continental-scale hypsodonty patterns,climatic … · [email protected]); alexey tesakov,...

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Mikael Fortelius1, Jussi Eronen1, Liping Liu1,2, Diana Pushkina1,Alexey Tesakov3,Inesa Vislobokova4 & Zhaoqun Zhang2

1 Department of Geology, University of Helsinki2 Institute of Vertebrate Paleontology and Paleoanthropology,Academia Sinica, Beijing3 Geological Institute, Russian Academy of Sciences, Moscow4 Paleontological Institute, Russian Academy of Sciences, Moscow

Fortelius M., Eronen, J., Liu, L.P., Pushkina, D., Tesakov, A., Vislobokova, I. & Zhang, Z.Q., 2003- Continental-scale hypsodonty patterns, climatic paleobiogeography and dispersal of EurasianNeogene large mammal herbivores - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND

MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 1-11 [ISSN 0923-9308] Published 1 December 2003

The dispersal of land mammals depends not only on the availability of physical connections butalso on the presence of habitats that can support viable populations. Here we use mean molar hyp-sodonty of large mammal plant-eaters to map environmental conditions in Eurasia during theNeogene in order to establish a framework for discussing the distribution and dispersal of Neogeneland mammals. The first Early Miocene centres of hypsodont faunas are seen in Iberia and centr-al Asia. In the Middle Miocene Iberian values are low and a strong East-West contrast is seen with-in Europe, with another centre of hypsodonty developing in eastern Asia. From the Late Mioceneonwards we show a pattern of high values in the central part and low values in the humid areas ofwestern Europe and southern China (no suitable data are now available for southernmost Asia).This pattern has remained relatively stable since the Late Miocene, with only regional changes anda general increase in the overall level of hypsodonty. These results suggest that the hypsodonty pat-tern is primarily controlled by climatic effects of Himalayan-Tibetan uplift, specifically to thedrying and increased seasonality of humidity predicted by climate models, rather than to the cool-ing that would have been most noticeable in the northern half of the continent. The strong and per-sistent relationship between position on the continent and relative degree of hypsodonty suggeststhat adaptation to local conditions by natural selection has been the main determinant of ungulatehypsodonty in the Neogene. A logical consequence of this would be that regional climatic condi-tions have been a major determinant of the geographic ranges of individual species throughout theNeogene, except perhaps at times of major faunal turnover. For example, an initial dispersal of hip-parionine horses across the arid Central Asia appears highly improbable compared with dispersalalong a more humid northern route, and the apparently early arrival of hipparions in Spain maywell reflect this circumstance.

Correspondence: Mikael Fortelius (to whom correspondence should be addressed), Jussi Eronenand Diana Pushkina, Department of Geology, P O Box 64, FIN-00014 University of Helsinki,Finland (e-mail: [email protected]; [email protected]; [email protected]); Liu Liping and Zhang Zhaoqun, Institute of Vertebrate Paleontology and Paleoanthropology,Academia Sinica, P.O. Box 643, Beijing 100044, China (e-mail: [email protected]; [email protected]); Alexey Tesakov, Geological Institute, Russian Academy of Sciences,Pyzhevsky 7, 109017 Moscow, Russia (e-mail: [email protected]); Inesa Vislobokova,Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya, 123, 117868 Moscow,Russia (e-mail: [email protected])

Keywords: Neogene, paleogeography, paleoclimate, paleodiet, fossil mammal dispersal

Continental-scale hypsodonty patterns, climaticpaleobiogeography and dispersal of EurasianNeogene large mammal herbivores

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DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

INTRODUCTIONIt has been customary to discuss the dispersalof fossil land vertebrates in terms of physicalbarriers and connections. While there can beno doubt that these factors are ultimatelydecisive, it must also be acknowledged thatland mammals appear to be very good atcrossing seemingly insurmountable barriers(Solounias et al. 1999), and that the low tem-poral resolution of the mammal record usual-ly makes it difficult to propose a direct rela-tionship between dispersal events and speci-fic geophysical changes with confidence.Moreover, the successful crossing of a bridgeor a barrier remains invisible to the fossilrecord unless it results in successful colonisa-tion of the new area, and this depends not onthe details of topography or land-sea configu-ration but (apart from luck) on the nature ofthe invaded habitat. Indeed, the distributionpatterns themselves - quite apart from disper-sal dynamics - must ultimately depend to amajor degree on environmental conditions.

