Mahmoo

Abstract

The fluvial Neogene Siwalik formatic$ns of northern Pakistan contain a long and richly fossiliferous sequence of
terrestrial vertebrate faunas in which patterns of fauna1 turnover and changes in diversity can be documented and
analyzed for intervals having durations of 0.5 m.y. The complete sequence extends from circa 18.5 to 1 Ma, but the
part between 18.5 and 5.5 Ma is best sampled, and most intervals within it are well represented
Thirteen orders of Siwalik mammals have been identified, with well-sampled intervals having 50 or more species.
Most Siwalik mammals, however, are either rodents or artiodactyls. Bovids arc the most common and most speciose
of the larger mammals, while murid and “:;r*icetid”rodents dl~minate the small mammal assemblages.
Between 18.5 and 5.S Ma species diversity varied considerably. Among artiodactyls and rodents the number of
species first increased between 15 and 13 Ma and then fell. Data on stratigraphic ranges of rodents and artiodactyls
show that fauna1 change in the Siwaliks v~asepisodic, occurring during short intervals with high turnover followed
by longer periods with considerably less change. Maxima of first appearances occurred at approximately 13.5 and 8.5
Ma, while maxima of last occurrences were at 12.5 and 8.0 Ma.
Some of the observed faunai events can be correlated to climatic and environmental changes. The Middle Miocene
diversification occurred during a period of global cooling, while the latest Miocene decline in diversity and increased
turnover accompanied oxygen rrr;:dcarbon isotopic changes that correlate to globally increasing seasonality and
aridity. Other correlatir\nh are ambiguous. The marked decrease in diversity and the major turnover events between
13 and 8 Ma do not correspond to known local or global events.
The Neogene Siwaliks and Paleogene Bighorn-Crazy Mountains sequence in Wyoming and Montana share many
similarities. They have equivalent levels of temporal resolution and similar levels of completeness of their fossil
records. Siwalik ordinal abundance and diversity patterns differ markedly from those of the Paleogene, but generic,
and probably species, diversity was approximately the same, although the Siwahk faunas may have been slightly less
diverse. Over time, changes in diversity were of comparable magnitude, with monotonic trends persisting for more
than 5 million years. The magnitude of fauna1 turnover was also similar, ranging from less than half to 3.5 times that
expected. In both sequences fauna1 change appears to have been episodic, with strong pulses between intervals of low
turnover. The Siwaliks, in contrast to the Paleogene sequence, may have had more distinct pulses and longer intervals
between pulses. Neither sequence has peaks of first occurrences coinciding with peaks of last occurrences.

003 l-0182/95/$9.50 0 1995 Elsevier Science B.V. All rights reserved

SSDI 0031-0182(94)00112-X
1. Introduction
Terrestrial deposits are notoriously incomplete
and discontinuous, making study of fauna1 change
in ancient mammal communities difficult because
of uncertainties about the ages and even the order
of fauna1 events. The Neogene Siwalik forma-
tions of Pakistan, however, contain thick sedi-
mentary sequences with good stratigraphic control
and abundant fossils, making it possible to docu-
ment fauna1 changes in Siwalik communities with
increased temporal resolution. In this regard, the
Siwaliks are similar to the Paleogene formations
of the Bighorn and Crazy Mountains Basins in
Wyoming and Montana, and both records present
opportunities to document and compare patterns
of fauna1 turnover and change in terrestrial eco-
systems over intervals of ten to twenty million
years.
In this paper we discuss the patterns of fauna1
turnover and species richness (or diversity as we
will generally refer to it throughout this paper) in
the Neogene Siwalik record, and make comparison
to the patterns documented in the Paleogene
Bighorn and Crazy Mountains Basins by Maas
et al. (this issue), Our principal objective is to
provide information with which to compare the
evolution of the mammals and terrestrial eco-
systems in the early and late Cenozoic, but we are
atso concerned with whether the apparent patterns
are simply artifacts of a biased, incomplete record.
In the following we first discuss the background
and methods of our study. Second, we describe
the quality of the Siwalik fossil record and the
patterns of change in species richness and turnover,
using our estimates of record quality to weigh the
significance of the observed patterns. We then
discuss possible correlations between Siwalik biotic
events and tectonic and climatic events, and end
by comparing the Meogene Siwa!ik and Paleogene
Bighorn-Crazy Mountains sequences.
There is a very large literature on Siwnlik mam-
mals, beginning with publication of discoveries in
the mid4830’s by British colonial ofhcers (e.g.,
Cautley and Falconer, 1835; Baker and Durand,
1836). These earliest works were primarily con-
cerned with the description of taxa, as were the
later publications of Falconer and Lydekker.
Pilgrim ( 1910, 1926) and later Colbert ( 1935)
commented on some of the large scaie evoiutionary
patterns of Siwalik mammals, but they too princi-
pally focused on phylogenetic and biogeographic
patterns. It is only recently that questions concern-
ing species diversity and the dynamics of fauna1
turnover have been addressed. Important conclu-
sions of recent studies include recognition that
Miocene fauna1 change in southern Asia was epi-
sodic, occurring as brief periods of high turnover
separated by intervals of considerably less change,
and recognition that pulses of increased extinction
did not necessarily accompany pulses of increased
first appearances ( Barry et al., 1990, 1991). Large
differences in species richness and relative abun-
dance between stratigraphic levels were also noted
in those studies and by Jacobs et al. ( 1990). Barry
et al. ( 1991) also documented trends in body size
increase among ruminant artiodactyls.
The geologic setting and important taphonomic
biases of the Siwalik formations are discussed in
this issue by Badgley and Behrensmeyer, Badgley
et al., Behrensmeyer et al., and Willis and
Behrensmcyer. Here it is only necessary to note
that the formations are very thick and comprise
alternating sandstones and fine grained sediments,
with typical fluvial environments such as channels,
crevasse splays, oxbow fills, and floodplain soils.
The Siwalik fossil accumulations have been inter-
preted as attritional assemblages that may combine
taxa of several distinctive local communities
( Badgley, 1986; Behrensmeyer, 1987). The verte-
brate fossils typically occur as modest concen-
trations of disarticulated bones, but low density
accumulations and isolated bones are also found.
Fossils are most common in the fill of the small
channels and the crevasse splays, and because these
facies become rare after circa 7 Ma ( Willis, 1991)
the evolving depositional system limits our ability
to determine stratigraphic ranges in the younger
parts of the sequence. For this reason we restrict
our analysis to sediments older than 5 Ma.
Thick deposits of Neogene Siwalik sediments
are known throughout India and Pakistan. Our
specific study area is on the Potwar Plateau in
northern Pakistan where fossiliferous sediments
are especially well exposed. Consequently, relative
stratigraphic positions of fossil sites are easily

