Chapter 4: Subtropical Forests

Richard T Corlett and Alice C. Hughes

Center for Integrative Conservation, Xishuangbanna Tropical Botanical Gardens, Chinese
Academy of Sciences, Menglun, Yunnan 666303, China

Citation: Corlett, R.T. and Hughes, A.C. (2015) Subtropical forests. In: Peh, K. S.-H., Corlett,
R.T. & Bergeron, Y. (eds.) The Routledge Handbook of Forest Ecology. Routledge, Oxford, UK,
pp. 46-55. ISBN 978-0-415-73545-2

Introduction
There is no widely agreed definition of what constitutes the ‘subtropics’, but in recent ecological
literature the term has been most often applied to the two belts between the tropics (+/-23.4o) and
approximately 30o north and south of the equator (Corlett, 2013; Figure 1). These boundaries
will be used in this chapter. Although this choice is fairly arbitrary, particularly at the outer (30o)
limit, applying a uniform standard makes it possible to compare forests in the same latitudinal
belt in different parts of the world, as well to make comparisons within regions between
subtropical forests and the forests of the tropical (i.e. <23.4o from the equator) and temperate (i.e.
>30o) zones. Moreover, defining the subtropics by latitude alone avoids the confusion between
latitudinal and altitudinal gradients that occurs in some of the literature.
Much of the subtropical belt, as defined above, is too dry (mean annual rainfall < 600-
700 mm) for forest because of the descending branches of the Hadley circulation (the subtropical
high), but there are, or were, extensive forests on the southeastern side of all the continents,
particularly in China and adjacent countries (71% of the total current forest area in the subtropics
according to the MODIS landcover layer for 2012), and in northern Argentina and southeastern
Brazil (23%), with smaller areas in northern Mexico and the southeast USA (4%), eastern
Australia (2%), and southern Africa (1%) and a very small area in southeastern Madagascar.
These extant areas of subtropical forest represent only part of their potential extent; around 40%
if forest is assumed to have covered all land areas < 4000 m a.s.l. and with a mean annual rainfall
> 700 mm (Figure 1). Moreover, much of the extant forest is secondary, degraded, or consists of
plantations. The miss- match between actual and ‘potential’ forest is greatest in southern Africa,
where climate predicts a much larger area than the scattered patches that exist.
Common features of the climates of these forested regions, apart from adequate rainfall,
include lowland temperatures suitable for plant growth over most or all of the year, despite the
occurrence of at least mild winter frosts in most areas in most years. In coastal regions, except in
South America, subtropical forests are vulnerable to cyclones, with repeated occurrences
reducing stature and biomass (e.g. McEwan et al., 2011). There have also been reports of
damage by ice-storms. In early 2008, the longest cold spell for more than 40 years brought snow
and ice-storms to a broad band of subtropical China, north of around 25°N, causing massive
mechanical damage to native broad-leaved forests and devastation to many plantations (Zhou et
al., 2011).
Subtropical forests have been intensively studied in China and have received
considerable attention in South America, but studies elsewhere have been largely descriptive.
The results of these studies are scattered across multiple journals, in multiple languages, since
there is no subtropical equivalent of the international journals that focus on the tropics.
Moreover, inconsistent nomenclature makes it hard to retrieve these papers electronically. The
overview presented here is thus inevitably incomplete, but we hope it will encourage more ‘pan-
subtropical’ comparisons.

