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Faunal and Floral Migration and Evolution in SE Asia-Australasia

Iain Davidson
Archaeolo gy and Palaeoanthropolog y
Hu m a n and Environmental Studies
Univ ersit y of New Engla n d
When Alfred Wallace emb a rked on his voyage from Bali to Lombok he
turned his back on the limitations of our homi nin ancestry and did
something distinctively human: he drew a line (Davidso n 1999b) (Figu r e
1). Morwood’s confirmat ion of dates for ston e artefa c ts earlier than 800
thousand years ago (Morwood et al. 1999; Morwood et al . 1998) has
shown how our homin in ancest o r s may have crossed simila rly along the
ch ain of islan ds of the Les ser Sundas. That may or may not have been a
step of simila rly huma n dimensi o ns. In this paper, I arg ue th at it is
implausibl e to interpr et that small step as an indic ator of the emerg ence
of human abilities and behaviour. The really “gian t leap for man kind”—
for our hu ma n ances t o r swas the journey to Australia (Davidso n &
Noble 1992).
The journey of our ancesto r s from the tim e of our common ancesto r with
other ap es can be represented as a mor e prosai c one from Canberra
(wher e we left the apes behin d) to Armid ale. In Sydney those ancesto rs
became bipedal, they first left stone tools in Newc a stle, made handaxe s
in Mus wellbrook and continued to make them almo st until they re ached
Armidal e. For those unfamiliar with the geog r a p h y of eastern Australia,
a simila r line can be draw n in the United St ates for a journey from San
Fr ancisc o to New York (Figur e 2). By this compa rison, bip edalism
eme rg ed in San Fr ancisc o, ston e tools in Salt Lake City. Acheule a n
handaxes first appeared aroun d Kansas City and continu ed to be mad e
until beyo nd Philade lphia. On this time scale, ho mini ns crossed into
Flor es around Indianapo lis.
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So mewhere close to Armidale, or bet ween Philadelp hia and New York,
our ancestors began that most distin ctive of human behavi ours: they
began to talk to each other in languages only they und erstand.
Langua g e, as a mean s of com m u n i cation usin g symbols, involve s,
pe rmits and perhaps requir es the generati on of me anings shared among
a community brought up to share the conventio n s of those symbolic
mea nings (Noble & Davidson 1996). This is true of those brough t up
speaki n g Australi an rather than American English, or those who learned
geology rathe r than archae ology, or are universi ty- educ a t e d rathe r than
not. Such distinctions draw lines between communiti e s as strong as or
stronger than the line that Wallac e cross e d.
Once lang u a g e emerged, all hell broke loose—all of it withi n Armidale, or
between Philade lphia and New York. Language- using peopl e coloni sed
Australia; they painte d cav es, invente d agric ulture. They star ted to write
down details of the wealth they had accumula te d (on clay table ts in
cu neiform of Linear B) and then they made up sto ries about it. They
invented gods, colo nialism an d the Y2K bug.
My arg u m e n t in this paper is that the early ston e tools from Flores, while
re m arkable and impor t ant, indicat e a sea- crossing by some mea n s, but
they do not cons titute evidence, as the later sea- cros sing into Australia
did, of the beginnin g of that proc ess of em ergence of the full rang e of
hum a n abilitie s.
Our bipedal early ancesto rs were quite distinc tive from the othe r apes of
Africa. Ther e ar e various ways of defining these distinctive differenc es
(eg Isaac 1978). Noble and I (Noble & Davidson 1996) have point ed out
severa l features: biped alism (Hunt 1994); ston e tool carrying, makin g
and use (Sc hick & Toth 1993; Wynn & McGr ew 1989); meat cuttin g
(Keeley & Toth 198 1); wood working (pointed sticks ) (Keeley & Toth
1981; Thieme 1997); aimed throwing (Calvin 1982; 1993); meat eating
(Aiello & Wheeler 1995); and the coloni satio n of temp e r a t e and lightly-
woode d envir onme n t s (Gam ble 1986).
The question, which is cruci al to evaluating the early evidence fro m
Flor es, is when these an cestors became more like humans and less like
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ap es. My answ e r to this que stion is that only humans can ask the
qu estio n and only we can answe r it, bec ause only hum a ns have the
ability to choos e where to draw the line and thus to draw it in differ e nt
places. Only huma ns can define different conve n tions for the arbitrary
relation between meaning and the symbols with whic h we discuss it
(Noble & Davidson 1996). Only hum a n s can give meanin g to Wallaces
line, Huxleys line and all the oth er lines that slice through our
understanding of the biogeographic divers ity of Wallacea. That ability to
dr aw the line, which Wallac e exem p lified so well, is the distin ctiv e ability
that permitt e d people to take that giant lea p into Australia. Is it likely
that the evidence from Flores also indicates such a leap? Does that
voyage de monstrate the emergence, 800 000 years ago, of that
combin a tion of arbitrarin es s and convention which langu ag e alon e
pe rmits?
