Variation in plant demographic trajectories: problems and opportunities
-
Upload
owen-jones -
Category
Education
-
view
242 -
download
3
description
Transcript of Variation in plant demographic trajectories: problems and opportunities
Variation in plant demographic trajectories:
problems and opportunities
Owen R. Jones & Roberto Salguero-Gó[email protected]
Laboratory for Evolutionary Demography,Max Planck Institute for Demographic Research, Rostock, Germany
Evolution is a demographic process
“Evolution is driven by the propagation of genes, traits and individuals with and between populations. This propagation depends on the survival, fertility and dispersal at each age or stage during their life history.”
Metcalf & Pavard (2007) TREE
Mor
talit
y/fe
rtili
ty r
ate
Surv
ivor
ship
(lx
)
Classical theories of the evolution of senescence
Hamilton (1966)Williams (1957) Kirkwood (1977)Medawar (1952)
Older ages are less important to evolution than younger ages: - force of selection declines with age.
Classical theories of the evolution of senescence
“No life schedule ... could escape my spectrum of forces of senescence … in the farthest reaches of almost any bizarre universe” (Hamilton, 1966)
Mor
talit
y
AgeRe
prod
uctio
nAge
COMPADRE DatabaseCOMParative Plant & Algae Demographic REsearch
Trees
Palms
Succulents
Lianas
Shrubs
EpiphytesHerbsAlgae
Miguel Franco
Jonathan Silvertown
COMPADRE DatabaseCOMParative Plant & Algae Demographic REsearch
Trees
Palms
Succulents
Lianas
Shrubs
EpiphytesHerbsAlgae
Miguel Franco
Jonathan Silvertown
Roberto Salguero-Gómez
33"1 " " """""""44" " """""""""""""""""88 " " """""""""""""132"
COMPADRE DatabaseCOMParative Plant & Algae Demographic REsearch
Trees
Palms
Succulents
Lianas
Shrubs
EpiphytesHerbsAlgae
817 species
Number of species
From matrices to trajectories
lxmx
Age
Age
Caswell (2001); Cochran and Ellner (1992)
Mortality trajectories in plants
Acknowledgements
Discussion
Life and death in the garden: mortality and recruitment trajectories in plantsOwen R. Jones* & Roberto Salguero-Gómez*Laboratory of Evolutionary Biodemography, Max Planck Institute for Demographic Research, Rostock, Germany{jones/salguero}@demogr.mpg.de; *Authors contributed equally to this work
One group of 'classical' theories of senescence focuses on the existence of trade-offs between early and late life performance. This body of literature, initiated by Hamilton1 and later supplemented by Williams2, Medawar3 and Kirkwood4, was developed with unitary organisms in mind. These theories are usually interpreted to predict that, after the onset of sexual maturity, survival rate will decline alongside reproductive output. Using data derived from the COMPADRE database5, we examined age-specific mortality and per-capita recruitment trajectories for 329 vascular plant and algae species using population projection matrices6 to examine the validity of these theories for the age trajectories of modular organisms.We demonstrate the existence of an astonishing variety of demographic trajectories in modular organisms, the majority of which do not adhere to the 'classical' Hamiltonian predictions of inevitable senescence. We argue that environmental filtering and phylogenetic ancestry explains much of the observed variation, and that the classical theories of senescence are not sufficient to explain demographic trajectories in modular organisms like vascular plants, red and brown algae.
Summary
Methods
Legend
Force of mortality
Recruitment
Survivorship
Age
Stan
dard
ized
forc
e of
mor
tality
and
recr
uitm
ent
Surv
ivors
hip
We first obtained the average, control population projection matrices for 329 species of vascular plants and algae from the COMPADRE database5. Each of these matrices contains at least three years worth of stage-based information (stage-specific survival, changes in stage, and sexual reproduction) describing the species’ population dynamics. The average, control population projection matrix represents the element-by-element average of all temporally and spatially replicated matrices in each publication but only under “natural” conditions (i.e., experimental treatments such as fires, floods, and droughts were excluded).We employed methods described by Cochran & Ellner7 to obtain age-specific survorship (lx) and per-capita recruitment (mx), as well as mean age at sexual maturity (Lα)6 and lifespan (L)6. The estimates of lifespan excluded seedbank stasis and thus inform only on the performance of photosynthetically active individuals. We then calculated the force of mortality (μx) from lx, and standardized μx and mx by their respective mean values from age zero up to the age of 99% mortality. These standardized trajectories are informative about the distribution of mortality and per-capita recruitment events throughout the life course of each species8. We have sorted the panels from top-left to bottom-right by decreasing standardized force of mortality at old-age. We have color-coded the background of each plot according to the species' growth form (see legend).
Algae Epiphyte Herbaceousperennial Succulent Shrub Liana Palm tree Tree
References
Despite predictions based on Hamiltonian theories of aging, we found no species where there was a simultaneous increase in the force of mortality (μx) together with a decrease in per-capita recruitment (mx) after maturity. A small number of species did show either increasing μx [Plot #: 2, 4, 5, 6, 7, 8, 11, 13, 19, 24, 25, 26, 49, 177] or declining mx values with age [72, 118, 189, 313]. However, the vast majority of species displayed an increase in mx and flattened or decrease in μx with age.Contrary to expectation, the top part of the aging continuum depicted in our plots is not exclusively occupied by short-lived species, nor by species that invest relatively little in tissue construction and maintenance, such as herbaceous perennials. Instead, a number of long-lived trees, palm trees, succulents, and shrubs appear throughout the continuum. The center of the continuum is mostly dominated by species where μx and mx plateau very quickly after sexual maturity. Although a plateau is ultimately expected from the type of stage-to-age transformations shown here6,7, their initial trajectories prior to the plateau are biologically meaningful properties of their populations. For example, the μx of the desert shrub Cassia nemophila [260] appears to gently fluctuate before eventually stabilizing.These trajectories offer a fascinating glimpse into the demography and senescence of modular organisms, which until now have been neglected. The factors shaping these patterns of aging, as well as the conditions under which modular organisms may escape senescence remain to be explored.Note: species which are lacking mx trajectories are either (i) strictly clonally reproductive [10, 13, 65, 103, 161, 213, 229], (ii) did not report recruitment [2], or (iii) are species where <1% of surviving individuals reproduce [306].
