Niches and Areas

of
Distribution I
Jorge Soberon
Museum of Natural History, University of Kansas
hoto by !arren A"el

Caveat Emptor
#
$his is a tal% about ideas being develo"ed as &e s"ea%'
#
Nothing is yet firmly established
#
!e encourage dissent and (uestioning
#
Key "a"ers)
*
ulliam, +,,,, -col' .ett' /)/012/34
*
earson 5 Da&son, +,,/, 6lob' -col' 7iogeog' 4+)/342/84
*
Arau9o 5 6uisan, +,,:, 6lob' -col' 7iogeog' //)4388243;;
*
Kearny +,,3, <i%os 44:) 4;32414
*
Soberon 5 eterson, +,,:, 7iodiv' Info' +)424,
*
Soberon +,,8, -col' .ett' 4,)444:244+/

ur"ose) to clarify very muddled
terminology
#
!e need conce"ts of
=area of distribution>
of a s"ecies &hich are
clear and o"erational'
#
$he same for niche'
#
?elate the conce"ts of
=areas of distribution>
and =niche>'

Around 411, three things ha""ened
4' .arge databases of
"resences of s"ecies
@mainly
com"uteriAed
scientific collectionsB
began being
accessible at
significant amounts

II' 6ISC
#
6eogra"hical
Information Systems
technology became
&idely accessible to
ecologists and
biogeogra"hers

III' =Niche> Modeling
#
Since the 41;,s, the
Australians began doing
=bioclimatic> modeling
for a""lied entomology
"ur"oses @DlimeEB
#
.ater they develo"ed
7ioclim, D<MAIN, and
then, 6A?
#
$oday, there are about 4:
methods to do niche
modeling, most of them in
free&are form

Using those toolsC
#
Since about 411: there has been an eE"losion in
the a""lication of ecological =niche> modeling
methods to estimate =areas of distribution>,
"otential or actual'
#
$he methods are used to "redict invasive2s"ecies
routes, s"read of vectors of diseases, restoration
ecology, conservation, and many to"ics in
fundamental ecology and biogeogra"hy'
#
.iterally hundreds of "a"ers have been "ublished,
about the "otential andFor actual distributions of
thousands of s"ecies

And yetC
#
=6eogra"hic distributional areas are the shado&s
"roduced by taEa on the geogra"hical screen' $o
study them one needs to measure ghosts>
*
A' ?a"o"ort' Areogra"hy) 6eogra"hical Strategies of
S"ecies, 41;+
#
=Most ecologists &ould agree that niche is a central
conce"t of ecology, even though &e do not %no&
eEactly &hat it means>
*
.' ?eal and S' .evin, $heoretical advances) the role of
theory in the rise of modern ecology
4114

In other &ords)
#
!e @the niche
modelersB are doing
&holesale estimation
of those shado&s
@geogra"hical areasB
by using a non2
conce"t @nicheBG
#
Not niceC

!e need to define -NM terms and
methodology
#
referably operational definitions
#
?elevant in the field of modeling
distribution areas
#
referably mathematical @formalB
#
Avoid brea%ing &ith tradition, if "ossible

.et us begin &ith area'
#
In boo%s they a""ear
as little ma"s
#
Some theorists even
treat them as if they
&ere "hysical entities'
-2 -1 0 1 2
-2
-1
0
1
2
7ut areas are com"osed
by the "resence @detectedB
of individuals
Jim 7ro&n suggested a 6edan%eneE"eriment

Imagine that &e ma%e individuals fluoresce'''
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2

-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2
-2 -1 0 1 2
-2
-1
0
1
2

!e see that to define =area of
distribution> &e need to)
#
Agree on &hat it is meant
by a ="resence> @sin% or
source "o"ulationsB'
otential or actual'
#
Agree at &hat resolutions
ma%es sense to tal% about
areas of distributionH
Normally, this &ould be
=lo& resolutions> I 4,
/
%m
+
#
Inside the cells, &e are in
the domain of ecology,
outside, it is biogeogra"hy
or macroecology

So areas are'''
#
?egions of the &orld defined by the detection of
individuals &ithin cells in grids
#
$here are several things that &e can detect)
re"roductive individuals, 9ust individuals, regions
&ith "ositive gro&th rates'''
#
Although a lot of those can be defined at high
resolutions, areas are defined at lo& resolutions'
#
It is still a dream to measure the movements of all
the individuals of a s"ecies'''