One way to demonst ra te the emergence of humanness would be to arg ue
that the evidence of homi nin evolution leaves unambiguous an swers.
Physical anth r opologists interpret the phylogeneti c relationshi p s
re presented by the various fossilise d frag m e n t s of the skeleto n s of tho se
ancestors on the basis of careful comp a rison of morphol ogical variations
of different parts of the skull (Groves 1989). The re sult ing classific atio n s
into specie s, of cou rse, canno t be tested agai nst any other real recor d of
specie s identification. They are artefacts of the meth o ds used by
physical anthr o p ologi sts, and are likely to be oversi m plified and speci es
num b e r s underestimated because of the tiny number of fossils that have
be en found (Foley 1991; cf. Tatt ersall 1992). Indee d, it has recent ly been
de monstrated that ther e is a mismatch between skeletal variation and
specie s identification based on genetic compari s o n s (Collar d an d Wood
2000), although similar methods of gener a t ing phylogenetic tree s ar e
able to match gen etic trees when applied to soft tiss ues of the same
ge nera (Gibbs, Collar d and Wood 2000). Natu r ally there are
disagree m e nts amo n g sp ecialis ts about the naming of species and the
attribution of particular sp ecime ns to particul ar position s in our
ancestry. One view of the phylogeny (Foley & Lahr 1997) suggests the
ho mini ns of modern Indo nesia do not seem to have cont ributed to the
modern popula tions in any region (thou gh they survive d for 700 000
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years). This is consis t ent with the more detailed analyses of the relations
among skel etal remain s in the Australian and East Asia region (Brown
1992; Storm 1995), and also with the developing cons e n s u s that modern
hum a n s emerged out of Africa less than 800 000 yea rs ago (Groves 1992;
Grov es & Lahr 1994; Rogers 1995; Rouh a ni 1989).
In unde r s t a n d i n g this lack of contributio n to modern huma ns from the
Asian Homo erect u s , two geogr a p h ic ba rriers appear fund a m e n t althos e
into Australia and into the America s. The Indones i an Ho mo erect u s did
not make it into Australia, and the East Asian one did not make it into the
northern region s of Asia or furt her, into the Ame rica s. We can evalu ate
the abilities of Ho mo erect us by con sideri n g the behavioural abilities of
the first peo ple who did make it into Australia (Davidson 1999a).
The eviden c e sugge s t s that both the Australian and the nort h e r n re gio ns
(inclu ding the Americas) were only coloni sed less than 60 thou sand year s
ago, after the emergence of communicatio n using symb ols (Davidson &
Noble 1998). Symbolic represe ntatio n is necessary for the plan ning and
anticipation involved in assembli ng the ma terial s to build a waterc r aft
and to have a purp ose in building it (Davidson & Noble 1992). It is also
necessary for ensuri n g the provisions nece s s a r y for the voyage. Similar
planning and anticipat ion were necessary on a much mor e continuing
basis for the coloni sation of cold enviro n m e n t s, to mak e suit able shelt e r
and provision for the winter months.
The eviden c e aroun d the world suggest s remarkably similar dates (on a
geological time- scale) for the earliest appearance of direct evidence for
symbols: 90 thousand in South Africa (Henshil wood & Sealy 1997; Singe r
& Wymer 1982); 35 thous an d in Europe (White 1989); 32 thousand in
Australia (Mors e 1993).
The Australian eviden c e , at the moment, is represented by the pier ced
se a- shells from Mandu Mandu, hun d r e d s of kilom etres from the sea at
the time of use (Morse 1993). As in Eu rope at the same time (White
1989), this suggests that among the earliest us es of symbolic
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re presentation was the use of bead s for pe rsonal decor ation—a rg u a b ly a
mea n s of dr awin g a line betwe e n bead wearers and others .
The use of sym bols in commu nication, necessary for the colonisatio n of
Australia, seems to be wide spread around the world at about the time of
that first colonisation (Davidson & Noble 1998), and was proba bly
necessary for some of the oth e r novel features of hu man behaviour that
eme rg ed in that time period. The issue for evaluating the evidence fro m
Flor es is whether there is evidence around the world for a simila r
context of emergence of langu a g e- base d abilities from that time period.
Beads and painted caves are merely obvious sign s of symb ols. The
watercraf t that brought people to Austra lia was much earlier, and Noble
and I (Davidson & Noble 1992) have argued that its constru ct ion
involved symbolic representation , without direct evidence of such
obvious signs. Was there evidence of such symbolic repre s e nt ati o n in
the earlier archaeolog ical record without the appe a r a n c e of objects th at
were the mselves symbolic?