1. Hamilton W. 1966. J. Theor. Biol. 12, 12-45. 2. Williams G. C. 1957. Evolution 11, 398-411. 3. Medawar P. B. 1952. An unsolved problem of biology. pp. 44-70. 4. Kirkwood T. B. 1977. Nature 270, 301-304. 5. Salguero-Gómez R. Unpubl. COMPADRE: COMparative Plant & Algae Demographic REsearch Database. MPIDR. 6. Caswell H. 2001. Matrix population models: Construction, analysis and interpretation. 7. Cochran M. E. & Ellner S. 1992. Ecol. Monogr. 62, 345-264. 8. Baudisch A. 2011. Methods in Ecol. & Evol. 2, 375-382.
This work was supported by the Laboratory of Evolutionary Biodemography, and the Max Planck Research Group: Modeling the Evolution of Aging, at the Max Planck Institute for Demographic Research, as well as two fellowships to R.S-G (Teece Family Foundation, University of Pennsylvania). We acknowledge help by the COMPADRE digitalization team: E. Briks, S. Zeh, C. Farack, G. Roemer, A. Henning & G. Hoppe.
Mean age at first reproduction
0.0
2.5
5.0
0 6Years
1Ceratozamia mirandae
(Cycadales)
0 5Years
2Prunus serotina
(Rosales)
0 37Years
3Pinus sylvestris
(Pinales)
0 109Years
4Escontria chiotilla
(Caryophyllales)
0 198Years
5Coccothrinax readii
(Arecales)
0 134Years
6Thrinax radiata
(Arecales)
0 52Years
7Fagus grandifolia
(Fagales)
0 6Years
8Pterygophora
californica(Laminariales)
0 62Years
9Podococcus berteri
(Arecales)
0 16Years
10Disporum smilacinum
(Liliales)
0 128Years
11Geonoma orbignyana
(Arecales)
0 14Years
12Rhododendron maximum
(Ericales)
0 27Years
13Clintonia borealis
(Liliales)
0 458Years
14Prioria copaifera
(Fabales)
0 309Years
15Pinus palustris
(Pinales)
0 40Years
16Callitris
intratropica(Pinales)
0 998Years
17Vouacapoua americana
(Fabales)
0 17Years
18Digitaria eriantha
(Lamiales)
0 8Years
19Carlina vulgaris
(Asterales)
0 672Years
20Dicorynia guianensis
(Fabales)
0 30Years
21Alnus indicata
(Liliales)
0 12Years0
0.5
122
Eriophorumangustifolium
(Poales)
0.0
2.5
5.0
0 15Years
23Anemone patens
(Ranunculales)
0 28Years
24Ulex gallii(Fabales)
0 86Years
25Pinus jeffreyi
(Pinales)
0 11Years
26Carex membranaceae
(Poales)
0 26Years
27Lonicera maackii
(Dipsacales)
0 18Years
28Carex aquatilis
(Poales)
0 37Years
29Aristida bipartita
(Poales)
0 16Years
30Ophrys sphegodes
(Asparagales)
0 8Years
31Centaurea jacea
(Asterales)
0 15Years
32Rosa multiflora
(Rosales)
0 998Years
33Balsamorhiza
sagittata(Asterales)
0 13Years
34Podophyllum peltatum
(Ranunculales)
0 57Years
35Iris germanica(Asparagales)
0 7Years
36Gaura neomexicana
(Myrtales)
0 24Years
37Achillea millefolium
(Asterales)
0 41Years
38Geonoma brevispatha
(Arecales)
0 14Years
39Neotinea ustulata
(Asparagales)
0 671Years
40Callycophyllum
spruceanum(Gentianales)
0 561Years
41Virola surinamensis
(Magnoliales)
0 482Years
42Carapa guianensis
(Sapindales)
0 396Years
43Pouteria sagotiana
(Ericales)
0 378Years0
0.5
144
Platymiscium filipes(Fabales)
0.0
2.5
5.0
0 355Years
45Mora paraensis
(Fabales)
0 136Years
46Arnica angustifolia
(Asterales)
0 271Years
47Licaria mahuba
(Laurales)
0 43Years
48Astragalus scaphoides
(Fabales)
0 14Years
49Cheirolophus
metlesicsii(Asterales)
0 50Years
50Abies sachalinensis
(Pinales)
0 92Years
51Dactylorhiza
lapponica(Asparagales)
0 140Years
52Hymenoxys herbacea
(Asterales)
0 68Years
53Trifolium pratense
(Fabales)
0 61Years
54Pinguicula ionantha
(Lamiales)
0 98Years
55Abies concolor
(Pinales)
0 35Years
56Himantoglossum
hircinum(Asparagales)
0 181Years
57Calochortustiburonensis
(Liliales)
0 706Years
58Taxus brevifolia
(Pinales)
0 475Years
59Heliconia acuminata
(Zingiberales)
0 82Years
60Laccospermasecundiflorum
(Arecales)
0 41Years
61Primula veris
(Ericales)
0 5Years
62Tillandsia
brachycaulos(Poales)
0 14Years
63Heteropogon contortus
(Poales)
0 15Years
64Bothriochloa
insculpta(Poales)
0 8Years
65Aechmea nudicaulis
(Poales)
0 10Years0
0.5
166
Braya fernaldii(Brassicales)
0.0
2.5
5.0
0 59Years
67Banksia ericifolia
(Proteales)
0 139Years
68Petrophile pulchella
(Proteales)
0 3016Years
69Prosopis glandulosa
(Fabales)
0 24Years