Biogeography of the subtropics

The subtropics of the northern and southern hemisphere are separated by the 5,185-km width of
the tropics. The forests of the northern subtropics, in Asia and North America, are floristically
quite similar to each other, reflecting the availability of land routes across the North Atlantic
during the early Tertiary, until global cooling broke this connection, probably at the end of the
Eocene, c. 34 million years ago (Givnish and Renner, 2004). In contrast, the subtropical forests
in the southern hemisphere have less in common with each other than they do with their nearest
northern counterparts. Although these southern forests are all on fragments of the ancient
southern supercontinent of Gondwana, the last terrestrial connections were in the early
Cretaceous (145–100 million years ago), before most modern groups of plants originated, while
tropical montane habitats have provided more recent stepping-stones from the north, across the
tropical lowlands.
In southwest China, subtropical forest floras similar those of today have existed since the
early Miocene (Jacques et al., 2013), although some tropical elements in the Miocene flora have
retreated to the south in response to global cooling and regional uplift (Jacques et al., 2014). In
North Africa, in contrast, the shrinkage of the Tethys Sea during the Late Miocene drastically
weakened the African summer monsoon, creating conditions at subtropical latitudes that were
too arid for forest (Zhang et al., 2014). A relic of the North African Miocene forests persisted in
the subtropical Canary Islands, which were buffered against excessive drying, but these isolated
forests have been impoverished by their small total area, the difficulties of dispersal over marine
barriers and changes in climate (del Arco Aguilar et al., 2010, Fernández-Palacios et al., 2011).
In eastern Australia, the rapid northward movement of the continent since the Eocene has
both progressively reduced the extent of subtropical rainforests and brought the region closer to
the extensive forests and floras of Southeast Asia. As a result, there is a north-south gradient in
the relative contribution of recent immigrants from the Asian tropics and ancient Gondwanan
lineages to humid forest floras, with immigrants enriching but not replacing the indigenous
clades in subtropical forests (Sniderman and Jordan, 2011). The South American subtropics has
a poorer paleoecological record, but this shows the replacement of subtropical forests by arid
vegetation west of the Andes from the Middle Miocene onwards as the Andes rose and the cold
Humboldt Current was established (Le Roux, 2012).
Northern subtropics
In the northern subtropics, forests are most extensive by far in East Asia. There is a much smaller
area in North America and none in North Africa, although a relic of the Miocene forests of North
Africa and southern Europe persists in the Canary Islands. The most characteristic common
feature of these forests is the abundance and diversity of Fagaceae (oaks and other genera) and,
particularly following disturbance and at higher altitudes, the abundance of pines (Pinus).
Fagaceae are most diverse at both the species and genus level in subtropical Asia. Pines attain
their greatest diversity in the subtropical forests of the Sierra Madre Occidental of Mexico (Cord
et al., 2014), which is also a secondary centre of diversity for oaks (Quercus). Other genera
shared between East Asia and North America include Abies, Alnus, Ilex, Magnolia and Picea.