!hat determines &here do &e
find individualsH
Autoecology, interactions, migration "atterns, historical factors,
o"erating &ith different strenghts at different s"atiotem"oral scales'
$here are e(uations @horribly difficultB describing this'
4
@ B @ J B @ J B
j
j j j j j
i
i i i
j
i
dx
r e x R x
x dt
= + T
r
r r r
S"ecies
6rid cell

@ B
j
i
r e
r
@ J B
j j j
i i
x R
r
r
@ J B
j
x T
r
4
@ B @ J B @ J B
j
j j j j j
i
i i i
j
i
dx
r e x R x
x dt
= + T
r
r r r
6rinnell,
sceno"oetic
-lton, bionomic
Movements
G
7AM Diagram
7
M
A

So &e can define stuffC
#
!e can define,
rigorously, areas of
distributions in terms
of &hether the x
i
j
(t) are
"ositive, or above a
threshold, or &hether
the intrinsic gro&th
rate is "ositive
@"otential area of
distributionB
,
K L ,M
j
j
C
x
dx
j
dt

= > = J G A B I
K L @ B ,M
F j
j r e = > = J G A
r
,
, 4
K L ,
and @ B ,, M
j
j
O
x
j
dx
j
dt
x t t t t

= >
>
=
J
A B M I I

And nichesH
#
Dlearly, the s"atial
regions determined by
the "arameters in those
horrible e(uations
have environments'
#
Niches may be
defined using
the environmental
variables in the
=occu"ied> cells'

A useful and forgotten distinction
of Hutchinson @418;B
#
Sceno"oetic variables'
Non2interacting, slo&ly
changing from a s"ecies
"oint of vie&'
#
7ionomic variables'
Dou"led, fast changing
#
HutchinsonNs nice
distinction &as (uic%ly
forgotten and then
reinvented by Austin,
7egon, and others'

Niches can be defined using the
same e(uations that govern area
4
@ B @ J B @ J B
j
j j j j j
i
i i i
j
i
dx
r e x x
x dt
= + R T
r r r
Oundamental niche
Intrinsic 6ro&th ?ate
@Sceno"oeticB
?esource2consumer
dynamics, com"etitors,
"redator2"rey @bionomicB'
Migration
coloniAation, history

So, the niches are the sets of
environments "resent in the different
areas)
#
An abiotic niche) the set
of environments in the
area &here the intrinsic
gro&th rate is "ositive
#
A colonizable niche) the
set of environments in the
area &here both biotic and
abiotic conditions are right
#
A occupied niche, the set
of the environments in the
area &here the s"ecies
actually occurs
@ B
O O
= J N
@ B
C C
= J N
@ B
F F
= J N

HutchinsonNs terminology
1. Fundamental niche'
Defined &ithout
reference to sceno"etic
and biotic variables'
2. Realized niche'
Oundamental reduced
by com"etition' $he
entire bioto"e occu"ied
1. Abiotic niche' Defined using
only sceno"oetic variables'
2. Colonizable niche' $he
abiotic reduced by
com"etition'
3. Occupied niche' $he
realiAed niche ta%ing into
account movement
restrictions

Some areas and niches
7
M
J
<
J
<
7
M
J
<
A
J
<
O C
= N N
<ccu"ied P coloniAableH
@ B
F
= A N
4
@ B
A A

= = N A J
Abiotic
N
@ N B
@ B @ N B
O O C
O O
O O
O O C

=
=
=
=
J J J
J J
J J
N N N
U
U
U
U
DoloniAable
and
occu"ied

$he -ltonian Noise Hy"othesis I'
#
At the scale at &hat
&e &or% @cells I 4,
,

%m
+
B, the biotic factors
become much less
relevant than the
abiotic ones) the signal
is dominated by
abiotic factors, and the
biotic factors act as
noise
7
M
J
<
A
J
<

-ltonian noise hy"othesis II'
#
$he -ltonian "rocesses
are of very high
resolution
#
In the large cells at
&hich sceno"oetic
variables are measured
=com"etitive eEclusion>
may ta%e "lace &ithout
affecting the totality of
"o"ulations'
A
7
M

