Quickstart guide

In this tutorial we will assume that you have imported matpopmod as

import matpopmod as mpm

Defining a model

The central object of matpopmod is the MPM, an abstract representation of a matrix population model that stores all the information about that model. You can create a MPM by specifying its projection matrix A:

>>> m1 = mpm.MPM(A = [[0.1, 2], [0.4, 0.8]])
>>> m1
  A = [[0.1, 2. ],
       [0.4, 0.8]],
  metadata = {}

However, whenever possible it is preferable to specify a matrix of survival probabilities S and a matrix of fertilities F. This will give you access to much more information about the model.

>>> m2 = mpm.MPM(S = [[0.1, 0], [0.4, 0.8]], F = [[0, 2], [0, 0]])
>>> m2
  S = [[0.1, 0. ],
       [0.4, 0.8]],
  F = [[0, 2],
       [0, 0]],
  metadata = {}

To avoid having to type matrices twice, it is also possible to specify A along with the list of its entries that correspond to fertilities. This has the same effect as specifying S and F.

mpm.MPM(A = [[0.1, 2], [0.4, 0.8]], fertilities = [(0, 1)])

If you find that specifying matrices using lists of lists is not convenient, note that a variety of formats are available (lists of lists, strings, NumPy arrays, external files…). In fact, much of the documentation of matpopmod uses strings, which are easier to type than lists of lists:

mpm.MPM(S = "0.1 0; 0.4 0.8", F = "0 2; 0 0")

See the documentation of MPM for details.

Computing descriptors

Computing the descriptors of the models we have just defined is straightforward:

>>> m2.lmbd # asymptotic growth rate
>>> m2.w # stable stage distribution
array([0.60414407, 0.39585593])
>>> m2.v # reproductive values
array([0.52602896, 1.72336113])
>>> m2.elasticities
array([[0.02253133, 0.29526595],
       [0.29526595, 0.38693677]])
>>> m2.R0 # net reproductive rate
>>> m2.T_R0 # R0 generation time
>>> m2.T_a # average age of mothers

Matpopmod gives you access to more than 70 descriptors of matrix population models. If you want to know something about a model, chances are that the answer is only one “dot” away. See the complete list of available descriptors, which contains detailed explanations about how each descriptor is defined and computed, along with precise references.

As mentioned above, not all descriptors can be computed for every model. For instance, some descriptors require knowing the survival and fertility matrices. If it was not provided, a NotAvailable exception will be raised:

>>> m1 = mpm.MPM(A = [[0.1, 2], [0.4, 0.8]])
>>> m1.R0
Traceback (most recent call last):
matpopmod.utils.NotAvailable: A = S + F decomposition required.

In addition to this, some descriptors may be undefined because of the properties of the projection matrix. For instance, the following projection matrix has no stable distribution due to periodicity:

\[\begin{split}\mathbf{A} = \begin{pmatrix} 0 & 2 \\ 0.5 & 0 \end{pmatrix}\end{split}\]

When trying to access a descriptor that is mathematically undefined, a detailed warning message will be printed and the value nan (“not a number”) will be returned.

>>> m3 = mpm.MPM(A = "0 2; 0.5 0")
>>> m3.w
UserWarning: A is not quasi-primitive. Most descriptors
are ill-defined. They will be set to NaN.
array([nan, nan])

Note that you do not have to worry about keeping track of the descriptors: they are automatically stored in the model.

Finally, and importantly, it is not possible to modify the descriptors of a model. Trying to do so will raise an error:

>>> m3.lmbd = 0
Traceback (most recent call last):
AttributeError: can't set attribute

This behavior is intended: it is meant to help you catch errors in your code.

The only exception to this rule is the metadata attribute: this is a dictionary that you can use however you wish to store contextual information about the model.

>>> m3.metadata["Comments"] = "Something relevant"

Trying ready-made models

The module examples provides ready-made models that can be used to get familiar with the library. The complete list of examples is

>>> mpm.examples.all_models
(this prints a lot of things)

If you want to work with a specific model, you can do so as follows:

>>> orcas = mpm.examples.orcinus_orca
>>> orcas
  S = [[0.    , 0.    , 0.    , 0.    ],
       [0.9775, 0.9111, 0.    , 0.    ],
       [0.    , 0.0736, 0.9534, 0.    ],
       [0.    , 0.    , 0.0452, 0.9804]],
  F = [[0.    , 0.0043, 0.1132, 0.    ],
       [0.    , 0.    , 0.    , 0.    ],
       [0.    , 0.    , 0.    , 0.    ],
       [0.    , 0.    , 0.    , 0.    ]],
  metadata = {
    'ModelName': 'orcinus_orca',
    'Species': 'Orcinus orca',
    'CommonName': 'Killer whale',
    'Classes': ['Newborns', 'Juveniles', 'Reproductive adults',
'Post-reproductive adults'],
    'Source': 'Example 5.1 from Caswell H. (2000). Matrix
Population Models: Construction, Analysis, and Interpretation.',
    'Comments': 'The projection matrix is not irreducible due
to the presence of a post-reproductive class.\n\nExample of a
K-strategy, with high survival and low fertility.'

As we can see here, those models come with predefined metadata. In particular, the Comments field gives a brief description of the specificities of the model:

>>> print(orcas.metadata["Comments"])
The projection matrix is not irreducible due to the presence
of a post-reproductive class.

Example of a K-strategy, with high survival and low fertility.

The documentation of the module examples contains a detailed description of each model, along with a brief discussion of its specificities, usually with code samples. We recommend reading it after finishing this quickstart tutorial.

Plotting trajectories

The module plot can be used to plot a variety of things, such as life-cycle graphs, eigenvalues or matrices.

Here we are going to focus on plotting trajectories. First, we need to create a Trajectory:

>>> dipsacus = mpm.examples.dipsacus_sylvestris
>>> n0 = [0, 0, 0, 0, 0, 1] # initial population vector
>>> traj = dipsacus.trajectory(n0, t_max=20)

To plot the total population size \(n(t)\) as a function of time, we simply use:

>>> mpm.plot.trajectory(traj)
<AxesSubplot:xlabel='time $t$', ylabel='Population size'>

This returns a matplotlib Axes that can be saved to an external file using

>>>  mpm.plot.savefig("figure.png")

or displayed interactively by calling

>>>  mpm.plot.show()

(those functions are aliases of the corresponding matplotlib.pyplot functions; see the documentation of matplotlib for details). To avoid having to run mpm.plot.show() every time you plot a figure, you can turn matplotlib’s interactive mode on with

>>>  import matplotlib.pyplot as plt
>>>  plt.ion()

After displaying the figure, you should see this:


As expected, the population growths exponentially and very rapidly \((\lambda \approx 2.33)\). As a result, this plot is not very informative; instead, it would be more instructive to plot a function of the population size. For instance, since we know that \(n(t) \sim \mathbf{v} \mathbf{n}(0) \lambda^t\), we could plot \(n(t) - \mathbf{v} \mathbf{n}(0) \lambda^t\) instead of \(n(t)\). This would enable us to see the oscillations of the second-order dynamics. The amplitude of these oscillations increases/decreases like \(\tau^t\), where \(\tau\) is the second-largest modulus of the eigenvalues, and here \(\tau \approx 1.77\) so it will grow rapidly. Thus, it would make sense to use a log scale. All of this can be done as follows:

>>> mpm.plot.trajectory(traj, second_order=True, log=True)

Of course, other transformations of the population size are possible. For instance, we could have plotted \(n(t) / \lambda^t\) by using the argument rescale instead of second_order, and \((n(t) - \mathbf{v} \mathbf{n}(0) \lambda^t) / \tau^t\) by using both.

So far, we have focused on the total population size. But we can also plot the abundance of each class thanks to the argument show_classes. For the sake of the example, we will also use show_period to add vertical lines at multiples of the period of the second-order oscillations.

>>> mpm.plot.trajectory(
...     traj,
...     show_classes = True,
...     second_order = True,
...     rescale = True,
...     show_period = True)

A useful way to plot the abundances of the classes while making the total population size and the relative abundances of the classes easy to read is to use a stacked plot. Of course, this can only be used with a linear scale and with positive quantities.

>>> mpm.plot.trajectory(
...     traj,
...     show_classes = True,
...     stacked = True,
...     rescale = True,
...     show_period = True)

This nicely illustrates the convergence of the population structure.

Finally, to conclude this brief overview of the function plot.trajectory, here is another stacked plot of the same trajectory, where we (1) only plot a subset of classes; (2) add a legend to the figure; and (3) set the plotting window manually.