The first serious attempts to understand thecontinental-scale evolution of Eurasian envi-ronments and biota seem to have been madeby Soviet workers of the 1940’s to 60’s (e.g.Borissiak & Beljaeva 1948; Orlov 1962;Dorofeyev 1966), probably the only profes-sionals at the time with access to fossil mate-rial of sufficient geographical range to con-template such a task. Although hampered bymajor stratigraphical problems these synthe-ses did in fact establish a coherent view,based on both the nature of the fossil assem-blages and multidisciplinary study on strati-graphy, tectonics, paleoclimatology and paleo-geography. It was thus clearly recognised thatthe general trend towards increasingly openenvironments observed over the entire areadid not proceed at a uniform pace over theentire continent, but took place much earlierand proceeded further in the continental inte-rior than in the climatically more moderatedmargins, especially Europe with its complica-ted system of large seaways. The influenceof environmental and climatic changes on thehistory of some dominant groups of mammals

were analysed in the series of the fundamen-tal works of next generations of Russianscientists (e.g., Zhegallo 1978; Vislobokova1990). But it was only with the seminal workof Bernor (1983; 1984) that the concept ofdiachronous biotic and environmental changerather belatedly became incorporated in wes-tern work on Eurasian Cenozoic mammalevolution.

It may seem surprising that fossil mammalsshould have been so central to developing ourideas about environmental change during theCenozoic, but there are in fact good reasonsfor this. Fundamentally, plants enjoy a moredirect relationship with climate than do ani-mals, of course, but neither macroscopicplant remains nor fossil pollen floras have thestratigraphic and geographic coverage offeredby fossil mammals, nor do they permit parti-cularly detailed or reliable reconstructions ofregional vegetation types. Fossil mammalsare therefore simply one of the best sourcesavailable for reconstructing past vegetationpatterns and thus of climate. Not only havefossil mammals been a main focus ofresearch for a long time and are thereforerelatively well understood, but the environ-mental signal reflected in mammalian mor-phology and community structure is also oneof the strongest and best resolved. Damuth(1982) showed that the original communitystructure is robustly preserved in mammalianfossil assemblages, and Janis (1984) arguedexplicitly for the use of fossil mammal com-munities to reconstruct past vegetation pat-terns. Fortelius et al. (1996) used fossil mam-mals to chart large-scale patterns of environ-mental change on a gradient from closed toopen vegetation in the Miocene of westernEurasia. More recently, we (Fortelius 2003;Fortelius & Hokkanen 2001) showed persis-tence during 20 Ma of a difference in meanmolar capability of ungulates from differentregions of Eurasia, over an interval duringwhich the same measure increased aboutthreefold in all areas compared. Finally, workin progress by a group coordinated by JohnDamuth (http://www.nceas.ucsb. edu/

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FORTELIUS et al.: Eurasian Neogene hypsodonty

fmt/doc?/nceas-web/results/projects/98DAMUT1) has shown that communitystructure and ecomorphology of living mam-mals can be used to successfully predictmodern vegetation patterns worldwide.