70Calochortus albus
(Liliales)
0 147Years
71Chamaedorea radicalis
(Arecales)
0 8Years
72Polemoniumvanbruntiae
(Ericales)
0 13Years
73Saussurea medusa
(Asterales)
0 32Years
74Fritillaria
camtschatcensis(Liliales)
0 25Years
75Actaea elata
(Ranunculales)
0 19Years
76Calluna vulgaris
(Ericales)
0 20Years
77Scabiosa columbaria
(Dipsacales)
0 6Years
78Pseudomisopatesrivas−martinezii
(Malpighiales)
0 32Years
79Fumana procumbens
(Malvales)
0 49Years
80Helianthusdivaricatus(Asterales)
0 27Years
81Viola fimbriatula
(Malpighiales)
0 33Years
82Tillandsia recurvata
(Poales)
0 34Years
83Cimicifuga elata(Ranunculales)
0 16Years
84Centaurea maculosa
(Asterales)
0 20Years
85Setaria incrassata
(Poales)
0 89Years
86Aquilaria crassna
(Malvales)
0 38Years
87Borderea pyrenaica
(Dioscoreales)
0 57Years0
0.5
188
Succisa pratensis(Dipsacales)
0.0
2.5
5.0
0 170Years
89Rosmarinus tomentosus
(Lamiales)
0 121Years
90Cirsium acaule
(Asterales)
0 401Years
91Scaphium borneense
(Malvales)
0 42Years
92Helianthemum juliae
(Malvales)
0 95Years
93Epilobium latifolium
(Myrtales)
0 44Years
94Primula farinosa
(Ericales)
0 34Years
95Calochortusmacrocarpus
(Liliales)
0 88Years
96Geum reptans
(Rosales)
0 125Years
97Eremophila maitlandii
(Lamiales)
0 375Years
98Borderea chouardii
(Dioscoreales)
0 62Years
99Nardostachys
grandiflora(Dipsacales)
0 26Years
100Geranium sylvaticum
(Arecales)
0 33Years
101Rubus saxitalis
(Rosales)
0 196Years
102Calochortusobisopensis
(Liliales)
0 16Years
103Cleistes varicata 1
(Asparagales)
0 84Years
104Periandra
mediterranea(Fabales)
0 99Years
105Guarianthe aurantiaca
(Asparagales)
0 35Years
106Calochortus lyallii
(Liliales)
0 9Years
107Arabis fecunda
(Brassicales)
0 85Years
108Lathyrus vernus
(Fabales)
0 25Years
109Clidemia hirta
(Myrtales)
0 39Years0
0.5
1110
Artemisia genipi(Asterales)
0.0
2.5
5.0
0 7Years
111Sarcocapnosenneaphylla
(Ranunculales)
0 34Years
112Acer saccharum
(Sapindales)
0 44Years
113Ipomoea leptophylla
(Solanales)
0 6Years
114Daucus carota
(Apiales)
0 37Years
115Aconitum
noveboracense(Ranunculales)
0 16Years
116Lomatium cookii
(Apiales)
0 16Years
117Manglietia fordiana
(Magnoliales)
0 7Years
118Hypericum cumulicola
(Malpighiales)
0 8Years
119Iridaea splendens
(Gigartinales)
0 14Years
120Asclepias meadii
(Gentianales)
0 237Years
121Ptychosperma
macarthurii(Arecales)
0 25Years
122Lespedeza cuneata
(Fabales)
0 24Years
123Hydrastis canadensis
(Ranunculales)
0 238Years
124Licania heteromorpha
(Brassicales)
0 5Years
125Cirsium vulgare
(Asterales)
0 9Years
126Atriplex acanthocarpa
(Caryophyllales)
0 399Years
127Eremophila forrestii
(Lamiales)
0 5Years
128Linum catharticum
(Malpighiales)
0 96Years
129Echinaceaangustifolia(Asterales)
0 24Years
130Themeda triandra
(Poales)
0 150Years
131Opuntia macrocentra
(Caryophyllales)
0 7Years0
0.5
1132
Gelidium sesquipedale(Gelidiales)
0.0
2.5
5.0
0 45Years
133Fritillaria meleagris
(Liliales)
0 37Years
134Coryphantarobbinsorum
(Caryophyllales)
0 419Years
135Lepanthes eltoroensis
(Asparagales)
0 45Years
136Aster amellus
(Asterales)
0 18Years
137Allium monanthum
(Liliales)
0 3Years
138Dipsacus sylvestris
(Dipsacales)
0 68Years
139Panax quinquefolium
(Apiales)
0 9Years
140Viola elatior(Malpighiales)
0 12Years
141Braya longii(Brassicales)
0 17Years
142Asarum canadense
(Piperales)
0 4Years
143Tragopogon dubius
(Asterales)
0 14Years
144Lupinus arboreus
(Fabales)
0 6Years
145Carnegiea gigantea
(Caryophyllales)
0 38Years
146Saxifraga cotyledon
(Saxifragales)
0 7Years
147Rumex rupestris(Caryophyllales)
0 6Years
148Taraxacum officinale
(Asterales)
0 21Years
149Chamaecrista keyensis
(Fabales)
0 20Years
150Tolumnia variegata
(Asparagales)
0 8Years
151Centaurea corymbosa
(Asterales)
0 11Years
152Zea diploperennis
(Poales)
0 11Years
153Aquilegia sp.(Ranunculales)
0 10Years0
0.5
1154
Tillandsia punctulata(Poales)
0.0
2.5
5.0
0 653Years
155Sarracenia alata
(Ericales)
0 5Years
156Cirsium dissectum
(Asterales)
0 10Years
157Lupinus tidestromii
(Fabales)
0 9Years
158Allium tricoccum
(Liliales)