-Eam"les and countereEam"les
#
Cactoblastis cactorum
#
<bligate "ollinators

-Etremely Im"ortant I'
#
7y assuming non2interactivity &e can re"resent
the environments @and therefore the 6rinnell
nichesB by !et! o" numbe#!'
#
$he interactive and dynamic -lton niches must be
re"resented by the "arameters of functions
@sometimes by the isoclinesB, because one needs to
measure both the use of a resource and its im"act
@see Dhase and .eibold, +,,/, Ecological NicesB

-Etremely Im"ortant II'
#
Since &e are defining our 6rinnell niches
using such =sceno"oetic> variables, it
means that &e can use)
*
!idely available datasets
*
At lo& resolutions

$o&ards concludingC
#
So it is useful to define areas as subsets of
cells in a geogra"hic grid, defined by
biological "ro"erties of s"ecies @"resence,
"resence of viable "o"ulations, suitable
conditions for survivorshi",CB
#
And 6rinnell niches as the subsets of
sceno"oetic variables associated to those
areas

And of course, 6rinnell niches are
multidimensional, but in the s"ace of
sceno"oetic variables
#
=$he conce"t of
niche'''is here defined
as the sum of all the
environmental factors
acting on an organismJ
the niche thus defined
is a region of an n2
dimensional
hy"ers"ace>

Multidimensionality a""lies to
both scales of niches
#
In 6rinnellNs @related
to distribution,
geogra"hicB, variables
are elevation,
orientation, geology,
climate'''
#
$hese variables are
=less interactive>
#
In -ltonNs @more
tro"hic, ecologicalB,
variables may be
nutrient concentrations,
food siAe distributions,
"rey, "redators,
com"etitors and
mutualist densities'''
#
$hese variables are
highly interactive

So, it is useful to acce"t to& levels
#
6rinnell Niches, lo&
resolution, sceno"oetic
variables'
#
-lton Niches, high
resolution, bionomic
variables' $hey may
be the cause of
reduction from
Oundamental to
?ealiAed
A
M
7

$his distinction is based on)
#
Acce"ting the assum"tion that differences in
certain eEtreme ty"es of variables) non2interactive,
meaningful at lo& resolutions, vs interactive,
meaningful at high resolutions do eEist'
#
Ho& valid is this sim"lificationH
#
$he fact that -NM is "redictive is evidence in
favor'
#
!e need to establish the sco"e of validity of the
assum"tion' At high resolutions "robably it &ill
brea% do&n'

$han%s to
#
$o&n eterson, ?ichard earson, -nri(ue
MartineA, Miguel Na%amura, Darlos
MartineA del ?io, the students of the Niche
Seminar

Niches and Areas
of
Distribution II
Jorge Soberon
Museum of Natural History, University of Kansas
hoto by !arren A"el

So ho& &e measure a 6rinnell
NicheH
4' !e go and see &here it is that a s"ecies occurs
@=C&ithout immigration>B
+' !e measure environmental variables in those
"laces'
/' !e eEtra"olate from a fe& ="oints> to an area'
0' !e obtain the corres"onding environmental
values'
:' $his is the niche'

$here are Several !ays of Dalculating
the Niche
hysical variable 4

h
y
s
i
c
a
l

v
a
r
i
a
b
l
e


+
7ioclim
Dlusters
About 4: others'''

5
10
25
50
75
100
125
150
175
200
225
250
275
above
Bivariate Histogram
62s"ace' $&o dimensions
.o& resolution @"iEels
of 4 %m
+
B
-2s"ace' Many variables'
Sceno"oetic' .o& resolution'
.o& "recision

Notice)
Donce"tually, every algorithm tries to =recover> the realiAed 6rinnell
niche, since the data "oints come from the actual distribution J
<
4' Ho&ever, &hen &e ma" bac% from $ to G, in general, &e do not
recover J
<
since there may be other s"atial regions &ith the same
niche' !e recover J
D
' In general, one has to deal &ith this
"roblem outside the niche modeling algorithm
J
<
N
<
J
<

# S# S
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!amandua mexicana

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# S # S# S
# S# S

!hen
24
@N
<
B is different from J
<
H
J
4
J
+
$
1
62s"ace -2s"ace

Using 6ar", MaEent, 6AM, C
estimate the <ccu"ied Niche
If you &ant the Jo @rather than J
D
B, then cli" the
geogra"hical eEtent of the $
?
using biological
connectivity or historical reasoning'
!ith 6IS dis"lay the
geogra"hical eEtent
of the <ccu"ied
Niche
7egin &ith "resence @absenceB data