The relationship between herbivore hypso-donty and a wear-inducing diet is now almostuniversally recognised (Janis & Fortelius1988; Romer 1970; Van Valen 1960;MacFadden 2000). It is well documentedempirically and easily understandable theore-tically in terms of functional demands andadaptive evolution. The factors potentiallyinvolved are many, but virtually all increasein effect with increasing aridity and opennessof the landscape (increased fibrousness,increased abrasiveness due to intracellularsilica or extraneous dust, and decreased nutri-tive value) (Fortelius 1985; Janis 1988).Hypsodonty should thus record a condition ofthe vegetation that might be termed generali-sed water stress, either in overall conditions,or (perhaps more commonly) as a regularlyoccurring extreme period, such as a dry seas-on. We therefore propose here to use thesimple property of mean hypsodonty of mam-malian large herbivores to map the develop-ment of this ‘generalised aridity’ duringNeogene time.

MATERIALS AND METHODSThe data used were downloaded from theNOW database on March 8, 2001 and inclu-ded non-public datasets for the Former SovietUnion (FSU) and China. The Chinese datasetwas further supplemented from an ongoingcompilation project. A subset limited toEurasia and large land mammals was selec-ted. All analyses reported here were limitedto species characterized as plant-eaters orplant-dominated omnivores. Hypsodonty wasranked according to the following scheme:brachydont=1, mesodont=2, hypsodont orhypselodont=3 and means were calculated forindividual localities or other subsets(SPLOC-analysis; see Fortelius & Hokkanen2001). The possible range of mean hypsodon-

ty is thereby constrained to the range 1-3,beyond which this metric cannot resolve(paleo)precipitation. Values based on singlespecies were excluded from all analyses.

The data were grouped into 1 Ma time sli-ces on the mean value of minimum and maxi-mum age estimates as reported in the databa-se. Except for some Chinese localities, locali-ties with a difference between minimum andmaximum estimates of more than 3 Ma wereexcluded. The correlation table of the NOWdatabase follows Steininger et al. (1996),with ad hoc additions by AL and ZZ for theFSU and Chinese data. For investigation ofgeographic gradients, the data were groupedinto sectors of 10 degrees longitude andzones of 5 degrees latitude.

For the GIS maps, the time slices werecombined into four larger temporal groupings(see Figs. 1 a-d for details). All GIS mapswere made in MapInfo Professional 6.0 usingthe inverse distance weighted (IDW) algo-rithm and the following settings: cell size 30km, search radius 3000 km, grid border 1100km, number of inflections 9, values roundedto 1 decimal. The inflection values weremanually set to the range 0.7-3 for all maps,and a mask was manually superimposed tofade out areas more than 1000 km from thenearest data point (opacity 50%). We delibe-rately use modern maps as a background forthese patterns partly because of the lack ofpalinspastic paleogeographic maps (on whichthe localities could be automatically plottedin their correct positions) and partly becauseour time slices span more time than any pale-ogeographic configuration. We also usemodern geographic names (such as ‘China’ or‘Iberia’) as neutral landmarks. It is devoutlyto be wished that palinspastic paleogeograp-hic maps of the Eurasian continent will beco-me available, since discussing dispersal inrelation to a modern basemap is difficult atbest and can easily be misleading.

The dataset used is available from the aut-hors, and the latest public NOW dataset canbe downloaded from http://www.helsinki.fi/

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science/now/, where further details regardingthe data may also be obtained.

RESULTS

Hypsodonty mapsThe Early Miocene hypsodonty pattern (Fig.1d) suggests two centres of early increase incrown height: Central Asia and the IberianPeninsula. In both cases the increase is seenonly in localities from the end of the interval,later than c. 18 Ma. Data for eastern Europeand Asia are scarce and there is little detail,but the western and central European covera-ge is satisfactory and the lack of hypsodontspecies outside Iberia is well established. Thehigh-crowned species involved in this earlyphase are primarily mesodont and hypsodontrhinoceroses of the ‘Hispanotherium fauna’,known in the Early Miocene only fromCentral Asia and Spain and later also fromsouth-western Asia (Heissig 1976; Köhler1987). The age of the Anatolian localitiesInönü I, Pasalar and Çandir is treated here asmiddle Miocene, following the NOW databa-

se, Bernor & Tobien (1990), and Steininger etal. (1996). It is, however, also possible tointerpret these localities as early Miocene(Sen 1990), and this is in fact done in thelatest rodent zonation for Anatolia (Ünay etal. this volume). If an Early Miocene age isaccepted for any of these localities the EarlyMiocene development of hypsodont ungulatecommunities would be circum-Mediterraneanand the difference between the Early andMiddle Miocene patterns smaller than it nowappears.