Asia
Subtropical forests were once very extensive in eastern China and the southern islands of Japan.
They used to cover 25% of China’s total land area, but this zone is highly suitable for agriculture
and has a high human population density. Most of the original forest has now been cleared and
the remnants are mostly badly degraded, except at high altitude (Corlett, 2014). Some of the
best-preserved examples are on Taiwan and the Ryukyu Islands. The total forest area is now
increasing in the Chinese subtropics, but most of this is young secondary forests and
monoculture plantations, often of exotic species.
Under the influence of the East Asian monsoon, the subtropical forests of East Asia
receive more rainfall (900-2000 mm) than most forests at these latitudes and the winter dry
season is ameliorated by cooler weather that reduces evapotranspiration. As a result, these forests
are composed largely of broad-leaved evergreen trees, with the Fagaceae and Lauraceae the
dominant tree families. Common genera including Ilex (Aquifoliaceae), Elaeocarpus
(Elaeocarpaceae), Castanopsis, Cyclobalanopsis (Quercus) and Lithocarpus (Fagaceae),
Distylium (Hamamelidaceae), Beilschmiedia, Cinnamomum, Cryptocarya, Lindera, Machilus,
Neolitsea, and Phoebe (Lauraceae), Magnolia and Michelia (Magnoliaceae), Symplocos
(Symplocaceae) and Schima (Theaceae) (Corlett, 2014). Young secondary forests, in contrast,
are often dominated by pines (e.g., Pinus massoniana, P. yunnanensis) or sometimes by winter-
deciduous species (e.g. Alnus nepalensis) (Tang et al., 2010). In cold, dry areas of northern
Yunnan, there is a distinctive evergreen sclerophyllous forest, dominated by hard-leaved
Quercus species (Tang, 2006).
Similar broad-leaved evergreen forests extend west across northern Myanmar and along
the foothills of the Himalayas (Davis, 1960; Singh and Singh, 1987), although pine forests are
more extensive here, probably reflecting the long history of human impacts. At higher altitudes,
the broad-leaved evergreen forests give way to forests dominated by conifers (Abies, Picea,
Pinus, Tsuga etc.) and broad-leaved deciduous species. Slope aspect appears to have a large
impact on forest composition in many parts of subtropical Asia but there has been little
systematic study of this.
In eastern China, the subtropical lowlands have been almost entirely deforested, but in
Myanmar, Northern India, Bhutan and Nepal, they support mostly dry-season deciduous or semi-
deciduous forests with a distinctly tropical flora. The semi-deciduous dipterocarp, Shorea
robusta (sal), dominates large areas, and in both Northeast India and northern Myanmar, lowland
evergreen rainforest with dipterocarps occurs at 27-28oN (Proctor et al., 1998). These forests
experience annual minimum temperatures below 10oC and most suffer at least occasional frosts,
but the impacts of winter cold on their structure and species composition are not obvious. It
seems likely that chilling-sensitive tropical taxa have been excluded, but testing this will require
comparative studies on a north-south transect across the Tropic of Cancer.
Eddy covariance measurements of carbon fluxes show that East Asian subtropical forests
are currently major carbon sinks: more important on a per-area basis than tropical and temperate
forests (Yu et al., 2014). Net Ecosystem Productivity (NEP) between 20o and 40oN shows a
strong negative relationship with forest age and a strong positive relationship with wet nitrogen
deposition (from industry and agriculture), suggesting that both recovery from past disturbance
and nitrogen fertilization contribute to the high carbon uptake. Year-round photosynthesis is
possible in at least some sites (Yan et al., 2013). Moreover, subtropical broad-leaved evergreen
forests dominated by Fagaceae can attain surprisingly high above-ground biomasses (c. 70-220
Mg C ha−1 in Taiwan; McEwan et al., 2011), making them among the most carbon-dense forests
in the world.

North America
The largest forest area in the North American subtropics is on the Sierra Madre Occidental, a
mountain range in northwestern Mexico that runs parallel to the Pacific coast and attains a
maximum altitude of 3300 m. The forests here are largely dominated by oaks (Quercus spp.),
pines (Pinus spp.), or both, with pine dominance increasing with altitude (González-Elizondo et
al., 2012). This mountain range (including the southern, tropical, parts) supports 54 species of
Quercus and 24 species of Pinus. Relatively minor forest types include a mixed coniferous forest
at higher altitudes, dominated by Abies, Pseudotsuga and/or Picea, along with oaks and pines.
There are also small areas of montane cloud forest with Magnolia, Styrax, Cedrela, Ilex, Tilia,
oaks and various Lauraceae. The Sierra Madre Oriental, in eastern Mexico, also supports diverse
pine-oak forests in the more humid areas. At lower altitudes on the dry northeastern coastal
plains, in areas with accessible ground water, there are forests with a tree layer dominated by
genera of Nearctic affinities (Alnus, Carya, Platanus, Quercus, Salix, Ulmus) (Encina-
Domínguez et al., 2011). Similar oak and pine-dominated forests occur in the southeastern
United States, where most have been severely disturbed, and on some of the islands of the
Bahamas.

Canary Islands
Remnants of subtropical broad-leaved forests persist on several of the Canary Islands. Similar
forests occur north of the subtropics on the islands of Madeira and the Azores, where oceanic
buffering has produced climates resembling those of the continental subtropics (Fernández-
Palacios et al., 2011). The Fagaceae were represented in the Canary Islands by an unidentified
species of Quercus in the Holocene forests of Tenerife, but this declined to extinction after
human settlement. The surviving broad-leaved evergreen forests are dominated by the family
Lauraceae (del Arco Aguilar et al., 2010). There are also extensive pine forests at higher
altitudes and in drier areas, and small areas of sclerophyllous woodland at lower altitudes.