1985), and Kappelman ( lYS4) solely on the basis
of superposition. On any single magnetic polarity
timescale, resolution of our age estimates typically
approaches 200,000 years or less. IIowever, since
differences between magnetic timescales are often
greater, resolution in an absolute time framework
is typically closer to 500,000 years. In this paper,
we use the Berggren et al. ( 1985) time scale and
our age estimates range from 100,000 to 400,000
years older than those based on the Zankin~n ;~d
Dalrymple ( 1979) time scale.
Johnson et al. ( 1982, 1985) estimated the base
of the Potwar Plateau Siwalik sequence to be older
than 18.3 Ma, and the top to be less than 0.6 Ma.
In this nearly 18 m.y. interval, thirteen orders of
fossil mammals have been identified. Despite the
ordinal diversity, however, most Siwalik mammals
belong to just three orders: rodents, artiodactyls,
and perissodactyls. Among the larger mammals,
ruminant artiodactyls and equids are the most
common and have the most species, while murid
and “cricetid” rodents dominate the small
mammal assemblages in both number of specimens
and number of species. Siwalik ruminants include
tragulids, “giraffoids,” and bovids. “Cricetids”
comprise five families or subfamilies ( Flynn et al.,
1985), three of which are now extinct, while one
is now exclusively African, These Siwalik ordinal
abundance and diversity patterns contrast strongly
with those of the Paleogene and are a consequence
of both Neogene radiations and extinction of
Paleogene groups. Temporal ranges for Siwalik
ruminants, murids, and “cricetids” are listed in
Barry et al. ( 1991) although some changes to that
listing are incorporated here.
Biogeographically, the Siwalik faunas have been
interpreted as representing a distinct, isolated pre-
cursor of the modern I~~domalayan province of
southern and southeast Asia (Bernor, 1983, 1984;
Barry et al., 1985; Thomas, 1985). Throughout its
history the degree of isolation has varied, and four
phases of isolation and interchange can be recog-
nized in the Siwaliks. The first phase is one of
isolation, which was most complete in the earliest
s ancl tragulids.
a second phase, which encompasses
ma1 exchange with northern Eurask and Africa.
ween circa 18 and 7 a, these and other taxa
diversified, with many of the resulting lineages
110~ being found only in the modern Indomalayan
realm. This third phase includes the Si~~~alikfauna1
zones of Pilgrim ( 19 13) and earlier autl+is. A
fourth phase can also be recognized following a
fauna1 turnover after 7.5 Ma. This Late Miocene
turnover made the Siwalik faunas more simila: to
r::\nrempor~neo11~faunas in northern and western
Eurasia and suggests a southeastward retraction
of the Indomalayan province onto peninsular
India. The third and fourth phases are the focus
of this paper.
While there are no significant fossil floras in
Pakistan, Miocene floral assemblages from north-
western India have been interpreted as showing
the Pliocene replacement of lowland semi-
evergreen forests by drier, more open forests
( Vishnu-Mittre, 1984; Prakash, 1973). similarly,
Quade et al. (1989) interpreted paleosol carbon
and oxygen isotope data from Pakistan to indicate
the existence of closed canopy forest before 7.5
Ma and increasingly open woodland and extensive
grasslands afterwards. These authors attributed
Late Miocene vegetational chmges to the inception
or intensification of the Indian monsoon, but
recent interpretations (Cerling et al., 1993; Quade
and Cerling, 1995) emph, I 1 decreasing atmo-
spheric CO2 as a contribc ~ng or alternative
factor. These Late Miocene vegetational changes
undoubtedly influenced the Late Miocene fauna1
turnover documented in Pakistan.
2. Materials and methods
In :his paper we USCstratigraphic range data for
artiodactyls and rodents, as well new data on total
mammal diversity of selected intervals, to discuss
fauna1 turnover in the Siwaliks. These data
(Table 1 ), which are updated and expanded over
previous studies ( Barry et al., 1990, 1991 ), are
Table 1
Number of faur~ul turnover events a:nd estimates of diversity
in Siwalik mammal assemblages. First occurrences ( Nf). last
occurrences (N,). wly occurrences (N,, ). SKI found bcforc.
during, and after (_Vbda),range through tnsa (N,,). maximum
species richness (iv,,,,, ), and standing species richness ( N,,) 14.0 7 I I 4 15 10.5
14.5 0 0 0 4 7 7.0
Age hrf N, No 15.0 1 1 0 3 8 7.0
(Ma) 15.5 0 1 0 3 8 7.5
16.0 0 0 0 6 8 8.0
Rodiwts 16.5 0 0 0 4 8 8.0
5.5 0 1 3 0 (I 4 2.0 17.0 3 0 0 4 8 6.5
6.0 0 0 n 0 1 1 1.0 17.5 0 0 1 0 5 6 5.5
6.5 1 ‘0
0 0 0 3 1.5 18.0 0 0 0 0 5 5 ‘i 5.0
7.0 2 7 2 c 0 11 5.5 18.5 5 0 1 0 0 6 3.0
7.5 3 i 3 1 3 I(! 7.0
8.0 I 5 3 3 1 13 8.5
8.5 6 3 4 7 1 15 9.0
9.0 0 6 1 5 0 12 8.5 used to determine: the number of species present
9.5 0 0 0 0 II II 11.0
within intervals, as well as to calculate rates of
10.0 3 5 1 8 0 17 12.5
I’;;lLiiWl t Ui~ilOWr. The stratigraphic data are based
10.5 1 3 1 12 0 17 14.5
11.0 z 1 I 13 0 17 15.0 on collections madb since 1973 as part of an
Il.5 0 0 0 z 12 14 14.0 ongoing research program. Our identifications are
12.0 1 4 0 12 1 IS 15.5 mostly at the species level. But, since our most
12.5 0 7 0 13 4 24 20.5
reliable data set includes only rodents and artio-
13.0 3 4 1 21 0 20 25.0
13.5 11 4 1 14 0 30 22.0 dactyls, which typically comprise about 60% of
14.0 I 1 1 15 I IO 17.5 the species in any stratigraphic interval, the data
14.5 5 1 1 II 1 19 15.5 encompass only part of the total mammal fauna.
15.0 0 0 0 0 13 13 I3.0 Thus our species-level data are not strictly compa-
13.5 7 I I to 1 1s 13.5
‘) rable to the more inclusive genus-level Paleogene
16.0 i 0 0 IO II! 12.0
16.3 3 0 0 0 0 I2 IO.5 data of Maas et al. ( 1995). Perissodactyls, and