The Middle Miocene pattern (Fig. 1c) isstrongly dominated by the contrast betweenwestern and eastern Europe, the ‘West’ and‘East’ previously recognised from taxonomicand biogeographic as well as ecomorphologi-cal comparisons of mammal data (Fortelius etal. 1996; Werdelin & Fortelius 1997;Fortelius & Hokkanen 2001). The data pointsfor Asia are few but there is an indication ofhypsodont faunas beginning to evolve innorthern China. These assemblages, typifiedby the famous Tunggur fauna (Qiu 1989), arecharacterised by mesodont and hypsodont

Figure 1 Colour-interpolation map of mean hypsodonty in Eurasia during five intervals. a Pliocene (5-2 Ma); b late Miocene (11-5 Ma); c middle Miocene (15-11 Ma); d Early Miocene (24-15 Ma). See Methods for technical details.

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bovids and rhinoceroses occurring in a con-text still dominated by brachydont ruminantsand suoids. The few data points in CentralAsia show little sign of increasing hypsodon-ty at this time.

The Late Miocene pattern (Fig. 1b) showsthe emergence of more hypsodont faunasdominated by bovids and horses in theMediterranean region and central Asia, espe-cially the areas north of the Tibetan Plateau.This pattern is quite unlike the modern distri-bution of mean annual temperature (Fig. 2a)and strikingly similar to the modern meanannual rainfall distribution (Fig. 2b), espe-cially in the areas of persistently low hypsod-onty in the humid western Europe and sout-hern China. A temporal breakdown of theLate Miocene localities shows that the laterpart of the interval appears relatively drier inthe western part of the continent, while thereverse is true for the eastern part. (In thispreliminary survey we have chosen not todelve too deeply into the effects of time-aver-aging, which obviously influence all ourmaps to a greater or lesser extent.)

The Pliocene pattern (Fig. 1a) generallyresembles the latest Miocene one but is evenmore similar to the modern rainfall pattern.Overall hypsodonty is higher, the main centreof hypsodonty has shifted from the easternMediterranean to Central Asia, and China isnow included in the realm of high hypsodon-ty. The low hypsodonty values seen in theCaucasus may reflect local climatic condi-tions the explaining presence there in thePliocene and Early Pleistocene of Africanelements, including early humans (Gabunia etal. 2000).

Geographic hypsodonty gradientsThe history of increasing hypsodonty illustra-ted by the maps shows considerable regulari-ty with respect to geography. A coarser viewwith hypsodonty values reduced to latitudinaland longitudinal group means shows that thisregularity developed gradually over time(Fig. 3 a,b). The appearance of the modernlatitudinal gradient with higher values in the

south took place in the Middle Miocene andremained in place while the overall levelalmost doubled during the Neogene (Fig. 3a).The relationship between hypsodonty andlongitude is less regular but shows the suc-cessive development of a pattern with highervalues in the interior of the continent andlower values near the coasts (Fig. 3b). Theshift of the highest values from the westtowards the east in the course of the Neogeneis also seen.