0 27Years
159Anthyllis alpicola
(Fabales)
0 33Years
160Echeveria longissima
(Saxifragales)
0 5Years
161Cleistes varicata 2
(Asparagales)
0 16Years
162Viola stagnina(Malpighiales)
0 81Years
163Primula vulgaris
(Ericales)
0 272Years
164Duguetia neglecta
(Magnoliales)
0 11Years
165Cryptantha flava
(Lamiales)
0 32Years
166Aspasia principissa
(Asparagales)
0 9Years
167Bothriochloaischaemum
(Poales)
0 32Years
168Betula nana
(Fagales)
0 10Years
169Tillandsia deppeana
(Poales)
0 15Years
170Hydrangea paniculata
(Cornales)
0 36Years
171Pedicularisfurbishiae
(Malpighiales)
0 33Years
172Cytisus scoparius
(Fabales)
0 82Years
173Iriartea deltoidea
(Asparagales)
0 8Years
174Hypochaeris radicata
(Asterales)
0 15Years
175Lomatium bradshawii
(Apiales)
0 313Years0
0.5
1176
Garcinia lucida(Malpighiales)
0.0
2.5
5.0
0 72Years
177Pseudophoenix
sargentii(Arecales)
0 33Years
178Annamocarya sinensis
(Malvales)
0 77Years
179Swallenia alexandrae
(Poales)
0 26Years
180Manilkara zapota
(Ericales)
0 5Years
181Espeletia spicata
(Asterales)
0 641Years
182Molinia caerulea
(Poales)
0 9Years
183Viola pumila(Malpighiales)
0 311Years
184Paronychia pulvinata
(Caryophyllales)
0 20Years
185Lobularia maritima
(Brassicales)
0 43Years
186Atriplex canescens
(Caryophyllales)
0 6Years
187Dicerandra frutescens
(Lamiales)
0 3Years
188Ailanthus altissima
(Sapindales)
0 3Years
189Mimulus guttatus
(Lamiales)
0 158Years
190Araucaria hunsteinii
(Pinales)
0 142Years
191Atriplex vesicaria(Caryophyllales)
0 102Years
192Rourea induta
(Oxalidales)
0 11Years
193Pinguicula villosa
(Lamiales)
0 5Years
194Ambrosia deltoidea
(Asterales)
0 33Years
195Tillandsia
macdougallii(Poales)
0 758Years
196Encephalartos
villosus(Cycadales)
0 171Years
197Opuntia rastrera(Caryophyllales)
0 5Years0
0.5
1198
Oenothera deltoides(Myrtales)
0.0
2.5
5.0
0 1276Years
199Acacia bilimekii
(Fabales)
0 15Years
200Astragalus tyghensis
(Fabales)
0 106Years
201Agrimonia eupatoria
(Rosales)
0 29Years
202Andropogon semiberbis
(Poales)
0 43Years
203Trifolium montanum
(Fabales)
0 297Years
204Sclerocarya birrea
(Sapindales)
0 147Years
205Ardisia
escallonioides(Ericales)
0 6Years
206Disporum sessile
(Liliales)
0 6Years
207Festuca gracillima
(Poales)
0 39Years
208Silene regia
(Caryophyllales)
0 59Years
209Geum rivale
(Rosales)
0 16Years
210Syzygium jambos
(Myrtales)
0 5Years
211Eryngium cuneifolium
(Apiales)
0 20Years
212Werauhia
sanguinolenta(Poales)
0 41Years
213Aechmea magdalenae
(Poales)
0 28Years
214Lindera umbellata
(Laurales)
0 41Years
215Achnatherumcalamagrostis
(Poales)
0 12Years
216Haplopappus radiatus
(Asterales)
0 13Years
217Gentiana pneumonanthe
(Gentianales)
0 186Years
218Quercus rugosa
(Fagales)
0 35Years
219Ramonda myconi
(Lamiales)
0 5Years0
0.5
1220
Oxalis acetosella(Oxalidales)
0.0
2.5
5.0
0 28Years
221Scorzonera hispanica
(Asterales)
0 35Years
222Hudsonia montana
(Malvales)
0 3Years
223Aquilaria microcarpa
(Malvales)
0 595Years
224Euterpe edulis
(Arecales)
0 13Years
225Dodonaca angustifolia
(Sapindales)
0 6Years
226Catopsis sessiliflora
(Poales)
0 105Years
227Stryphnodendron
excelsum(Fabales)
0 22Years
228Calocedrus macrolepis
(Pinales)
0 5Years
229Carex humilis
(Poales)
0 6Years
230Neodypsis decaryi
(Arecales)
0 33Years
231Pterocarya rhoifolia
(Fagales)
0 51Years
232Abies magnifica
(Pinales)
0 186Years
233Borassus aethiopum
(Arecales)
0 30Years
234Machaerium cuspidatum
(Fabales)
0 302Years
235Juniperus procera
(Pinales)
0 24Years
236Lepidium davisii
(Brassicales)
0 6Years
237Aquilegia chrysantha
(Ranunculales)
0 11Years
238Hilaria mutica
(Poales)
0 998Years
239Myrsine guianensis
(Ericales)
0 78Years
240Chamaelirium luteum
(Liliales)
0 7Years
241Laminaria digitata
(Laminariales)
0 3Years0
0.5
1242
Limonium carolinianum(Caryophyllales)
0.0
2.5
5.0
0 209Years
243Kosteletzkyapentacarpos
(Malvales)
0 42Years
244Acer rurinerve
(Sapindales)
0 18Years
245Sanicula europaea
(Apiales)
0 499Years
246Roupala montana
(Proteales)
0 38Years
247Pinguicula alpina
(Lamiales)
0 4Years
248Austrostipaaristiglumis