I’/,0 4 0 1 4 I 10 7.5 especially equids. are also rich in species, but are
17.5 0 0 0 0 s ‘S 5.0 not included because of the difficulties of identi-
_
18.0 0 0 0 0 2 5 5.0 fying their species from fragmentary material.
18.5 5 0 2 0 0 7 Z.S
Because the data are most robust, we restrict our
analysis to the time period between 5.5 and 18.5
5.5 2 0 0 3 6 4.5 Ma. As in previous studies, we have divided this
6.0 2 0 0 5 7 6.0 period into half-million year intervals. Some of the
6.5 3 0 3 4 IO 8.5 terms used in our discussion are defined and
7.0 3 I 8 I 14 11.5
7.5 I 0 9 3 illustrated in Fig. 1.
13 12.0
8.0 2 1 11 0 15 13.0 Fauna1 turnovers are the result of local extinc-
8.5 3 0 12 0 16 14.0 tion, immigration, anagenetic evolution, or clado-
9.0 0 1 IO 1 16 13.5 genesis. Immigrant species are those that dispersed
w 3 0 6 3 I3 11.o from other geographic regions, wilile in situ specia-
IO.0 3 0 4 5 14 11.5
IO.5 I 0 3 7
tion, whether anagenetic or cladogenetic, took
13 11.5
11.0 5 2 Ir) 0 I8 14.0 place within the region. Because in situ speciations
11.5 1 0 13 3 16 IS.5 and immigrations may not be distinguishable,
12.0 I I 13 2 18 16.5 we have not attempted to separate them and
12.5 0 I 11 3 17 15.5
7
treat them all as “first appearancss.” Preliminary
13.0 1 0 II I5 14.0
13.5 I 1 12 0
studies, however, indicate that nearly half of all
16 14.0
Siwalik first appearances are probably immigration
 Before
Interval
Only Occurrences
FirstOccurrences
Last Occurrences
Before,During,After
RangeThrough
Fig. I. Definitions of categories of Uaunal occurrences and nomenclature
events, while anagenetic and cladogcnetic
tion are less common.
Completeness, the fraction of the original living
assemblages preserved in the fossil assemblages,
critically affects inferences we might draw from
the fossil record. That is, taphonomic factors
influence how abundant fossils are at localities,
and localities or horizons with abundant fossils
ordinarily have more taxa than those with fewer
( Badgley and Gingerich, 1988; Badgley. i 990).
Thus to some unknown extent, taphonomic pro-
cesses determine how well fossil assemblages record
original diversity and how accurately the magni-
tude and timing of changes in diversity can be
estimated, Here we use two methods to assess the
completeness of the Siwalik record. The first is a
modified version of the interval-quality scores used
by Barry et al. ( 1990), where each 0.5 m.y. interval
is graded on criteria that include the number of
specimens, number of localities, and preservation
of the fossils. In this paper we use seven classes,
with Class 0 having no data and Class 6 the best.
Because of differences in taphonomy and collecting
techniques, the rodents and artiodactyls are cval-
uated separately. The rationale and limitations of
this method were discussed by Barry et al. ( 1990).
As discussed below, we consider only ClasseL; 4
through 6 to be reliable, although Classes 1
through 3 also provide some useful information.
The interval-quality method has the advantage
During After
Interval Interval
N max
1 Nbda
1 N,., I baN
used in text. A’= present. O= absent.
evolu- of using several different kinds of information, but
the disadvantage of being sequence specific. since
scores for intervals in the Potwar cannot be com-
pared directly to scores from the Bighorn-Crazy
Mountains sequence. However, two other mea-
sures of completeness (Cl, and CY2)overcome this
limitation to some extent. Both involve calculating
an index using an estimated or expected diversity
and some measure of observed or recorded diver-
sity ( Krause and Maas, 1990; Maas et al., 1995).
For CI,, the index used by Krause and Maas
( 1990), recorded diversity (IV, ) is defined ‘as the
number of species that have actually been found
in the interval (first and last occurrences, taxa
known only in the interval, and species known
before, during, and after), while estimated diversity
is derived from the sum of all species known in
the interval (N,) plus those known before and
after, but not during the interval (A$,). (See Fig. 1 )
??
Thus:
We also calculate a second index of completeness
(C12), in which expected diversity is the number of
taxa known in intervals before and after the
interval of interest (i.e. the subset of N, + N,, from
which first, last, and only occurrences have been
excluded), while recorded diversity (Nbda) is the
number of species that are known not only before
and after, but also during the interval ( Fig. 1).
Thus, the expected value is the sum of Nbdiland
N,, and the index is the percent of the expected
taxa that are actually found in the interval, or:
This index is more conservative because it gives
lower values than CI,, although the two indices
track each other. Because C12 has expected values
that are independent of the observations, confi-
dence intervals can be placed on the estimates.
This allows sample-size effects to be taken into
consideration in judging how good any specific
estimate might be, which is not possible with CI,.
The Cl2 confidence intervals can be determined
using the binomial distribution (Simpson et al.,
1960), assuming that taxa in older and younger
intervals should also be present in the interval of
interest and that the discovery of a taxon in the
interval is independent of the discovery of other
taxa. We have calculated the upper and lower
bounds using formulas given by Ehrenfeld and
Littaucr ( 1964, p. 373).
Maximum species richness or di~versity(N,,,, ) is
the total number of taxa that actually or poten-
tially occur in each interval ( Fig. 1). This is calcu-
lated using the range-through method ( Roltovsky,
1988; Barry et al., 1990). which assumes that taxa
occurring in older and younger intervals were also