DISCUSSIONIt is helpful to discuss the present results inrelation to present-day climatic patterns,especially the basic variables temperature andprecipitation (Fig. 2). It is immediatelyobvious from a comparison of the hypsodontymaps with these that the pattern seen fromthe Late Miocene onwards is quite similar tothe distribution of rainfall and quite differentfrom the regular zonation of temperature.This suggests that the basic assumption ofour approach is valid, and that hypsodontycan indeed be used as a proxy for variablesthat correlate with rainfall, such as the ‘gene-ralised aridity’ proposed in the Introduction. The development of the modern climatic pat-tern of the Eurasian continent is clearly aLate Miocene phenomenon, and the earlierNeogene development of less humid regionsapparently took place under a different clima-tic regime. One does not need to search farfor a plausible mechanism for dividing theclimatic history of Eurasia near theMiddle/Late Miocene boundary. Indeed, theliterature on the later Neogene climate historyof Eurasia has recently focused on the issueof the effects of uplift of the Tibetan Plateauon winds and precipitation patterns (apartfrom high-latitude cooling). The distinctchange from the Middle to the Late Mioceneseen here can be taken to coincide with amajor phase of uplift in the HigherHimalayas (Amano & Taira 1992) and theTibetan Plateau at about 10 Ma (An et al.2001) and with evidence of major contempo-raneous environmental change (Tanaka

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1997). The ‘Mid-Vallesian Crisis’ (Agustí &Moya-Sola 1990) of western Europe is per-haps the best-documented (Fortelius et al.1996) example of a major Eurasian mammalevent at about 10 Ma, and recent reviews ofthe evidence suggests a continent-wide eventthis time (Agustí et al. 1999; Fortelius &Hokkanen 2001).

Specific predictions from general circula-tion models concerning the climatic effects oflate Miocene Tibetan uplift include decre-asing mean annual precipitation to the northand west of the plateau and increasing preci-pitation to the south and east (Kutzbach et al.1993). The hypsodonty pattern shows justsuch an effect, although with considerably

more detail. It is particularly striking that therise of hypsodonty values is much lower inChina than elsewhere on the continent, andthat the main single difference between theMiddle and the Late Miocene pattern is in thearidification of Asia to the north of theTibetan Plateau. The development of persis-tent geographic gradients (Fortelius 2003;Fortelius & Hokkanen 2001) is reflected herein the relatively stable latitudinal and longitu-dinal hypsodonty profiles, where the increasein overall hypsodonty proceeds without majorchanges of the relative proportions (Fig. 3).This seems to suggest that these patterns fun-damentally reflect overall climatic relations-hips of a continent, such as the difference

Figure 2 Maps of present-day mean annual temperature (a) and mean annual precipitation (b) in Eurasia. Created from publicdata provided by the Food and Agriculture Organization of the United Nations, Environment and Natural Resources Service(Leemans & Cramer 1991) using the public WinDisp application. For temperature, reds indicate the highest values, blues and whites the lowest. For precipitation, greens indicate the highest values, browns and greys the lowest. Data and software weredownloaded from http://www.fao.org/WAICENT/FAOINFO/SUSTDEV/eidirect/CLIMATE/EIsp0002.htm.

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between the oceanically moderated rim andthe ‘continental’ interior, and that this deeppattern is modified but not fundamentallyaltered by tectonic changes.

As regards the geographic distribution ofspecies, the close relationship with expectedand observed climate patterns strongly sugge-sts that natural selection tracking environ-mental change is the main mechanismresponsible for the persistence of thegeographic contrasts. This is further suppor-ted by the observation that the geographiccontrasts may break down temporarily attimes of major taxonomic turnover, but aresoon re-established (Fortelius 2003). In theabsence of evidence to the contrary, we pro-pose that the case of hypsodonty is typicaland that the long-term ranges of mammalianspecies are typically under strong and directenvironmental control. This is not to say thatall species are equally restricted in thisregard. It is quite clear, for example, that her-bivores enjoy (or perhaps more appropriatelysuffer from) a more direct relationship withvegetation and climate than do carnivores, acircumstance reflected in the greater geograp-hic ranges and stratigraphic ranges of carni-vores compared with herbivores (unpublishedanalysis of NOW data 1999). Nevertheless,there seems to be a clear relationship betweenoverall taxonomic provinciality and the

regional hypsodonty patterns shown here, inthat the breakdown of the environmentalEast-West contrast in Europe from the LateMiocene onwards is accompanied by a corre-sponding reduction in provinciality at thistime (Fortelius et al. 1996; Werdelin &Fortelius 1997). Times of great environmentaldiversification are thus indeed likely to beaccompanied by reduced dispersal of species,and vice versa. Koufos (this volume) provi-des multiple examples of open-adapted spe-cies of the eastern Mediterranean realm thatfailed to disperse into the closed environ-ments of central Europe during the earlierpart of the Late Miocene, while the East-Westcontrast was still strong.