Southern subtropics
The absence of Fagaceae and native pines distinguishes the forests of the southern subtropics
from those of the north, although pines are now widely planted and have become invasive in
some areas (Simberloff et al., 2010). Gondwanan elements (e.g. Cunoniaceae, Proteaceae.
Podocarpaceae) are usually present, but most other species in these forests appear to have
tropical (or northern subtropical) rather than ‘southern’ origins.

Madagascar
The subtropical southeastern section of Madagascar has steep altitudinal and rainfall gradients,
resulting in a wide range of forest habitats, including coastal forests on sandy soils, lowland and
montane humid forests and dry spiny forests, as well as a transition between rainforest and spiny
forest (Goodman 1999; Helme and Rakotomalaza 1999; Rakotomalaza and Messmer 1999). The
flora of these forests is essentially tropical, which may reflect the absence of land to the south,
but elevational zones are lower than is typical in the tropics to the north.

Southern Africa
Southern Africa is mostly too dry for forests, but there are a few thousand square kilometres of
forest patches scattered through the wetter areas inland from the east coast (Midgley et al.,
1997). These forests have been divided into two basic types, the interior Afromontane forests,
mostly above 1000 m elevation, and the coastal lowland Indian Ocean forests, which can then be
divided further on the basis of floristics and habitats. However, there is considerable overlap in
the flora of the various types, except at the extremes. Giant emergent Podocarpaceae with shade-
tolerant seedlings are characteristic of old-growth Afromontane forests, but angiosperms
dominate after human disturbance (Adie et al., 2013). In the northern subtropics of South Africa
and Mozambique, a distinctive ‘sand forest’ with a more tropical flora grows on deep sands
(Gaugris et al., 2008).

South America
In northwest Argentina, subtropical forests occur across a wide range of environments, from the
dry (< 1000 mm), warm Chaco lowlands, to the cool, moist (< 3000 mm) Yungas montane
forests (Ferrero et al., 2013), although the complex topography of this region produces many
different combinations of rainfall and temperature, so these forest are not easily arranged along a
continuum. Most of the surviving natural vegetation in the dry Chaco region would not meet
most definitions of forest, but there are also areas where soil, topography and rainfall (> 500
mm) allows the formation of a more or less closed woody canopy. In comparison with tropical
forests with similar dry season length, these subtropical forests are lower, structurally simpler,
smaller-leaved and floristically less diverse (Sarmiento, 1972). In comparison with the
subtropical forests China, those in South American generally have a larger deciduous
component, presumably reflecting the harsher dry seasons.
In northeast Argentina, where there is no marked dry season, the subtropical forests
represent an extension of the Atlantic forests of Brazil, including the largest remaining
continuous area. As in subtropical China, satellite measures of greenness (NDVI) suggest that
humid subtropical forests maintain high photosynthetic capacity throughout the year and could
thus be large carbon sinks (Cristiano et al., 2014). At the same latitude in the extreme south of
Brazil, the subtropical Atlantic forest can be classified into rainforests, semi-deciduous,
deciduous and mixed confer-hardwood forests, with the southern conifer, Araucaria angustifolia,
forming the upper canopy over a diverse broad-leaved middle and lower storey (Souza, 2007).
In the southern subtropics of Chile, to the west of the Andes, small areas of evergreen
forest occur in patches on coastal mountains facing the Pacific, where they depend on fog
subsidies in areas receiving only 150 mm of rainfall (van Zonneveld et al., 2012). Easter Island,
3700 km west of South America, apparently supported a palm-dominated forest until human-
settlement in recent millennia (Cañellas-Boltà et al., 2013).

Australia
Most of Australia is too dry for forest, particularly in the subtropical zone, but a broad belt along
the east coast receives enough rainfall (> c. 400 mm) to support eucalypt (Eucalyptus spp.)
woodland and the narrow strip nearest the coast (> 800-1000 mm rainfall) supports largely
evergreen rainforest patches in a matrix of tall open eucalypt forest, with a more or less dense
understorey. Rainforest was much more extensive at subtropical latitudes in Australia in the early
Tertiary until the late Eocene, but has become severely restricted subsequently. The richest
rainforest floras are found in areas where there is evidence for continuity of a moist climate
(Weber et al., 2014). These areas shelter both ancient Gondwanic lineages and species derived
from Asian lineages that invaded Australia from the Miocene onwards.