present through the intervening interval. This
method also a~s~l~~s that all extinctions occur at
the end of the interval and all appearances at
the beginning, so that it may count taxa as
co-occurring in an assemblage that actually did
not overlap in time. Following Maas et al. ( 1995).
we have therefore also calculated standing richness
as an estimate of diversity in the middle of the
interval. Standing richness (N,,) is the number
of taxa known before and after the interval
M&da+ N,,). plus the sum of first occurrences. last
occurrences, and only occurrences, divided by two,
or:
(Our formula differs from that used by Maas
et al. ( 199% because in Table 1 we tabulate only
occurrences separately from the values for fj’rst
and last occurrences.) This method assumes that
half of the first and last occurrences lie before the
midpoint of the interval, which will only be true
if the occurrences arz evenly distributed through-
out the interval (Maas et al., 1995) or at least
distributed symmetrically about the midpoint. The
estimate will bc off, however, if the distributions
of first or last occurrences are markedly asymmetri-
cal about the midpoint. The method also assumes
that only occurrences have a duration of approxi-
mately one half interval, and are symmetrically
distributed.
Fauna1 turnover is calculated from the number
of first and last occurrences in each interval (only
occurrences are counted as one first and one last
occurrence). Turnover may be expressed simply as
the number of events or as rates adjusted for
interval taxonomic richness or duration (Webb,
1969). Gingerich ( 1987) has proposed use of a
rate quotient (RQ). calculated as the ratio of
observed to expected turnover, where expected
turnover is derived from multiple regression of the
number of events on richness and duration (see
also Maas et al., 1995), Because our intervals are
of equal duration, neither the maximum nor the
standing diversity need to be adjusted for interval
Icngth. The problem of interval richness is dis-
cussed further below.
The apparent ages of first and last occurrences
are strongly distorted by the quality of the fossil
record, with intervals of poor quality making some
last occurrences seem too old and some first occur-
rences too young. To correct for this bias, we have
extended the observed stratigraphic ranges. The
method and its justification are discussed in Barry
et al. ( 1990). As in that study, when extending a
species’ range, we assume; ( 1) that the potential
range can be determined from our estimates of
data quality, and; (2) that the probability of
finding the first (or last) occurrence is equal for
each additional interval within the potential range.
This procedure is conservative with respect to our
inferences and conclusions, in that it underesti-
mates the magnitude of first and last occurrence
maxima and minima. It also gives the best visual
expression to the uncertainties of the ages of the
 Age QualIt\’ Q‘I: c-1, C-I?
( Ma) rank
P lower hpper
Age Quality CI, C‘I, G!
t Ma rank l-l.5 z 0.57 0.57 I). 1x 0.90
P lower P UPPer 15.0 2 0.63 0,SO 0.12 0.88
15.5 3 0.50 0.43 0.10 0.82
Rocim ts 16.0 3 0.75 0.75 0.35 0.97
5.5 4 1.oo ne 16.5 2 0.50 0.50 0.16 0.84
6.0 0 ne ne 17.0 3 0.88 0.80 0.28 0.99
6.5 1 1.oo ne 17.5 1 0.17 0.0 0.0 0.52
7.0 2 1.oo ne 18.0 1 0.0 0.0 0.0 0.52
7.5 5 0.70 0.25 0.01 0.81 18.5 3 11C ne
-_
8.0 5 0.92 0.75 0.19 0.99
8.5 5 0.93 0.67 0.09 0.99
9.0 4 1.OO 1.oo 0.48 1.oo
9.5 0 0.0 0.0 0.00 0.28 maxima and minima. In many cases where we can
10.0 5 1.oo 1.O@ 0.63 1.oo
identify deposits of relevant age and where intense
10.5 5 1.oo 1.oo 0.74 1.OO
11.0 5 1.OO 1.OO 0.75 1.oo collecting will yield good fossil samples, we can
11.5 1 0.14 * 0.14 0.02 0.43 test these assumptions. Identification of apparently
12.0 5 0.94 0.92 0.64 1.oo important intervals of low quality has changed our
12.5 3 0.83 0.76 0.50 0.95 collecting priorities in the past.
13.0 6 1.oo 1.oo 0.84 I .oo
To assess the significance of the observed turn-
13.5 6 1.oo 1.OO 0.77 1.oo
14.0 5 0.95 0.94 0. 711 1.oo over patterns, we use chi-square goodness-of-fit !o
14.5 5 0.95 0.92 0.62 I .oo test the departure of the observed pattern from an
15.0 0 0.0 (?.CI 0.00 0.25 expected distribution of first and !ast occurrences
15.5 3 0.93 0.91 0.59 1.00 (Barry et al., 1990). The null hypothesis in both
16.0 3 0.83 0.83 0.52 0.98 cases is that occurrences are distributed evenly
16.5 3 1.oo 1.oo 0.66 I .oo
9 0.90 0.80 0.28 0.99
among the 27 intervals. First occurrences in the
17.0
17.5 0 0.0 0.0 0.00 0.52 basal interval and last occurrences in the final
18.0 0 0.0 0.0 0.00 0.52 interval are excluded from the analysis. In contrast
18.5 3 IlC ne to the bootstrapping method of Maas et al. ( 1995).
our approach does not treat species ranges as of
5.5 2 ne ne
fixed duration, but rather assumes that first and
6.0 1 1X -ne last occurrences are independent of one another.
6.5 3 0.60 0:89
? 43 0.10 _ 0.82 Barry et al. ( 1990) used an even distribution as
7.0 4 0.93 0.52 J i.00 the expected, in which case the last as well as the
7.5 5 0.85 0.82 0.48 0.98 first occurrences were considered equally likely in
8.0 4 1.oo 1.oo 0.72 1.oo
I .oo 1.oo 0.74 1.oo
all intervals. use the same expected distribu-
8.5 6
9.0 5 0.94 0.91 0.59 1.oo tion here.
9.5 3 0.77 0.67 0.30 0.93
10.0 3 0.64 0.44 0.14 0.79
10.5 2 0.46 0.30 0.07 0.65 3. Results
11.0 6 1.OO 1.oo 0.69 1.oo
11.5 5 0.88 U.87 0.60 0.98
12.0 5 0.89 0.87 G.60 0.98
12.5 4 0.82 0.79 0.49 0.95
13.0 4 0.87 0.85 0.55 0.98 Data 011 interval quality are presented in ‘able 2
13.5 5 1.oo 1.oo 0.74 1.oo and Figs. 2 and 3. Both interval-quality scores
14.0 4 0.87 0.67 0.22 0.96
( Figs. 2A and 3A) and completeness indices ( Figs.
216