>>> ax = mpm.plot.trajectory(
...     traj,
...     show_classes = [2, 3, 4, 5],    # (1)
...     stacked = True,
...     show_legend = "upper left")     # (2)
>>> ax.set(xlim=(0, 6), ylim=(0, 1200)) # (3)

Plotting multiple trajectories on the same graph is also possible, and is done with the function plot.multiple_trajectories. Besides comparing deterministic trajectories corresponding to different initial conditions / models, this is useful to show different realizations of a stochastic process. The following code generates 1000 independent replicates of a population of Orcinus orca with demographic stochasticity, and overlays them with their expected value (i.e. with the deterministic trajectory of the corresponding matrix population model).

>>> orcas = mpm.examples.orcinus_orca
>>> trajs = orcas.stochastic_trajectories(
...     n0 = [10, 0, 0, 0],
...     t_max = 100,
...     reps = 1000,
...     reproduction = "bernoulli")
>>> mpm.plot.multiple_trajectories(trajs)

See the documentation of stochastic_trajectories() for the precise assumptions of the stochastic model, and of plot.multiple_trajectories for the various plotting options (which, in addition to rescaling, etc, include showing the empirical standard deviation and quantiles). Combining these can give aesthetically interesting plots, as the following trajectories of a stochastic population of Dipsacus sylvestris show.

Loading models from COM[P]ADRE

The COMPADRE and COMADRE databases contain more than 12000 matrix population models that have been curated from the literature. The databases are distributed as RData files that must be converted to JSON before they can be used with matpopmod. If you do not have a suitable version of R installed, you can use the following pre-converted versions of the databases:


converted on 18/11/2021


converted on 18/11/2021

Those files can then be loaded in Python using the function load():

>>> db = mpm.compadre.load("COMPADRE_v.")
>>> db
MPM Collection (7907 models) at 0x7fe0bfc2be10

If you have a working installation of R, you can also use fetch() to download, convert and load any version of the database:

# Fetch the latest version
latest = mpm.compadre.fetch("compadre")

# Fetch a specific version
older = mpm.compadre.fetch("compadre", "")

Loading a database yields a MPMCollection. MPM collections are essentially lists of MPMs with a custom interface. General information that corresponds to the fields of the COMPADRE/COMADRE variable version is stored in the info attribute of the collection:

>>> db.info
{'filename': 'COMPADRE_v.',
 'Database': 'COMPADRE',
 'Version': '',
 'Type': 'Release',
 'DateCreated': 'Aug_20_2021',
 'TimeCreated': '19:00',
 'Agreement': 'https://www.compadre-db.org/UserAgreement',
 'GeneratorScriptVersion': '1.3'}

Models that could not be loaded (e.g, because of NA entries or of survival probabilities greater than 1) are kept in invalid_models, where they are represented by InvalidMPM objects. Invalid MPMs cannot be used to compute anything, but they have the same matrix and metadata attributes as regular MPMs. In addition, they have an attribute error explaining why they are not valid models.

>>> db.invalid_models[0].error
InadequateMatrix('Matrix S has a column whose sum is > 1.')

Use db[i] to access the i-th model of a collection, and for m in db to iterate over the models of the collection (only valid models are considered). For instance, the following line of code shows that about half of the models in COMPADRE correspond to decreasing populations:

>>> numpy.median([m.lmbd for m in db])

Models from COMPADRE/COMADRE have unique IDs known as their MatrixID. To access a model from its MatrixID, use