(Poales)
0 59Years
249Pinus lambertiana
(Pinales)
0 38Years
250Miconia prasina
(Myrtales)
0 45Years
251Viburnum furcatum
(Dipsacales)
0 32Years
252Pinus kwangtungensis
(Pinales)
0 4Years
253Eupatoriumperfoliatum(Asterales)
0 65Years
254Magnolia salicifolia
(Magnoliales)
0 4Years
255Phyllanthus emblica
(Malpighiales)
0 61Years
256Geonoma macrostachys
(Arecales)
0 51Years
257Trillium grandiflorum
(Liliales)
0 22Years
258Actaea spicata(Ranunculales)
0 43Years
259Tachigali vasquezii
(Fabales)
0 33Years
260Cassia nemophila
(Fabales)
0 12Years
261Gardenia actinocarpa
(Gentianales)
0 25Years
262Ulex minor(Fabales)
0 158Years
263Sarcocapnos baetica
(Ranunculales)
0 40Years0
0.5
1264
Pentaclethramacroloba(Fabales)
0.0
2.5
5.0
0 40Years
265Parashorea chinensis
(Magnoliales)
0 150Years
266Sarracenia purpurea
(Ericales)
0 3Years
267Rhopalostylis sapida
(Arecales)
0 277Years
268Ardisia elliptica
(Ericales)
0 215Years
269Hyparrhenia diplandra
(Poales)
0 76Years
270Eremospathamacrocarpa
(Arecales)
0 40Years
271Calochortuspulchellus
(Liliales)
0 19Years
272Cleome droserifolia
(Brassicales)
0 12Years
273Arisaema serratum
(Alismatales)
0 13Years
274Arisaema triphyllum
(Alismatales)
0 21Years
275Pinus ponderosa
(Pinales)
0 12Years
276Tillandsiamulticaulis
(Poales)
0 92Years
277Rhododendron ponticum
(Ericales)
0 409Years
278Silene acaulis(Caryophyllales)
0 56Years
279Scorzonera humilis
(Asterales)
0 36Years
280Pinguicula vulgaris
(Lamiales)
0 17Years
281Acer mono(Sapindales)
0 4Years
282Chlorocardium rodiei
(Laurales)
0 198Years
283Minuartia obtusiloba
(Caryophyllales)
0 550Years
284Encephalartos
cycadifolius(Cycadales)
0 3Years
285Cirsium palustre
(Asterales)
0 81Years0
0.5
1286
Sarcocapnospulcherrima
(Ranunculales)
0.0
2.5
5.0
0 386Years
287Calocedrus decurrens
(Pinales)
0 18Years
288Armeria maritima
(Caryophyllales)
0 10Years
289Mimulus lewisii
(Lamiales)
0 4Years
290Mimulus cardinalis
(Lamiales)
0 30Years
291Harrisia fragrans(Caryophyllales)
0 31Years
292Mauritaria flexuosa
(Arecales)
0 70Years
293Tsuga canadensis
(Pinales)
0 239Years
294Dicymbe altsonii
(Fabales)
0 30Years
295Sabal yapa(Arecales)
0 69Years
296Pterocarpusangolensis(Fabales)
0 36Years
297Silene
douglasii(Caryophyllales)
0 45Years
298Primula elatior
(Ericales)
0 52Years
299Phytelephas seemannii
(Arecales)
0 9Years
300Styrax obassia
(Ericales)
0 16Years
301Acer amoenum
(Sapindales)
0 22Years
302Lepanthes rupestris
(Asparagales)
0 294Years
303Rumex obtusifolius
(Caryophyllales)
0 84Years
304Pinus nigra
(Pinales)
0 41Years
305Dacrydium elatum
(Pinales)
0 16Years
306Neobuxbaumiamacrocephala(Caryophyllales)
0 20Years
307Pterocereus gaumeri
(Caryophyllales)
0 3Years0
0.5
1308
Mammillariapectinifera
(Caryophyllales)
0.0
2.5
5.0
0 23Years
309Mammillaria
magnimamma(Caryophyllales)
0 48Years
310Avicennia germinans
(Lamiales)
0 5Years
311Cynoglossumvirginianum(Lamiales)
0 20Years
312Tillandsia juncea
(Poales)
0 3Years
313Eupatorium resinosum
(Asterales)
0 5Years
314Bouteloua rigidiseta
(Poales)
0 178Years0
0.5
1315
Miconia albicans(Myrtales)
0.0
2.5
5.0
0 6Years
316Zamia amblyphyllidia
(Cycadales)
0 101Years
317Avicennia marina
(Lamiales)
0 8Years
318Astrocaryum mexicanum
(Arecales)
0 137Years
319Bertholletia excelsa
(Ericales)
0 16Years
320Neobuxbaumia tetetzo
(Caryophyllales)
0 100Years
321Nothofagus fusca
(Fagales)
0 320Years0
0.5
1322
Aesculus turbinata(Sapindales)
0.0
2.5
5.0
0 9Years
323Araucaria
cunninghamii(Pinales)
0 6Years
324Lindera benzoin
(Laurales)
0 3Years
325Neobuxbaumiamezcalaensis(Caryophyllales)
0 4Years
326Calathea ovandensis
(Caryophyllales)
0 25Years
327Prunus africana
(Rosales)
0 4Years
328Plantago media
(Lamiales)
0 9Years0
0.5
1329
Trilliumcamschatcense
(Liliales)
Demographic trajectories in plantsM
orta
lity
(red
)R
ecru
itmen
t (b
lue)
Surv
ivor
ship
(gr
een)
Age (yrs)
Red fir (Abies magnifica)
Demographic trajectories in plants
Demographic senescence DOES occur in some species:
...but mostly only in mortality, not recruitment.