The first, most obvio us claim is from ston e artefa cts, of whic h the most
re m arkable are the bifacially flaked objects archaeol ogists call Acheulean
The refutati on of this claim is still difficult, and very technic a l. I can not
go into details here (but see Davidson & Noble 1993; and Davidson 2000
xxxx). The main claim to “symb olicnature of Acheulean han daxes is
their astonishing uniform ity around the world for ove r a million yea rs
(Wynn 1979; 1995 ; Wynn & Tierson 1990). But the unifor mity may mask
a more co mplex reality which includes the selection by archaeolog ists of
thing s which are uniform becaus e they were sele cted (Dibble 1989), and
may involve a converg en ce on a form without any int entio n al selection by
the ma kers.
The clinching evidence is that Australia was coloni sed from a region said
never to have had handaxes (Schick 1994) and after the final appearance
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of stone indust ries based on handaxes (Cab rera Valdes 1989), yet there
ar e han daxes from Australia (Noble & Davidson 1996; Rainey 1991). The
producti on of bifacially flaked object s occurred many time s quit e
indep e n d e n tl y, as has been known and un derstood for many year s by
their appearance towards the end of the Europe a n Middle Palaeolithic
(Mellar s 1973). In this context, they wer e quit e separa te d from
Acheulea n indust ries in time and thus in every othe r way. Such
separatio n implies that they do not belong to a long tradition of
cultu r ally- designed objects.
The first Australians clea rly had the ability to impose form on the objects
they made. These were tthe ances t o r s of Austr alia n peoples such as the
Kalkad oons of Mt Isa (Davidson 1993) or the Anaiwan of the Armid ale
re gion (Godwi n 1997),. This was true both for the water c r aft they used
to get her e, and the othe rs th ey made to voyage off the northe as t edge of
the continent to the Bismarck Archipe lago (Allen & Gosd e n 1991).
But early tools also indicate this ability. In early sites, there are bone
tools shaped by grindin g unr estrained by the limitations of the raw
material , as were the ground- edged hatchet heads in the Huon Penin sula
and in Arnhem Land (Davidso n & Noble 1992). These hatchet heads
were mad e at least ten thousand of years before the appearance of
similar hatche t s in western Asia and Europe marked the beginning of the
destructi ve effects of agricult u r e.
The othe r great claim for early symbol use is the persisting belief that
the Neandertals of Euro p e and west e r n Asia buried their dead. I believe
this claim canno t survive the recent reanalysis by Rob Gargett (1989;
1999). He has show n that there are only two circumstanc e s in which the
skele tal remains of these homini ns (and the early mode r n humans of
Qafzeh) survived with almos t compl ete skeletons. Gargett has shown
that if they were complete, then they were killed by rockfall when the sky
fell in—“the Chicken Licken Effect . Sometimes the rock crushe d the m
like a foot stampi n g onto an emp ty beer can. Garget t also sho wed that if
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they were not crushed by rockfall, the n they seem to have decom p o s e d in
natural hollows in the sedimen t befor e those holes were naturally filled
with more dirt. Whole disar ticul a t ed segments are always missing,
indicati ng that partial deco m p ositio n of the body had taken plac e before
bu rial was complete. Ther e was somethi n g rotten in the st ate of
Ne andertals.
In Austr alia, th e story is quite different. The ea rli est burials in the world
ar e found here. The Mungo I cremation, burned, smashed, collecte d and
bu ried (Bowler et al. 1970), indicati ng by the repeated attention to the
body an order of magni tude differen t from anything even claimed for
Ne andertals, is dated to 26 thousand years ago (Bowler 1998). The
Mungo III burial (Bowler & Thorne 1976), quit e distinctively compl ete in
a way that Ne andertals are not, is probably as old as 40 000 year s (by
TL) (Bowle r 1998) but ther e is unc e rtainty about the dating (Brown &
Gillespi e 2000).
So metime after 100 thous a n d years ago somet hing major change d.
There were no people in the cold climat e s of the north at 50 tho usand.
After 40 thousand the north filled up, despit e the worse ning clim ate, and
so did Australia. Are these two events relat ed?
If the Ameri cas were first colonised at 14 thou sand year s ago (Beato n
1991) (and there is now some doubt about this [Meltzer et al. 1997 ]),
then it is worth pointing out that the previous time that such condi tion s
existed was back at 80 thousand. I believe that this evidence will
de monstrate that it was after 80 thous a n d that there was an emergence
of plan ning abilities. These planning abilities enable d the colonisa tion of
cold environment s, sea- cros sings such as were needed to get to
Australia, and whatev e r combinatio n of water- cross ing and cold climate
ad aptati on was needed to get to the Ame rica s.