5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

1.0 A
6 .a
.- .6
c-i .4
.2
0 ne ne
I
111111111~111111111IIII~III1
5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

1.0
C .8
cv -6
u .4
.2
0 nc ne ne ne ne
c
I I I I I I I I II I I I I I I I I I I I I II III1
5.0 6.0 7.0 8.0 9.0 10.0 Il.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

2B.C and 3B.C ) indicate that there is considerable (Table 2) ranges from 0.16 tti 0.90, indicating
variation in completeness in the Siwalik sequence considerable variation in the quality of the esti-
for both rodents and artiodactyls. Although all mates of completeness. That is, some are good
intervals have some artiodactyls and therefore estimates of completeness, while others are very
score at least as Class 1, there are five intervals poor.
that have no rodents and score as Class 0. Both The completeness indices generally track the
rodents and artiodactyls have two Class 6 intervals quality ranks, although the quality ranks may
and each has ten to eleven Class 4 or 5 intervals. allow finer discrimination. There are also impor-
The two completeness indices ( Figs. 2 and 3 ) tant differences between the quality ranks and
clo~ly track each other, with values ranging from completeness indices for the rodents. These include
0 to 1. However, the Cl, index is lower and the intervals between 7.5 and 9.5 Ma. where the
significant discrepancies appear between the two quality ranks suggest a more comp!ete record, and
indices with respect to the rodents after 9 Ma. between 15.5 and 17.0 Ma, where the quality ranks
These discrepancies are due to the larger propor- indicate a more mediocre record. In both cases,
tion of short-ranging species counted in the CI, the C12 confidence intervals indicate that the com-
index after 9 Ma (Table 2; Flynn et al., this issue). pleteness estimates are poor. This is especially true
For the rodents the frequency distribution of both for the intervals between 7.5 and 9.5 Ma.
CI, and C& values is bimodal ( Table 2). while These estimates of record quality are meant to
that of the artiodactyls is more continuous. For evaluate how reliably the fossil. assemblages might
the C12 index the 95% confidence intervals represent the original living assemblages, and to
 J. C.

1 .o
B .8
_ .6
u .4
.2
0 ne nc

5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

1.o-
c .8 -
N *6-
c .4-
.2 -
O- ne ne ne

I I I I I I I I I I I I I I I I I I I I I I I I I I 11 I
5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

Milllons of Years

Fig. 3. Record quality l’or artiodactyls in the Fotwar Siwalik sequence. A. Interval quality ranks. 6. C’1, completcnw indtx C. C’l,
complctencss index with 95 “,0 confidence limits. II(’= no cstimatc.

determine whether data from different intervals 4 and 5, but also for Classes 3 and 2. All these
can be compared. Turning to the latter issue first, classes of intervals are possibly comparable.
Barry et al. (1990) showed that intervals with the although we regard most of the Class 3 (and all
fewest species also had the the poorest-quality of the 2) intervals as suspect, and do not use either
data, and they argued that poor quality class class in the foIlowing analysis. In addition, the
intervals should not be compared to better quality Class 6 intervals at 13.0 and 13.5 Ma have very
class intervals. At the same time, they found that high species number, suggesting they may not
there were no differences between the two best be comparable to the lower ranking intervals.
classes in the number of species and therefore However, comparison of the number of species
those classes could be directly compared to each previously reported (Jacobs et al., 1990) for these
other. This same effect is strongly evident in our intervals (which were based on much smaller
artiodactyl data, in which a plot of quality rank samples) to those given here (Table 1) shows that
against the number of species actually found (N, ) only four species were added to each interval
(Fig. 4B) shows much overlap among Classes 4, because of new discoveries. This occurs despite at
5, and 6, and wide separation from the Class 1, 2, least a 3-fold increase in the number of specimens,
and 3 intervals. We, therefore, consider the artio- and suggests that the two Class 6 intervals can be
dactyl Class 4, 5, and 6 intervals to be comparable. compared to the Class 4 and 5 intervals. Class 4,
The rodent data ( Fig. 4A) are more ambiguous. 5, and 6 intervals arc included in our analysis.
There is considerable scatter and overlap in the The second question, that of the reliability of
number of species per interval not only for Classes the data, is more difficult to address, because there
 A. Rodents
6 6 6

g 5 j 5 5 5 %A
“,4 4 4
5 3 3 3 3 3
YG
12- 2 2 2
o I- 1 1

II I I I II I I I 11 1 l l 11 11 11 l 11 ’ 1 l 11 11
0 2 4 6 8 IO 12 14 16 18 20 22 24 26 28 30

B. Artiodactyls
6 6
3 s 5

E4
6 “1 3 3333 3 3
a3 2 2 ?J 2 2
l- 111

II IIIII'Il1 II I 111 II1 IIII’I I Ii 17

0 2 4 6 8 IO 12 14 16 18 20 22 24 26 28 30

Number of Species (N,)