Demographic trajectories in plants
Most species are strongly negatively senescent... they get better with age.
Mortality trajectories in plants
Acknowledgements
Discussion
Life and death in the garden: mortality and recruitment trajectories in plantsOwen R. Jones* & Roberto Salguero-Gómez*Laboratory of Evolutionary Biodemography, Max Planck Institute for Demographic Research, Rostock, Germany{jones/salguero}@demogr.mpg.de; *Authors contributed equally to this work
One group of 'classical' theories of senescence focuses on the existence of trade-offs between early and late life performance. This body of literature, initiated by Hamilton1 and later supplemented by Williams2, Medawar3 and Kirkwood4, was developed with unitary organisms in mind. These theories are usually interpreted to predict that, after the onset of sexual maturity, survival rate will decline alongside reproductive output. Using data derived from the COMPADRE database5, we examined age-specific mortality and per-capita recruitment trajectories for 329 vascular plant and algae species using population projection matrices6 to examine the validity of these theories for the age trajectories of modular organisms.We demonstrate the existence of an astonishing variety of demographic trajectories in modular organisms, the majority of which do not adhere to the 'classical' Hamiltonian predictions of inevitable senescence. We argue that environmental filtering and phylogenetic ancestry explains much of the observed variation, and that the classical theories of senescence are not sufficient to explain demographic trajectories in modular organisms like vascular plants, red and brown algae.
Summary
Methods
Legend
Force of mortality
Recruitment
Survivorship
Age
Stan
dard
ized
forc
e of
mor
tality
and
recr
uitm
ent
Surv
ivors
hip
We first obtained the average, control population projection matrices for 329 species of vascular plants and algae from the COMPADRE database5. Each of these matrices contains at least three years worth of stage-based information (stage-specific survival, changes in stage, and sexual reproduction) describing the species’ population dynamics. The average, control population projection matrix represents the element-by-element average of all temporally and spatially replicated matrices in each publication but only under “natural” conditions (i.e., experimental treatments such as fires, floods, and droughts were excluded).We employed methods described by Cochran & Ellner7 to obtain age-specific survorship (lx) and per-capita recruitment (mx), as well as mean age at sexual maturity (Lα)6 and lifespan (L)6. The estimates of lifespan excluded seedbank stasis and thus inform only on the performance of photosynthetically active individuals. We then calculated the force of mortality (μx) from lx, and standardized μx and mx by their respective mean values from age zero up to the age of 99% mortality. These standardized trajectories are informative about the distribution of mortality and per-capita recruitment events throughout the life course of each species8. We have sorted the panels from top-left to bottom-right by decreasing standardized force of mortality at old-age. We have color-coded the background of each plot according to the species' growth form (see legend).
Algae Epiphyte Herbaceousperennial Succulent Shrub Liana Palm tree Tree
References
Despite predictions based on Hamiltonian theories of aging, we found no species where there was a simultaneous increase in the force of mortality (μx) together with a decrease in per-capita recruitment (mx) after maturity. A small number of species did show either increasing μx [Plot #: 2, 4, 5, 6, 7, 8, 11, 13, 19, 24, 25, 26, 49, 177] or declining mx values with age [72, 118, 189, 313]. However, the vast majority of species displayed an increase in mx and flattened or decrease in μx with age.Contrary to expectation, the top part of the aging continuum depicted in our plots is not exclusively occupied by short-lived species, nor by species that invest relatively little in tissue construction and maintenance, such as herbaceous perennials. Instead, a number of long-lived trees, palm trees, succulents, and shrubs appear throughout the continuum. The center of the continuum is mostly dominated by species where μx and mx plateau very quickly after sexual maturity. Although a plateau is ultimately expected from the type of stage-to-age transformations shown here6,7, their initial trajectories prior to the plateau are biologically meaningful properties of their populations. For example, the μx of the desert shrub Cassia nemophila [260] appears to gently fluctuate before eventually stabilizing.These trajectories offer a fascinating glimpse into the demography and senescence of modular organisms, which until now have been neglected. The factors shaping these patterns of aging, as well as the conditions under which modular organisms may escape senescence remain to be explored.Note: species which are lacking mx trajectories are either (i) strictly clonally reproductive [10, 13, 65, 103, 161, 213, 229], (ii) did not report recruitment [2], or (iii) are species where <1% of surviving individuals reproduce [306].
1. Hamilton W. 1966. J. Theor. Biol. 12, 12-45. 2. Williams G. C. 1957. Evolution 11, 398-411. 3. Medawar P. B. 1952. An unsolved problem of biology. pp. 44-70. 4. Kirkwood T. B. 1977. Nature 270, 301-304. 5. Salguero-Gómez R. Unpubl. COMPADRE: COMparative Plant & Algae Demographic REsearch Database. MPIDR. 6. Caswell H. 2001. Matrix population models: Construction, analysis and interpretation. 7. Cochran M. E. & Ellner S. 1992. Ecol. Monogr. 62, 345-264. 8. Baudisch A. 2011. Methods in Ecol. & Evol. 2, 375-382.
This work was supported by the Laboratory of Evolutionary Biodemography, and the Max Planck Research Group: Modeling the Evolution of Aging, at the Max Planck Institute for Demographic Research, as well as two fellowships to R.S-G (Teece Family Foundation, University of Pennsylvania). We acknowledge help by the COMPADRE digitalization team: E. Briks, S. Zeh, C. Farack, G. Roemer, A. Henning & G. Hoppe.