Late Miocene hipparionine horsesWe would like to end our discussion by tre-ating briefly what is probably the most spec-tacular and well-known dispersal event of theEurasian Neogene: the arrival and adaptiveradiation of hipparionine horses at the begin-ning of the Late Miocene. The NorthAmerican ancestors of the earliest Eurasianhipparions are best known from quarries inTexas and Nebraska, but the actual popula-tion from which the first Eurasian individualswould have been derived must have existedin Alaska, which would have been a relative-ly cool and humid place compared with the

Figure 3 Development of Eurasian latitudinal and longitudinal hypsodonty gradients over Neogene time. a Latitudinal gradient,b longitudinal gradient. Bars show standard error.

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North American interior. (Alaska is in theregion where plateau uplift would inducecooling and well outside the area wheremodel simulations predict drying; Ruddiman& Kutzbach 1991) On arrival, having per-haps crossed the Bering Strait during oneconvenient eustatic sealevel low, these firstEurasian hipparions would have found them-selves in a place that was climatically similarto their point of departure. Although the evi-dence is most unfortunately lacking, suchconditions must have extended westwardsalong the Eurasian coast all the way to wes-tern Europe, whereas vast arid regions wouldhave been encountered to the south.Preliminary results indicate that conditionswere quite dry as far south as central Chinain the earlier part of the Late Miocene (Zhanget al., 2002).

A trajectory across central Asia, such as istraditionally shown on maps of hipparioninedispersal, therefore appears to us quite unli-kely compared with the obvious choice ofexpanding westwards in a familiar environ-ment. We wish to point out that although thedata are woefully inadequate, such expansionwould fit the raw pattern of hipparion appe-arances quite well, with the earliest recordfrom the best sampled region of westernEurope for that time interval (The Vallès-Penedès basin of Spain; Garcés et al. 1997).The initial spread would have been geologi-cally instantaneous, whereas the subsequentcolonisation of more arid regions would haveproceeded at the pace of the subsequent adap-tive radiation, fast but potentially detectablein long sequences with good stratigraphiccontrol (Bernor et al. 2000). The processwould have resulted in the secondary evolu-tion in more arid regions of the Old World ofanimals eco(morpho)logically more similar toNorth American species known from the con-tinental interior than to their own immediate(presumably Alaskan) New World ancestors.One could speculate that such a processmight cause an evolutionary reversal of somemorphological characteristics, explainingsome of the perplexing systematic patterns

observed. Without going into further detailwe would finally like to point out that similardispersal scenarios might potentially apply tocamels, and perhaps even to canids.

It is obvious that large herbivore hypsodon-ty is only one – and a crude one at that – ofmultiple mammal-based environmentally sen-sitive indicators available. In the present con-text we would like to end with the wish for astudy of continental-scale patterns usingsmall mammal ecomorphology.

ACKNOWLEDGEMENTSWe thank Hans de Bruijn for providing theoccasion for an excellent meeting and theorganisers for inviting us to contribute.Special thanks to Albert van der Meulen formuch patience and encouragement. We thankLinda Ayliffe, Alan Gentry, HannuHyvärinen, Jukka Jernvall, Anu Kaakinen,Juhani Rinne, Jan van Dam, and Zhou Lipingfor valuable discussions and comments. Thework was supported by Academy of Finlandgrant nr 44026 (to Mikael Fortelius).

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Received 18 June 2001

Accepted 16 September 2002

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