Fig. 4. Number of species against interval quality rank. Data are the number of species actually recovered in each interval (N,). A.
Rodents. B. Artiodactyls.

is 110 certain way to estimate original diversity of good quality for the rodents and twelve for the
from the number of species found in a fossil artiodactyls. All but one of these inter\. a!s lies
assemblage, Therefore, even in the best of circum- between 14.5 and 7.0 Ma. the most completely
stances, we have no way of knowing with certainty known part of the sequence. It is noteworthy that
how closely the Siwalik fossil assemblages record the intervals after 7 Ma are particularly poor for
the original assemblages. For example, bats were both rodents and artiodactyls.
presumably a diverse and abundant component of
the original faunas, but are rarely found even in
lass 6 intervals. Nevertheless, modern faunas
from geographic regions of comparable size have
similar numbers of rodent and large mammal Figs. 5 and 6 show the number of artiodactyl
species as the Siwalik fossil assemblages (Barry and rodent spec: present throughout the
et al., 1990). In addition, as previously noted, sequence in the better quality intervals, with Fig. 5
although the sample sizes used here are signifi- being the maximum count (N,,,) and Fig. 6 the
cantly larger than those used by Barry et al. standing richness (N,,) (see also Table 1 where
( 1990). the number of species from well-sampled values are given for all intervals). Fig. 5 also shows
intervals has changed little. We therefore conclude our best estimates for the total number of mammal
that for at least the Class 4, 5, and 6 intervals, the species in three of the best known intervals (these
number of species of rodents and artiodactyls include all species that have been documented by
found or assumed to be present approximates the our or other collections; the original assemblages
diversity of the life assemblages. Class 3 intervals undoubtedly had more bats, small carnivores, and
may also approximate original diversity for the insectivores). As can be seen from the figures, the
rodents, but because of our uncertainty we do not curves of Figs. 5 and 6 closely match each other,
include them in our analysis of diversity and with standing richness smoothing out differences
turnover. between intervals, as well as having lower values.
Thirteen of the Siwalik intervals are considered It is also apparent that there is a strong record-
 76 species
(59 genera)

63 mecles

Millions of Years

Fig. 5. Maximum number (A’,.,,,,) of rodent and artiodactyl species found or inferred to be present using th: ra!!se-through method
for 0.5 m.y. intervals. Total numbers of species and genera for all Siwalik mammals are given for 8.5. 11.Q, .-wd 13.5 Ma intervals.
Closed triangles are intervals with both rodent and artiodactyl records of Class 4 or better. Open circles are intervals with on<
record Class 3 and the other Class 4 or better.

48
44

UY 4o
E 36
% 32
: 28
; 24
f 20
ii 16
12
8
4

5mO 6.0 7.0 8.0 9.0 10.0 11.0 12.0 i3.C 14.0 IS0 160 17.0 180 19.0

Millions of Years

Fig. 6. Standing species richness (N,,) for rodents and artiodactyls in 0.5 my. intervals. Calculated from N,, =
(Nbda+Nrt)+(Nf+N,+ N,)/t. Clued triangles are intervals with both rodent and artiodactyl records of Class 4 or better. Opel1
circles are intervals with one record Class 3 and the other Class 4 or better.

quality effect (Table I), but for the region under
the solid line we think we have reliable estimates
for the diversity of the life assemblages. Therefore.
in the following discussion we focus primarily on
standing species diversity and the interval between
14.5 and 7 Ma.
There are four notable features in the overah
standing diversity pattern. First, throughout mosl
of the sequence there are more species of rodents
than artiodactyls, and, as shown in Fig. 5, the non-
rodent, non-artiodactyl component of the fauna is
usually about 30 species. This is true in spite of
the txonomic composition of this component
changing markedly over time. Second, there is an
increase in the number of artiodactyl and rodent
species between 14 and 13 Ma that seems to be
independent of the record-quality effect, although
it should be noted that the highest number of
species are in the two Class 6 intervals. Third,
there is a marked decline in the number of species
between 14 and 9 Ma, with almost all of the
decline occurring in the rodents, while the number
of artiodactyls stays nearly constant. And finally,
there may be a small, but significant, decline in
the number of species in both groups bt:tween
8.5 and 7 Ma. If we accept our previous assertion
that the intervals of quality ranks 4 through 6
are comparable, then these observed differences
between intervals should approximate changes in
diversity.
Turnover can be expressed as the number of
first and last occurrence events within each
interval, with the central question being whether
the pattern is of steady or pulsed turnover relative
to the 0.5 m.y. intervals. There are, however, two
confounding issues which need to be considered.
The first is the problem of record quality. As noted
above, intervals of lower quality bias estimates of
where first and last occurre’;lces lie by making last
occurrences seem too old and first occurrences too
young. We have therefore developed a correction
to compensate for low quality data, This correction
involves first applying the estimates of data quality
to extend the species’ranges and then partitioning
the events along the extended range. The results
are shown in Fig. 7,
In addressing the second problem, that of com-
paring records from different sequences, Gingerich
( 1987) suggested that the number of first and last
IO-
129
--------_--____
14
16 Last Occurrences
Millionsof Years
Fig. 7. Number of first and last accurrences in each 0.5 m.y. interval. Data have been transformed using equal probability
wtitioning. Scale on the right shows occurrences expressed as rate quotients, calculated from the number of events in each interval
divided by the nvcr~~ge number of events.
occurrences needs to be adjusted for the duration
of the interval and the number of contained taxa.
To accomplish this, he applied a multiple regres-
sion of (In) duration and (In) assemblage size on
(In) occurrences to calculate expected turnover,
which can be related to the observed turnover as
a rate quotient (RQ), calculated as the ratio of
observed to expected turnover (see Maas et al.,
1995). The rate quotients can then be compared
between sequences. Our intervals are of equal
duration, and thus the duration correction does
not need to be made for our data. Fig. 8 shows
the least-squared regressions of the first and last
occurrences on standing species richness. The
hypothesized dependence of the number of events
on assemblage size is clearly not present in the
Siwalik data, and therefore we have not made this
adjustment either. However, to facilitate com-
parison to the Paleogene Bighornk and Crazy
Mountains sequence we have calculated an ana-
logue of RQ, using the average number of events
per interval as the expected value. This is the same
as using a regression with zero slope. The RQavy
scale is shown on the right of Fig. 7, where RQavg
equals the number of events observed in each
interval, divided by the average per interval (4.3
for first occurrences and 4.4 for last occurrences).
40
3.5
_-------_-_ 3,o
2.5
se------__._ 1.0
- 1.5
- 25
-----_ 3.0
3.5
4.0
 221