Mean age at first reproduction
0.0
2.5
5.0
0 6Years
1Ceratozamia mirandae
(Cycadales)
0 5Years
2Prunus serotina
(Rosales)
0 37Years
3Pinus sylvestris
(Pinales)
0 109Years
4Escontria chiotilla
(Caryophyllales)
0 198Years
5Coccothrinax readii
(Arecales)
0 134Years
6Thrinax radiata
(Arecales)
0 52Years
7Fagus grandifolia
(Fagales)
0 6Years
8Pterygophora
californica(Laminariales)
0 62Years
9Podococcus berteri
(Arecales)
0 16Years
10Disporum smilacinum
(Liliales)
0 128Years
11Geonoma orbignyana
(Arecales)
0 14Years
12Rhododendron maximum
(Ericales)
0 27Years
13Clintonia borealis
(Liliales)
0 458Years
14Prioria copaifera
(Fabales)
0 309Years
15Pinus palustris
(Pinales)
0 40Years
16Callitris
intratropica(Pinales)
0 998Years
17Vouacapoua americana
(Fabales)
0 17Years
18Digitaria eriantha
(Lamiales)
0 8Years
19Carlina vulgaris
(Asterales)
0 672Years
20Dicorynia guianensis
(Fabales)
0 30Years
21Alnus indicata
(Liliales)
0 12Years0
0.5
122
Eriophorumangustifolium
(Poales)
0.0
2.5
5.0
0 15Years
23Anemone patens
(Ranunculales)
0 28Years
24Ulex gallii(Fabales)
0 86Years
25Pinus jeffreyi
(Pinales)
0 11Years
26Carex membranaceae
(Poales)
0 26Years
27Lonicera maackii
(Dipsacales)
0 18Years
28Carex aquatilis
(Poales)
0 37Years
29Aristida bipartita
(Poales)
0 16Years
30Ophrys sphegodes
(Asparagales)
0 8Years
31Centaurea jacea
(Asterales)
0 15Years
32Rosa multiflora
(Rosales)
0 998Years
33Balsamorhiza
sagittata(Asterales)
0 13Years
34Podophyllum peltatum
(Ranunculales)
0 57Years
35Iris germanica(Asparagales)
0 7Years
36Gaura neomexicana
(Myrtales)
0 24Years
37Achillea millefolium
(Asterales)
0 41Years
38Geonoma brevispatha
(Arecales)
0 14Years
39Neotinea ustulata
(Asparagales)
0 671Years
40Callycophyllum
spruceanum(Gentianales)
0 561Years
41Virola surinamensis
(Magnoliales)
0 482Years
42Carapa guianensis
(Sapindales)
0 396Years
43Pouteria sagotiana
(Ericales)
0 378Years0
0.5
144
Platymiscium filipes(Fabales)
0.0
2.5
5.0
0 355Years
45Mora paraensis
(Fabales)
0 136Years
46Arnica angustifolia
(Asterales)
0 271Years
47Licaria mahuba
(Laurales)
0 43Years
48Astragalus scaphoides
(Fabales)
0 14Years
49Cheirolophus
metlesicsii(Asterales)
0 50Years
50Abies sachalinensis
(Pinales)
0 92Years
51Dactylorhiza
lapponica(Asparagales)
0 140Years
52Hymenoxys herbacea
(Asterales)
0 68Years
53Trifolium pratense
(Fabales)
0 61Years
54Pinguicula ionantha
(Lamiales)
0 98Years
55Abies concolor
(Pinales)
0 35Years
56Himantoglossum
hircinum(Asparagales)
0 181Years
57Calochortustiburonensis
(Liliales)
0 706Years
58Taxus brevifolia
(Pinales)
0 475Years
59Heliconia acuminata
(Zingiberales)
0 82Years
60Laccospermasecundiflorum
(Arecales)
0 41Years
61Primula veris
(Ericales)
0 5Years
62Tillandsia
brachycaulos(Poales)
0 14Years
63Heteropogon contortus
(Poales)
0 15Years
64Bothriochloa
insculpta(Poales)
0 8Years
65Aechmea nudicaulis
(Poales)
0 10Years0
0.5
166
Braya fernaldii(Brassicales)
0.0
2.5
5.0
0 59Years
67Banksia ericifolia
(Proteales)
0 139Years
68Petrophile pulchella
(Proteales)
0 3016Years
69Prosopis glandulosa
(Fabales)
0 24Years
70Calochortus albus
(Liliales)
0 147Years
71Chamaedorea radicalis
(Arecales)
0 8Years
72Polemoniumvanbruntiae
(Ericales)
0 13Years
73Saussurea medusa
(Asterales)
0 32Years
74Fritillaria
camtschatcensis(Liliales)
0 25Years
75Actaea elata
(Ranunculales)
0 19Years
76Calluna vulgaris
(Ericales)
0 20Years
77Scabiosa columbaria
(Dipsacales)
0 6Years
78Pseudomisopatesrivas−martinezii
(Malpighiales)
0 32Years
79Fumana procumbens
(Malvales)
0 49Years
80Helianthusdivaricatus(Asterales)
0 27Years
81Viola fimbriatula
(Malpighiales)
0 33Years
82Tillandsia recurvata
(Poales)
0 34Years
83Cimicifuga elata(Ranunculales)
0 16Years
84Centaurea maculosa
(Asterales)
0 20Years
85Setaria incrassata
(Poales)
0 89Years
86Aquilaria crassna
(Malvales)
0 38Years
87Borderea pyrenaica
(Dioscoreales)
0 57Years0
0.5
188
Succisa pratensis(Dipsacales)
0.0
2.5
5.0
0 170Years
89Rosmarinus tomentosus
(Lamiales)
0 121Years
90Cirsium acaule
(Asterales)
0 401Years
91Scaphium borneense