Discussion
The subtropics have usually been viewed by biologists as merely a transition between the tropics
and the temperate zone, although there has been little agreement on where exactly this transition
occurs (Corlett, 2013). Are subtropical forests sufficiently distinct from tropical and temperate
forests to be worth distinguishing as a separate ecological entity? Although the precise latitudes
of the Tropics of Cancer and Capricorn—the equatorward boundaries of the subtropics—are
ecological arbitrary, they are generally fairly close to the ‘frost line’, which seems to be a general
barrier to tropical plants because of the devastating impact of ice-crystal formation inside plant
cells (Corlett, 2014). In China, the poleward limit of the subtropics also coincides, more or less,
with an apparent eco-physiological boundary, where the dominant broad-leaved evergreen trees
give way to winter-deciduous species. This limit seems to coincide with an absolute minimum
temperature of around −15°C degrees (corresponding to a January mean of around 0°C) (Corlett,
2014). However, it is not clear from the available literature if a similar boundary occurs in other
parts of the world. Subtropical faunas, in contrast, are generally subsets of those nearer the
equator. This presumably reflects the ability of many animals to avoid short periods of extreme
cold behaviourally or, for mammals and birds, by thermoregulation, so that their poleward limits
are more likely to be set by a seasonal gap in food supply than by climate directly (Corlett,
2014).
In China, where there is a good paleoecological record, forests that would be considered
subtropical today have expanded north in warmer periods and retreated south in colder ones, to
be replaced by broad-leaved deciduous forests or more open vegetation types (Corlett, 2014; Ni
et al., 2014). This suggests that it is the climate rather than the latitude that makes a ‘subtropical
forest’. However, while mean temperature isotherms have shifted north and south with global
climate change, temperature seasonality is largely a function of latitude, so thermal regimes have
not simply shifted back and forwards.
Subtropical forests, as defined here, are probably less extensive today than at any time in
the last 20-30 million years. Long-term drying has limited their extent, except in monsoon Asia,
and human impacts have further reduced their area in those regions where suitable climates still
exist. These forests have received much less attention from conservationists than those in the
tropics, despite their importance for both biodiversity and carbon sequestration, but much of the
remaining forest area is included within Conservation International’s 35 global biodiversity
hotspots (Sloan et al., 2014). In East Asia, breeding-bird diversity reaches a maximum at around
25°N, rather than near the equator, with the larger land area available at subtropical latitudes than
on the tropical peninsula and islands further south a plausible explanation for this pattern (Ding
et al., 2006). Their relatively small extent reduces the global significance of subtropical forests
for carbon sequestration, but as pointed out above, some have both a high biomass and a high
sink capacity on a per-area basis.
Simple models of future vegetation distribution under anthropogenic climate change
suggest that subtropical forests will shift polewards as the temperature rises. Even in the absence
of the now ubiquitous forest fragmentation, however, tree species migration rates would
probably be too slow to track temperature change (Corlett & Westcott, 2013). Real-world
responses are likely to be more complex, reflecting both the complexities of climate change and
the many additional anthropogenic impacts that these forests are subject to. Long-term
monitoring of large, permanent, sample plots is our best hope of understanding these changes.
Recent observation of rapid structural and compositional changes in subtropical forests in both
China and Argentina have been interpreted as reflecting regional drying associated with warming
in the former case (Zhou et al., 2014) and the possible impacts of CO2 fertilization in the latter
(Malizia et al., 2013).

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Figure 1. Potential forest cover between 23.4o and 30.0o, under the assumption that all areas
with >700 mm annual rainfall and <4000 m elevation are capable of supporting forest. This
map was produced in ArcView 10.1 using climate data from WorldClim.