5
t I - -,139x + 2.12, R-squared: ,004

0
28 29 3 31 32 33 33 35 36 37 3
In(x) of StandIng Species Richness

25

Y” ,055x + 1.564, R-squared: ,001

28 29 3 31 32 33 34 35 36 37 38
In(x) of Standing Species Richness

Inspection of Fig. 7 indicates that there was
considerable variation between intervals in number
of first or last occurrence events, with jntervals of
very low as well as very high turnover. When
expressed as rate quotients, the values range
between less than 0.5 to almost 3.5 times the
expected. The presence of intervals of low and
high turnover gives a strong impression that turn-
over was episodic, or pulsed, on a time scale of
500,000 years or less. That is, the temporal
sequence of first or last occurrences appears to be
clustered in short pulses separated by longer
intervals with minimal turnover, as predicted by
the turnover hypothesis of Vrba (1985). To assess
the statistical significance of these pulses, we have
tested the departure of the observed pattern from
an expected distribution of first and last occur-
rences using a chi-square good ess-of-fit test. C&r
null hypothesis is that first (or last) occurrences
are considered to be equally likely in all intervals
and therefore should be distributed evenly among
the 27 intervals. With first appearances in the basal
interval and last occurrences in the final interval
excluded from the analysis, the chi-square values
(first appearances: 58.7; last occurrences: 59.8)
have very low probabilities (d.f. = 25, p < 0.005 ).
This leads us to reject the null hypothesis and
lends support to our conclusion that the turnover
events are clustered. However, peaks of first and
last occurrences do not coincide. For example, the
peak of first occurrences at 13.5 Ma has only a
weak extinction counterpart. This lack of coinci-
dence is a finding different from that predicted by
V&a’s turnover hypothesis. in which the peaks
should fall within 0.2 m.y. of each other.
Maxima of first appearances in the Siwaliks
occurred at approximately 13.5 and 8.5 Ma, while
maxima of last occurrences occurred at 12.5 and
8.0 Ma. There also appears to be a subtle change
in the pattern over time in Fig. 7, with more pulsed
turnover between 14 and 12 Ma and more constant
turnover after 11 Ma. This is expressed most
strongly in the record of first appearances, but
also appears in the last occurrences.
4. Discussion
Correlations between the records of Siwalik
biotic events and regiona! tectonic events are
imprecise, largely because the timing and magni-
tude of critical tectonic events are uncertain. In
addition, the effects of tectonic events are likely to
have been complex and transmitted only indirectly
through the intervening fluvial system. There is,
however, evidence from the Indus and Bengal
submarine fhns for a strong pulse of uplift that
began around 11 Ma, had peak values at 9 Ma,
and lasted until 7.5 Ma OI even later ( Amano and
Taira, 1992; Rcn, 1992). On the Potwar Plateau,
major changes in lithology and greatly increased
sedimentation rates have also been documented at
circa 11 Ma and related to tectonic events (Johnson
et al., 1985; Willis, 1991), while Cerveny et al.
( 1990) interpreted a change in heavy mineral
assemblages at circa 11 Ma as due to a change in
sediment-source area. There is also evidence that
the Indus and perhaps more regional drainages
were substantially reorganized between 7.5 and 6.5
Ma (Amano and Taira, 1992; Behrensmeyer,
1987). The latter event may have contributed to
the poor preservation of fossils after 7.5 Ma.
Correlations between Siwalik biotic events and
global or regional climate change are also uncer-
tain, primarily because of imprecise correlations
between the marine and terrestrial records. There
is a general correlation between the Middle
Miocene diversification of Siwalik faunas prior to
13 Ma and a long period of global cooling and
climatic instability ( Woodruff et al., 1981; Kennett,
1986; Vincent et al., 1991). However, age estimates
for the onset and end of the isotopic event differ
among authors by 0.5 m.y. (e.g. Kennett, 1986;
Vincent et al., 1991), making correlations with the
terrestrial Siwalik record uncertain.
Correlations with Late Miocene climatic events
are better, especially the correlation with the
oxygen and carbon isotopic changes noted by
Quade et al. (1989) at circa 7.5 Ma. These excLu--
sions were initially thought to docutnent the origin
of a local south Asian monsoonal systetn with
greater seasonality of precipitation, and their age
accords well with a change in monsoon driven
upwelling patterns in the Arabian Sea ( Kroon
et al., 1991). More recently Ccrling and others
(Cerling et al., 1993; Quade and Cerling, 1995)
have argued the isotopic changes also reflect cli-
mate change on a more global scale or change in
atmospheric composition, or both. Regardless of
the underlying cause, the decreased first and
increased last occurrences and subsequent small
diversity decrease noted after 8 Ma are contempo-
raneous with the isotopic transition, and may
result frotn the vegetational changes reflected in
the isotopic shift. Both the fauna1 and isotopic
changes in the Siwaliks correlate to global cooling
trends and increasing seasonality and aridity gcn-
erally noted in the Late Miocene.
The Siwahk isotot%c I record. however, does not
explain the much greater decline in species number
and attending fauna1 turnover apparent before 10
Ma, and it is difficult to find satisfactory correla-
tions to more global events. A sea-level lowstand
at about 10.5 Ma (Haq et al., 1987) corresponds
to the first of two major marine isotopic excursions
between 11 and 8.5 Ma (Kennett, 1986, using the
nannoplankton correlations of Berggren et al.,
1985). These isotopic events were probably the
result of large-scale climatic events that could be
related to the appearance of equids in southern
Asia and in Africa at circa 10 Ma. However, the
isotopic excursions are too late to explain either
the Middle Miocene diversity decrease or the larger
peaks of first and last occurrences between 14 and
12 Ma. This suggest there were other, as yet
undetected, 1
changes in the
Comparisons of the patterns of Neogene Siwalik
nge to those noted by Maas e
leogene Bighorn and Crazy
Basins lead to the following generalizations.