We should not exaggerate the ease with which the new achieveme n t s
were won from the emergence of plannin g abilities. The dates for first
colonisa tion of different regions in Australia show a rapi d movement
from first landfall to Tasmani a , but a rel atively slow occup a tion of
“difficultenvi ronments. The short time between the earlies t dates in
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northern Australi a and those for Tasmani a fits with Birdsell’s (195 7)
ra pid colonisati on model prop osed long before archaeolo gical evidenc e
for the timing of colonisation on the basis of purely theoreti c al
simul ation s of the demo gr a p hic condi tion s for colonising all of Australia.
Rindos & Webb (1992) suggested an “optimal malada p t a tionmodel in
which peopl e not well adjust ed to the poorly known and un predict a b ly
varying Australian envir onme n t s died out or moved on rath e r than
staying to find ways of coping with resource variatio n locally.
This rapid assess ment of homini n abilities around the world at the time
of the ston e artefa cts of Flores suggests that there is little evidence
around the world 800 000 years ago, or indeed until much lat er, for a
ho mini n ability to impose form on artefacts . The evidence of the abiliti es
of the first people in Aust rali a suggest s that they were eng aged in
plann e d activities, and had sufficien t be havi oural flexibility to cope
ra pidly with a wide range of environmentsbut not all of the m. So what
can we say about the early stone artefa cts in Flores as a result of this
ra pid asse s s m e nt of the compa r isons with events contemporary wit h the
artefa cts, and with the majo r sea cros sing much later in the story?
The crossing of Wallac e’s line by homin ins was sig nificant, but it seems
unlikely that it involved the suit e of abilities, based on plan ning, that are
indicated by the first colonisation of Australia. Such planning does not
se em to be characteristi c of any hominin behaviour elsewhere at the time
of the crossing to Flore s. On the basis of the evide nce from their other
abilities reflec ted in the archaeolog ical recor d , I doubt hominins
constru ct ed a water c r a ft.
It may be that the artefact s in Flores indica t e a previously un documente d
ability of early Homo , but if th e sea crossing was short, we may need to
make compar i son with similar crossings elsew h e r e. It will be nec e ssary
to con sider how early Homo react e d to the larg e rivers it undoubtedl y
encountered elsewhere in the world. If rafti ng on vegetation ma ts or
logs is a serio us possibility (see Smith this volume), woul d this have be en
an option in crossin g the Nile, the Tigri s or the Gan ges?
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What Morwood has demonstrated by showing an early date for stone
artefa cts east of the Wallace Line is that previous finds of such
significan ce may have been ignored unjus tifiably. Many other regions
may not have been adequatel y explor e d as a result of the easy
acceptance of model s that appea r to account for the known evidence.
Where these mod els are aimed at under st a ndi n g the evolut ionary
eme rg e n c e of abilitie s we take for grante d , they are often based on
inade q u a t e concep t u a lisati o n of those abilities. We need new concept ual
models to provoke approp r iate research.
It is claimed (Morwood et al 199 8; Morwo od this volu me) that the Flores
finds represent the achievement of human abilities at 800 000 years ago,
which previousl y we re only documente d wit h the colonis ation of
Australia after 80 thou s a n d year s ago. In order to acce pt this, it is
essential to expl ain why, as I have sough t to show, the claim appears so
su rprisi ng in the contex t of what else is known about the behaviour of
ea rly homini n s. The ability to construct waterc r aft at this early date in
the Early Pleistocene demands an explanati on for the lack of coloni sation
of other island s around the world until the end of the Pleistoce ne or
later. The Medit e rr a n e an islands, many of which are visible from the
mainla n d, or beco m e so at glacial perio d low sea- levels, were not
colonise d until after 11 thousand yea rs ago (Cherry 1990). Any
development of theory that arises from the Flores finds must also
consi der why ther e is suc h a differenc e in dat es betwe e n the first sea-
crossi ngs in the se two regions. Human ancesto r s were pre sent in
Eu rope at about the same time as they cross e d into Flores (Pare s &
Pérez- Gonzalez 1995) but ther e were no crossings to the islan ds visible
off the coast. Did Middle Pleistocene Medite r r a n e an environ me nts not
produce rafts of vegetatio n in the way the island s of Wallacea do?
Hu m a n s were present in Europe by 40 000 years ago (Bocquet- Appel &
Demars. 2000), but their be haviour does not appe a r to have include d
mes sing about in boats. It is sugg e s t e d that fishing was an impo rtant
be havi our befo re the cros sing into Aust rali a (Balme 1995; Davidson
1999a). Was there any evide nce for fishin g in th e Medit erra n e a n
between 40 000 and 11 000 years ago? Hum an ne ss seems to be a
necessary conditi on for regul ar sea crossi n g, but not a sufficien t one.