>>> db.get_from_id(239145)
  S = [[0.   , 0.   , 0.   , 0.   ],
       [0.   , 0.   , 0.045, 0.   ],
       [0.125, 0.   , 0.091, 0.   ],
       [0.125, 0.   , 0.091, 0.333]],
  F = [[0.   , 0.   , 0.245, 2.1  ],
       [0.   , 0.   , 0.   , 0.   ],
       [0.   , 0.   , 0.   , 0.   ],
       [0.   , 0.   , 0.   , 0.   ]],
  metadata = {
    'MatrixID': 239145,
    'SpeciesAuthor': 'Anthyllis_vulneraria',
    'SpeciesAccepted': 'Anthyllis vulneraria',
    'CommonName': 'Kidney vetch',
    'Kingdom': 'Plantae',
    'Phylum': 'Magnoliophyta',
    'Class': 'Magnoliopsida',
    'Order': 'Fabales',
    'Family': 'Leguminosae',
    'Genus': 'Anthyllis',
    'Species': 'vulneraria',
    'OrganismType': 'Herbaceous perennial',
    'DicotMonoc': 'Eudicot',
    'AngioGymno': 'Angiosperm',
    'Authors': 'Davison; Jacquemyn; Adriaens; Honnay; de Kroon; Tuljapurkar',
    'Journal': 'J Ecol',
    'SourceType': 'Journal Article',
    'YearPublication': '2010',
    'DOI_ISBN': '10.1111/j.1365-2745.2009.01611.x',
    'AdditionalSource': 'Piessens Oecologia 2009',
    'StudyDuration': '4',
    'StudyStart': '2003',
    'StudyEnd': '2006',
    'ProjectionInterval': '1',
    'MatrixCriteriaSize': 'Yes',
    'MatrixCriteriaOntogeny': 'No',
    'MatrixCriteriaAge': 'No',
    'MatrixPopulation': 'G',
    'NumberPopulations': 9,
    'Lat': 50.0914,
    'Lon': 4.5919,
    'Altitude': 160,
    'Country': 'BEL',
    'Continent': 'Europe',
    'Ecoregion': 'TBM',
    'StudiedSex': 'A',
    'MatrixComposite': 'Mean',
    'MatrixSeasonal': 'No',
    'MatrixTreatment': 'Unmanipulated',
    'MatrixCaptivity': 'W',
    'MatrixStartYear': '2003',
    'MatrixStartSeason': 'Summer',
    'MatrixStartMonth': '8',
    'MatrixEndYear': '2004',
    'MatrixEndSeason': 'Autumn',
    'MatrixEndMonth': '9',
    'MatrixSplit': 'Divided',
    'MatrixFec': 'Yes',
    'Observations': 'The GPS coordinates were approximated to the closest geographic location described in the reference.',
    'MatrixDimension': 4,
    'SurvivalIssue': 0.333,
    '_Database': 'COMPADRE',
    '_PopulationStatus': 'Released',
    '_PublicationStatus': 'Released',
    'MatrixClassAuthor': ['seedling', 'vegetative adults', 'small (1-2 flowering stalks)', 'large (>3 flowering stalks)'],
    'MatrixClassOrganized': ['active', 'active', 'active', 'active'],
    'MatrixClassNumber': [1, 2, 3, 4]

As we can see from this last example, the metadata of a COMPADRE/COMADRE entry are stored in the metadata dictionary of the corresponding MPM. For instance, the SpeciesAccepted field of a model m is obtained with


As an elementary example of how the databases can be used, the following code snippet will print the 5 most common species in COMADRE:

comadre = mpm.compadre.fetch("comadre")
species = [m.metadata["SpeciesAccepted"] for m in comadre]
from collections import Counter
for s, k in Counter(species).most_common(5):
    print(f"{s}: {k} projection matrices")

The result is perhaps unsurprising:

Homo sapiens sapiens: 1120 projection matrices
Gorilla beringei beringei: 84 projection matrices
Vireo atricapilla: 56 projection matrices
Brachyteles hypoxanthus: 52 projection matrices
Orcinus orca: 47 projection matrices

For a more complete presentation of the tools provided to work with COMPADRE/COMADRE – which include filtering and saving MPM collections – see the documentation of the module compadre.

Individual-based simulations

(coming in version 0.2.0)

Advanced use

Collapsing matrices

Collapsing a projection matrix consists in merging some classes in order to obtain a smaller projection matrix. This can be useful, e.g, to get models with similar classes that can be compared; or to test how much information is lost when adopting a coarser description of a model. The module collapsing implements the tools needed to do this.

Generating random MPMs

There are several reasons why generating random MPMs with prescribed properties can be useful. Maybe you have observed an empirical relationship between some descriptors and would like to test it a bit more thoroughly; maybe you have proved something for Leslie models, and would like to know whether that result also holds for other types of models; or maybe you would just like to do some code testing. The module random gives you a way to do just that.

(coming in version 0.2.0)

Studying kinship

The module kinship implements the calculation of the kinship matrices, as described in [CBRR21]. The (ij)-th entry of the kinship matrix \(\mathbf{K}(g, q)\) is the expected number of (gq)-kin of a focal individual (in a monoparental population, two individuals A and B are said to be (gq)-kin if the least common ancestor of A and B is the g-th ancestor of A and the q-th ancestor of B).

(coming in version 0.2.0)

Custom calculations

Users who would like to write their own mathematical formulas are invited to have a look at the module mathtools, which contains useful mathematical functions.

Note that some of these functions are also implemented in the SciPy library, which we recommend using in addition to NumPy for slightly more advanced linear algebra.