The two sequences have similar geographic
scope, and span approximately equal lengths of
time with equivalent levels of temporal resolution.
Both sequences have similar levels of completeness,
with a few poorly known intervals within otherwise
more complete sections. In both cases these poorly
known intervals limit our understanding of fauna1
change, due to uncertainties in species’ranges.
Although expressed in different ways, generic
and presumably species richness in the two
sequences was broadly similar. In the Siwaliks,
generic richness (expressed as a generic analogue
of N,,, ) varied between 50 and 59 genera (lkble 1,
Fig. 5 ), while in well-sampled Paleogene zones
standing generic richness (the analogue of /&,)
varied between 31 and 62.5 genera (Maas et al.,
1995; Table 2 and Fig. 5). Since standing richness
is generally lower than maximum richness, this
suggests that the standing diversity of the richest
Siwalik faunas may have been slightly less than
the richest Paleogene faunas and that the number
of Siwalik species changed less between intervals.
Aiternativeiy, differences in the quality of the
Neogene and Paleogene records and intensity of
sampling in the two areas could also account for
the differences.
Unlike the patterns of species diversity, the
number of higher-level taxonomic groups was very
different. Three out of thirteen orders dominate
the Siwalik faunas in both number of species and
number of fossils, a pattern which is characteristic
of African and Eurasian Neogene faunas. In con-
trast, the Paleogene assemblages have a more
equable representation of genera and species within
a larger number of ordinal-level taxa.
Species and generic diversity changes of compa-
rable magnitude occur in both sequences, with
both demonstrating trends that persisted for as
long as 5 or 6 m.y. Some fauna1 turnover is always
the Neogene a aleogene
ewes, and whe;.l expressed as rate quotients.
Iess than 0.5 to almost 3.5 times the
expected amount. T ese rates are very modest
compared to the rates of major sxtinctions in the
marine record (Cingerich, 1983).
There is evidence in both records suggesting that
fauna1 change occurred episodically, with turnover
concentrated in brief pulses separated by longer
intervals of low turnover. The Siwalik sequence
may have bepn the more strongly pulsed, with
longer intervals of low turnover separating the
peaks. axima of first appearances in the Siwaliks
occurr at approximately 13.5 and 8.5 Ma, while
maxima of last occurrences where at 12.5 and 8.0
Ma. In neither record do peaks of first appearances
coincide with peaks of last occurrences,
Changes in diversity and pulses of turnover can
cases be related to climatic or tectonic
0th the Siwalik and Palcogene sequences,
however, have other significant fauna1 changes
whose origin does not seem related to climatic or
tectonic events.
Patterns of fauna1 turnover and changes in
diversity in terrestrial Neogene vertebrate faunas
in north+_hipt
Pakistan can be analyzed for 0.5 m.y.
long intervals. On the Potwar Plateau the sequence
estends from circa 18.5 Ma to 1 Ma, with the
period between 18.5 and 5.5 Ma being best sampled
and having the highest data quality. In this time
range, well-sampled intervals have 50 or more
species of mammals, belonging to thirteen orders,
three of which are most common.
Between 18.5 and 5.5 Ma, species diversity and
the rate of turnover varied considerably. Strati-
graphic range data for rodents and artiodactyls
show that the number of species increased between
15 and 13 Ma, and then decreased after 13 Ma.
The data also show fauna1 change in the Siwaliks
was episodic, occurring as short intervals with high
turnover, followed by longer periods with much
less change. Peaks of first appearances occur at
approximately 13.5 and 8.5 Ma, while peaks of
last occurrences occur at 12.5 and 8.0 Ma. It is
thus apparent that in the Siwaliks increased extinc-
tion did not coincide with increased first
appearances.
Some of these fauna1 events can be correlated
to global climatic trends. The best case is the latest
Miocene decline in diversity and increased turn-
over that accompanied oxygen and carbon isotopic
changes at 7.5 Ma and correlates to globally
increasing seasonality and aridity. A weaker exam-
ple is the Middle Miocene diversification of Siwalik
faunas during a period of global cooling.
Comparisons to the Bighorn and Crazy
Mountains sequence establish a number of signifi-
cant similarities and differences. Both sequences
span long periods of time with similar levels of
temporal resolution, and both have similar lcvcls
of completeness. Diversity in the two sequences
appears to be broadly similar, with well-sampled
intervals having 30 to over 60 genera. Changes in
species and generic diversity were also of compara-
ble magnitude, with monotonic trends persisting
for more than 5 million years. The relative magni-
tude of fauna1 turnover was also similar in both
sequences, ranging from less than 0.5 to almost
3.5 times that expected, and there is always some
fauna1 turnover present. In both sequences fauna1
change appears to be episodic, with short pulses
of high turnover between longer intervals of low
turnover, In neither record do peaks of first occur-
renccs coincide with peaks of last occurrences.
Ic d~~~~e~~esinclude the pattern of genus
and species representation within higher-level taxo-
nomic categories. In addition the Siwalik faunas
may have been slightly less diverse than the
Paleogene faunas and the number of species may
have changed less over time. Siwalik fauna1 change
may also have been more episodic, with more
distinct pulses and longer intervals between pulses.
Acknowledgments
We wish to thank Catherine Badgley and Anna
Kay Behrensmeyer for the chance to participate
in the NAPC-V symposium and an earlier work-
shop, and Mary Maas for providing an early
version of her accompanying paper, The comments
of David Webb and an snonymous reviewer were
very helpful. Our close collaboratiotl with members
of the Geological Survey of Pakistan has made
this research possible. Will Downs generously
made data and his time available. Financial sup-
port derives from NSF grants BSR U-00145 and
BNS-88 12306 and SFCP grant 7087 120000.
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