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Noble and I have repeat e dly pointed out that the evidenc e from the first
hum a n colonisatio n of Austr alia is crucial to unde r s t a n d i n g the
evoluti onary emergence of language. I per sist in this claim in the face of
the finds from Flor es. Somet h ing different happened shortly before first
colonisa tion of Australia, which has no parallel in the world of homi nins
at the tim e of the first stone tools in Flores. If we are right, then the
nature of the first colo nisa tion must be understood in ter m s of the
symbolic con struction of the physical and social envir onm e n t s by the
first people to make the corss ing. I have begun to develop that argument
elsew h e r e (Davidson 1999a) .
There are now sufficie nt numbers of sites excavat e d and dated in most
re gion s of Australia that we can begin consid er details of the patt e r n of
colonisa tion. We need to consid e r the changing patterns of adapt a tion
which ma de it possible for people to cope wit h difficult environ ments
(Davidson 1990; O'Connor et al. 1993). But, along with the story of how
the descendants of a small number of initial coloni sts (speakin g a small
num b e r of languages) succeeded in occupying the whole of the
conti nent, there is another story that has not been told. That is a story of
the em e rgenc e of the many different peoples who lived in Aust rali a at the
time of the invasion from Europ e. How did the Australi an continent
(Sahul) come to have so many differen t people, so man y langua ge s? In
the perio d of historic docu m e n ta ti on, there were more than 250
langu a g e s on the mainland subdivid ed (by some) into 27 families (Wurm
1972). New Guinea (connected to the mainland by Pleistoce ne low sea-
level until 8 thous a n d years ago) had 200 languages in the Austronesian
family, and abou t 700 Papu a n lang uages in about 60 families (Dixon
1997). In seeking to explain the generati on of such diversity, the
theoreti c al argume n t s mu st be consistent with those that accoun t for the
long delay betw een the sea- crossi ng to Flores and that to Austr alia.
I believe the argum ent Noble and I have prod ucedthat only the second
crossi ng reflects the recent emergence of languagemeets the
re quirement of accounting for different classes of eviden c e before and
afte r coloni sation (Noble & Davidson 1996). The diversi ty that emerged
in Aboriginal Austra lia proba bly resul ts from the human propensi ty to
IA I N D A VI D S O N 1 0
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dr aw and define lines betwee n categ ories, the common result of the use
of lang uage. Wallace was doing something distinct ively human when he
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... Among the fauna of this continent, there are many examples of species extinction of vertebrates due to hunting, introduction of foreign species, and various forms of environment degradation caused by human settlement, which started in the 19 th century (see e.g. Matcalfe et al. 2001; Invasive Species Fact Sheet: The feral cat (Felis catus), publications/pubs/cat.pdf; ...
The authors of the study present an analysis of the structure and changes in the examined community of ptyctimous mites (Acari: Acariformes: Oribatida) found in Dorrigo National Park in Australia. The research was conducted during two periods: between 1990 and 1993 and later in 2007. The analysed mite community comprises 35 species, though, the dominance and frequency of particular species were different for each period. In the first research period (1990–1993) in the area of Dorrigo National Park, 28 species were recorded, whereas in 2007 – 23 species were found. There were 16 species that occurred in both research periods, and 12 species only in the samples collected in the 90’s, and 7 species only in those collected in 2007. The analysis also embraces the geographical distribution of the species in the area of Australia. Three species were designated as endemic, occurring only in the area of the examined national park (Austrophthiracarus dissonus Niedbała et Collof, 1997, Austrophthiracarus parapulchellus Niedbała, 2006 and Notophthiracarus distinctus Niedbała, 1989). The analysed samples contained only few specimens of these species. Due to the low abundance, great rarity of the local populations and high endemism, these species should be regarded as potentially endangered (EN according to the IUCN scale). A comparative analysis of the community from Dorrigo National Park (New South Wales) with those found in other larger areas of Australia in Victoria (Otway Ranges Area, Yarra Ranges Area, Strzelecki Ranges Area and Errinundra Plateau Area) examined by Niedbała and Szywilewska-Szczykutowicz (2017) has revealed that the communities found in Dorrigo National Park contained far more species, which constituted 30% of the whole fauna of Australia. In contrast to the communities of ptyctimous mites from Dorrigo, the individual communities in the area of Victoria contained only between 5% and 14% of all known species in Australia from this group.
... Core areas of Malesia accreted from terranes that broke off from Gondwana (which was part of Pangaea until the Late Jurassic) and moved northward through the Mesozoic (Metcalfe et al. 2001). These areas represent the first involvement of Gondwanan terranes with the formation of Southeast Asia and presumably included some floristic contributions. ...