A real2life eEam"le) "aronia brevicornis and
its single food "lant, Acacia cocliacanta

Oor this eEam"le the follo&ing layers
&ere used, all at 4)4,,,,,,,,)
#
As"ect
#
-levation
#
Mean $em"erature
#
Average minimum
daily tem"erature
#
Average maEimum
daily tem"erature
#
Qearly "reci"itation
MM- about :,, meters

"# brevicornis Abiotic Niche using 7S 6ar"

II) -stimating the =Area of Accesibility>
#
Orom &hereH !hat is the initial conditionH
#
At &hat scaleH In relation to &hat vagility
"arametersH
#
At certain scales, one can assume that
biogeogra"hy is a good surrogate for the
accesibility areas, this is, &e assume that if
a s"ecies is "resent in a given
biogeogra"hical region, it can reach all of it'

"# brevicornis 7iogeogra"hical rovinces

"# brevicornis# 7iogeogra"hical rovinces
as Surrogates for Accesibility Areas

"# "revicornis Niche R Accesibility
%

III' -stimating an <bligate Interaction
#
"# brevicornis only food "lant is the legume
Acacia cocliacanta
#
A similar analysis is the re"eated for A#
cocliacanta

A# cocliacanta Abiotic Niche

A# cocliacanta accesibility region @biogeogra"hyB

A# cocliacanta niche R accesibility

"# brevicornis @redB R A# cocliacanta @blueB


Oinally, is it the occu"ied or the
abiotic nicheH
7
M
J
<
A
@ B @ B
O
A O O


= =
A J
A N N J
B
M
J
<
A
@ B @ B
O
A O O

= =
A J
A N N J
;
;

Donclusion' !e still have a lot to
learn, for eEam"le about)
#
$he "rocess of =reduction> from Abiotic to
<ccu"ied, by -ltonian factors' $his is a "roblem
in scaling
#
$he basic assum"tion of using -NM to study
invasions is that N
<
P N
D'
Is this trueH !henH
#
A more standardiAed "rocedure to obtain different
Areas and 6rinnellian Niches using -NM
#
Develo"ing validation techni(ues based on
different configurations of the 7AM diagram
#
And many other thingsC

Ac%no&ledgments
#
roviders of museum data, for the huge
effort they have "ut in collecting, curating
and identifying s"ecimens, and then sharing
their results
#
$he o&ners of the land &here those
s"ecimens &ere originally ta%en

Sery Oundamental Donce"ts)
#
$o each region in
62s"ace there are
corres"onding
ones in -2s"ace
@6B P -
#
$here is an
inverse o"eration
that ma"s from 6
to -)
1
@-B P 6
-2S"ace
n dimensions
62S"ace,
+ dimensions

More Oundamental Donce"ts'
#
Niches @6rinnellB are subsets of
s"aces of environmental
variables corres"onding to
subsets of geogra"hical s"ace
&here a s"ecies live, or can
live'
#
$he function that ma"s from
G to $ is defined very sim"ly
by the tables in the 6IS that
lin% coordinates &ith
environmental values'
#
And viceversa, for the relation

24
that ma"s bac% $ to G

$edan%enexperiment I) !e %no& all regions &here
intrinsic gro&th rate is "ositive @this is, &e disregard
biotic interactions and movement limitationsB G
O
!e agree to call the corres"onding
-nvironmental subset the Oundamental Niche, $
O
P @G
O
B
It ought to be true that
24
@$
O
BP
24
@@G
O
BBPG
O

G
O

$edan%enexperiment II) !e %no& no& the area of
distribution of a s"ecies G
?
!e agree to call the corres"onding
-nvironmental subset the ?ealiAed Niche, $
?
P @G
?
B
No& it is obvious that
24
@$
?
B P may not be P G
?

G
?

Bio1
100 140 180 220 20 40 60 80 120
1
8
0
2
2
0
2
6
0
1
0
0
1
4
0
1
8
0
2
2
0
TempRng
AnnPrecip
1
0
0
0
4
0
0
0
7
0
0
0
180 220 260
2
0
6
0
1
0
0
1000 4000 7000
Coe!arPrecip
!amandua mexicana

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# S# S

!hen
24
@$
?
B is different from G
?
H
G
4
G
+
$
1
62s"ace -2s"ace