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Unraveling the origins of Malesia's once vast, hyperdiverse rainforests is a perennial challenge. Major contributions to rainforest assembly came from floristic elements carried on the Indian Plate and montane elements from the Australian Plate (Sahul). The Sahul component is now understood to include substantial two-way exchanges with Sunda inclusive of lowland taxa. Evidence for the relative contributions of the great Asiatic floristic interchanges (GAFIs) with India and Sahul, respectively, to the flora of Malesia comes from contemporary lineage distributions, the fossil record, time-calibrated phylogenies, functional traits, and the spatial structure of genetic diversity. Functional-trait and biome conservatism are noted features of montane austral lineages from Sahul (e.g., diverse Podocarpaceae), whereas the abundance and diversity of lowland lineages, including Syzygium (Myrtaceae) and the Asian dipterocarps (Dipterocarpoideae), reflect a less well understood combination of dispersal, ecology, and adaptive radiations. Thus, Malesian rainforest assembly has been shaped by sharply contrasting evolutionary origins and biogeographic histories. Expected final online publication date for the Annual Review of Ecology, Evolution, and Systematics Volume 50 is November 4, 2019. Please see for revised estimates.
... Metcalfe et al., 2001; Bruyn et al., 2014) and into the Indian subcontinent, no earlier than the Paleogene when it collided with the Eurasian Plate (e.g.Aitchison et al., 2007). The morphological similarity of western Laurasian Campanian †Palaeoleptochromus and south-eastern Laurasian Cenomanian Clidicus (= Cretoleptochromus), and the location of the present-day biodiversity center of Clidicus in SE Asia support this scenario of Clidicini evolution and dispersal. ...
Mastigitae is a supertribe of ant-like stone beetles (Scydmaeninae) that includes over 40% extinct genera, and whose evolutionary history is documented from the Upper Cretaceous through to today. Based on the results of phylogenetic analyses combining the most extensive taxon sampling to date, we reclassify Mastigitae into six monophyletic units: Leptomastacini, Clidicini sensu nov., Papusini trib. nov., Leptochromini trib. nov., †Baltostigini trib. nov. and Mastigini sensu nov. †Cretoleptochromus is placed as a junior synonym of Clidicus, and †Cascomastigus as a junior synonym of †Clidicostigus. Euroleptochromus setifer sp. nov. and Baltostigus striatipennis sp. nov. are both described from Upper Eocene Baltic amber. We postulate that Mastigitae have undergone differentiation into major lineages before the Cenomanian and that the Eurasian part of Laurasia was their ancestral distribution area. The reconstructed ancestor of Mastigitae was similar to the extant Scydmaenini, but with broadly separated antennal insertions and deep elytral striae. Four independent wing losses were inferred in Mastigitae. We present the first complete μCT reconstruction of the aedeagus in a fossilized scydmaenine, crucial for understanding the more than 99 million year long evolution of one of the most bizarre, asymmetrical aedeagi in the Coleoptera.
... The environmental values of Indonesia's forests are globally significant. Indonesia's position as an archipelagic bridge between Asia and Australia means that it evolved extraordinary and globally significant biological diversity (Metcalfe et al. 2001). Indonesia's forests rank first in the world in endemic birds and mammals (Birdlife International 2010; Schipper et al. 2008) and sixth in endemic amphibians (Stuart et al. 2004). ...
In recent months, strong global growth, rebounding commodity prices, and relatively accommodative financial conditions have benefited the Indonesian economy. The first quarter of 2017 in Indonesia saw resilient GDP growth, moderate inflation, stable exchange rates, an increase in the growth of non-oil exports, and an investment upgrade from ratings agency Standard & Poor's. Investment growth, however, did not pick up enough to drive overall growth to a higher rate. The poor quality of banking-sector assets and the gaps in tax revenue—despite the fulfilment of the government's tax-amnesty program—are two of the most immediate economic concerns. President Joko Widodo (Jokowi), who is well into the second half of his term, is under pressure to deliver on his development platform, which includes making progress in sustainable development and climate change mitigation. The effective management of forests is key to this platform. There has been longstanding tension over Indonesia's forests between the protection of environmental values, including carbon storage, and the production of valuable commodities, including timber, palm oil, and pulpwood, which generate revenue and employment. We survey recent developments in four storylines related to forestry and climate change: first, Indonesia's commitment to reducing emissions to 29%–41% below projected business-as-usual levels by 2030, as well as the international climate agreements and finance that can help achieve this commitment; second, land-use rights and regulations, including a moratorium on clearing, draining, or setting fires on peatland; third, measures to prevent catastrophic forest fires like those during the 2015 El Niño, including the establishment of the Peatland Restoration Agency; and, fourth, the actions of non-state actors, especially large agribusinesses, in managing forests and peatland. We conclude by discussing differences in the approaches of Jokowi's administration and those of former president Susilo Bambang Yudhoyono's administration and by questioning whether Indonesia's budgeted resources, actions, and results to date are commensurate with its climate commitments.
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ABSTRACT A growing corpus of finds dated to the Palaeolithic from islands in the Aegean has changed the archaeological community’s perspective on the earliest occupation of oceanic islands and the capacity of pre-sapiens hominins to make successful sea crossings. Some scholars have chosen to relate these marine dispersals with ancestral species such as Homo erectus or Homo heidelbergensis. This data has been compared with evidence of Pleistocene seafaring from the Pacific Ocean, as well as proposed sea-crossings of the Atlantic Ocean. Although the correlation of local finds with global perspectives of the past can and has been fruitful in many cases, the marine-route hominin dispersals theory in the Mediterranean has not thus far been well documented based more on inference rather than direct evidence. The majority of scholars deem the existing evidence, mainly from the Aegean, insufficient to support systematic sea faring activities on the Mediterranean or a sustained hominin presence on the Greek islands. The suggestion that a non-modern species, such Homo erectus, had inhabited islands in the Aegean or the Pacific during the Lower Palaeolithic is difficult to prove on the existing corpus of evidence, however it may equally be difficult to disprove. The debate has led both sides to extreme views, ultimately offering little to the better understanding of human mobility and modern behaviour during the Pleistocene. An alternative approach to the idea of early seaward dispersals will be discussed, interpreting local data based on the concept of glocalisation.
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This paper reviews some of the main advances in our understanding of human evolution over the last 1 million years, presenting a holistic overview of a field defined by interdisciplinary approaches to studying the origins of our species. We begin by briefly summarizing the climatic context across the Old World for the last 1 million years before directly addressing the fossil and archaeological records. The main themes in this work explore (i) recent discoveries in the fossil record over the last 15 years, such as Homo naledi and Homo floresiensis; (ii) the implications of palaeogenetics for understanding the evolutionary history of, and relationships between, Neanderthals, Denisovans and Homo sapiens; (iii) the interplay between physiology and metabolic demand, landscape use, and behavioural adaptations in the evolution of morphological and behavioural innovation; and (iv) recent advances in archaeological understanding for the behavioural record, in particular that of the Neanderthals. This paper seeks to provide a broad‐scale, holistic perspective of our current understanding of human evolution for the last 1 Ma, providing a reference point for researchers that can be built upon as new discoveries continue to develop the landscapes of human evolution.
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Since the mid 19th century, the biogeography of island South-East Asia has been the subject of much study. Early researchers explained many of the species distribution patterns by the rise and fall of sea levels and land. This and the work of other researchers culminated in a theory that emphasized the role of Pleistocene sea level low stands in species evolution. With the advent of newly developed molecular techniques, however, it became clear that many species divergence events had taken place before the Pleistocene and a biogeographical theory focusing on Pleistocene sea level changes was inadequate. In this research, I have developed a new biogeographic model that explains present-day distribution patterns and evolutionary relationships between species. I use this new model to explain 10 ‘mammalian riddles’, i.e. evolutionary or distribution patterns in selected mammal species groups that could not be explained with the existing theories. I developed the new model by analyzing the geological literature for this region, and by mapping palaeogeographical and palaeoenvironmental changes for the last 20 million years. In addition I compiled information on the palaeontological record for the region and on divergence times between taxa using a molecular clock assumption. These phylogenetic data were compared with the palaeomaps to assess whether particular divergence events could be correlated with certain palaeogeographical or palaeoenvironmental changes. The combination of these two information sources has resulted in a much-improved understanding of mammalian evolution in island SE Asia. Using this model it is now possible to relate important palaeoenvironmental events, such as the Late Miocene cooling, an Early–Middle Pliocene highstand, or the emergence and submergence of a land bridge between the Malay Peninsula and Java to evolutionary changes in species. I test the accuracy of the new model by analysing the relationships within several mammal groups using craniometric and molecular analysis. The observed relationships and deduced timing of divergence between taxa could in many cases be explained by the model, which indicates that it is relatively accurate. In addition, with the new model I have been able to find solutions to most mammalian riddles, although these results require further testing. Overall, I therefore believe I have made a significant contribution to the biogeographical understanding of island SE Asia.
Amborella trichopoda (Amborellaceae) is a shrub endemic to New Caledonia in the Southwest Pacific region. This plant suddenly became famous when molecular phylogenetic studies revealed that this sole species is likely the sister taxon to all other angiosperms. It has thus been a prime research model for reconstructing plant evolution and gaining insight into what the earliest angiosperms looked like. A wealth of studies on Amborella have now shed considerable light on its genome, morphology, anatomy, physiology, development, and architecture – this research is reviewed in this article. While Amborella likely retained some ancestral traits, critical character reconstructions have also highlighted some derived and sometimes unique characters in this species. The history of Amborella is also tied to the South Pacific archipelago of New Caledonia, its homeland. It was part of the New Caledonian biogeography puzzle and its genetic history shed light on the dynamics of its ecosystem, the rainforest understorey. Amborella is now cultivated in botanical gardens and has been the focus of some conservation measures that will also benefit other species in this biodiversity hotspot.
Natural rafting is an easy, non‐evidence‐based solution often used to explain the presence of a variety of species on isolated islands. The question arises as to whether this solution is based on solid scientific grounds. It is a plausible colonisation route only if intricate networks of variables are considered and many different conditions satisfied. This review provides a descriptive account of some of the most critical issues underlying the theory of natural rafting that should be addressed by its supporters. These include: (i) biological variables; (ii) characteristics of the vessels; and (iii) physical variables. Natural rafting may explain the dispersal of poikilotherms with low metabolic rates and low resource requirements that could withstand trans‐oceanic crossings, but explaining the transport of homeothermic terrestrial mammals to oceanic islands is more problematic. Drifting at sea exposes organisms to high concentrations of salt, high temperature and humidity excursions, starvation, and above all to dehydration. A sufficiently large group of healthy reproductive individuals of the two sexes should either be transported together, or be able to reassemble after separate crossings, to prevent inbreeding, genetic drift and ultimately extinction. Any vessels of flotsam occupied must minimally provide the animals they transport with sufficient provisions to survive the journey, offer minimum friction and drag through water, and be transported by appropriately directed, sustained, high‐speed currents. Thus, a ‘sweepstakes colonisation’ event would be the result of a lucky combination of all, or at least the majority, of these factors. Some cases throw doubt on the use of a natural rafting model to explain known animal colonisations, with one of the most striking examples being Madagascar. This island is far from the nearest mainland coasts and the sea currents in the Mozambique Channel are directed towards Africa rather than Madagascar, yet, the island was colonised by terrestrial mammals (e.g. extinct hippopotamuses, lemurs, carnivores, rodents and tenrecs) unable to swim and to survive long journeys at sea. In order to assess the feasibility of the natural rafting model in a case such as Madagascar, tests were performed using three variables for which enough information could be obtained from the literature: length of survival without food, survival without water, and sea current speed. The distributions of these variables appear to be log‐normal and multiplicative, or follow a power‐law, rather than being Gaussian. The tests suggest that a distributional analysis is a more suitable approach than the use of geometric probability to calculate the probabilities associated with the examined data. Such non‐linear and self‐organising systems may reach a critical point governed by different competing factors. Mammals with high survival requirements, such as lemurs and hippopotamuses, thus may have a virtually zero probability of reaching distant islands by natural rafting. Our results raise doubts as to the validity of a natural rafting model, and we urge a rethinking of the modes in which numerous islands were colonised by land mammals and a careful revision of past geological and phylogeographic work.
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Sundaland, the continental core of SE Asia, is a heterogeneous collage of continental blocks and volcanic arcs bounded by narrow suture zones that represent the remnants of ancient ocean basins. All the continental blocks of Sundaland were derived directly or indirectly from the Arabia-India–Australia margin of eastern Gondwana by the opening and closure of three successive ocean basins, the Palaeo-Tethys (Devonian-Triassic), Meso-Tethys (Permian-Cretaceous) and Ceno-Tethys (Jurassic-Cretaceous), and assembled by the closure of these ocean basins. Core Sundaland comprises a western Sibumasu block and an eastern Indochina–East Malaya block with an island arc terrane, the Sukhothai Island Arc, sandwiched between. The Palaeo-Tethys is represented by the Changning–Menglian, Chiang Mai-Chiang Rai, Chanthaburi and Bentong–Raub Suture Zones that form the boundary between Sibumasu and the Sukhothai Arc. The Indochina block was derived from Gondwana in the Devonian when the Palaeo-Tethys opened. The Sukhothai Arc formed on the margin of Indochina in the Carboniferous, and then separated by back-arc spreading in the Permian. The Jinghong, Nan–Uttaradit and Sra Kaeo Sutures represent this closed back-arc basin. The Sibumasu Terrane separated from Gondwana in the late Early Permian when the Meso-Tethys opened and collided with the Sukhothai Arc and Indochina in the Middle-Late Triassic. The Cathaysian West Sumatra block possibly represents a part of the Sukhothai Arc and was emplaced by strike-slip tectonics outboard of Sibumasu in the Triassic. The West Burma Block was already attached to Sundaland before the Late Triassic and is likely a disrupted part of Sibumasu. East Java-West Sulawesi and South West Borneo are tentatively identified as the missing “Argoland” and “Banda” blocks which must have separated from NW Australia in the Jurassic and subsequently accreted to SE Sundaland in the Cretaceous.
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