(Malvales)
0 42Years
92Helianthemum juliae
(Malvales)
0 95Years
93Epilobium latifolium
(Myrtales)
0 44Years
94Primula farinosa
(Ericales)
0 34Years
95Calochortusmacrocarpus
(Liliales)
0 88Years
96Geum reptans
(Rosales)
0 125Years
97Eremophila maitlandii
(Lamiales)
0 375Years
98Borderea chouardii
(Dioscoreales)
0 62Years
99Nardostachys
grandiflora(Dipsacales)
0 26Years
100Geranium sylvaticum
(Arecales)
0 33Years
101Rubus saxitalis
(Rosales)
0 196Years
102Calochortusobisopensis
(Liliales)
0 16Years
103Cleistes varicata 1
(Asparagales)
0 84Years
104Periandra
mediterranea(Fabales)
0 99Years
105Guarianthe aurantiaca
(Asparagales)
0 35Years
106Calochortus lyallii
(Liliales)
0 9Years
107Arabis fecunda
(Brassicales)
0 85Years
108Lathyrus vernus
(Fabales)
0 25Years
109Clidemia hirta
(Myrtales)
0 39Years0
0.5
1110
Artemisia genipi(Asterales)
0.0
2.5
5.0
0 7Years
111Sarcocapnosenneaphylla
(Ranunculales)
0 34Years
112Acer saccharum
(Sapindales)
0 44Years
113Ipomoea leptophylla
(Solanales)
0 6Years
114Daucus carota
(Apiales)
0 37Years
115Aconitum
noveboracense(Ranunculales)
0 16Years
116Lomatium cookii
(Apiales)
0 16Years
117Manglietia fordiana
(Magnoliales)
0 7Years
118Hypericum cumulicola
(Malpighiales)
0 8Years
119Iridaea splendens
(Gigartinales)
0 14Years
120Asclepias meadii
(Gentianales)
0 237Years
121Ptychosperma
macarthurii(Arecales)
0 25Years
122Lespedeza cuneata
(Fabales)
0 24Years
123Hydrastis canadensis
(Ranunculales)
0 238Years
124Licania heteromorpha
(Brassicales)
0 5Years
125Cirsium vulgare
(Asterales)
0 9Years
126Atriplex acanthocarpa
(Caryophyllales)
0 399Years
127Eremophila forrestii
(Lamiales)
0 5Years
128Linum catharticum
(Malpighiales)
0 96Years
129Echinaceaangustifolia(Asterales)
0 24Years
130Themeda triandra
(Poales)
0 150Years
131Opuntia macrocentra
(Caryophyllales)
0 7Years0
0.5
1132
Gelidium sesquipedale(Gelidiales)
0.0
2.5
5.0
0 45Years
133Fritillaria meleagris
(Liliales)
0 37Years
134Coryphantarobbinsorum
(Caryophyllales)
0 419Years
135Lepanthes eltoroensis
(Asparagales)
0 45Years
136Aster amellus
(Asterales)
0 18Years
137Allium monanthum
(Liliales)
0 3Years
138Dipsacus sylvestris
(Dipsacales)
0 68Years
139Panax quinquefolium
(Apiales)
0 9Years
140Viola elatior(Malpighiales)
0 12Years
141Braya longii(Brassicales)
0 17Years
142Asarum canadense
(Piperales)
0 4Years
143Tragopogon dubius
(Asterales)
0 14Years
144Lupinus arboreus
(Fabales)
0 6Years
145Carnegiea gigantea
(Caryophyllales)
0 38Years
146Saxifraga cotyledon
(Saxifragales)
0 7Years
147Rumex rupestris(Caryophyllales)
0 6Years
148Taraxacum officinale
(Asterales)
0 21Years
149Chamaecrista keyensis
(Fabales)
0 20Years
150Tolumnia variegata
(Asparagales)
0 8Years
151Centaurea corymbosa
(Asterales)
0 11Years
152Zea diploperennis
(Poales)
0 11Years
153Aquilegia sp.(Ranunculales)
0 10Years0
0.5
1154
Tillandsia punctulata(Poales)
0.0
2.5
5.0
0 653Years
155Sarracenia alata
(Ericales)
0 5Years
156Cirsium dissectum
(Asterales)
0 10Years
157Lupinus tidestromii
(Fabales)
0 9Years
158Allium tricoccum
(Liliales)
0 27Years
159Anthyllis alpicola
(Fabales)
0 33Years
160Echeveria longissima
(Saxifragales)
0 5Years
161Cleistes varicata 2
(Asparagales)
0 16Years
162Viola stagnina(Malpighiales)
0 81Years
163Primula vulgaris
(Ericales)
0 272Years
164Duguetia neglecta
(Magnoliales)
0 11Years
165Cryptantha flava
(Lamiales)
0 32Years
166Aspasia principissa
(Asparagales)
0 9Years
167Bothriochloaischaemum
(Poales)
0 32Years
168Betula nana
(Fagales)
0 10Years
169Tillandsia deppeana
(Poales)
0 15Years
170Hydrangea paniculata
(Cornales)
0 36Years
171Pedicularisfurbishiae
(Malpighiales)
0 33Years
172Cytisus scoparius
(Fabales)
0 82Years
173Iriartea deltoidea
(Asparagales)
0 8Years
174Hypochaeris radicata
(Asterales)
0 15Years
175Lomatium bradshawii
(Apiales)
0 313Years0
0.5
1176
Garcinia lucida(Malpighiales)
0.0
2.5
5.0
0 72Years
177Pseudophoenix
sargentii(Arecales)
0 33Years
178Annamocarya sinensis
(Malvales)
0 77Years
179Swallenia alexandrae
(Poales)
0 26Years
180Manilkara zapota
(Ericales)
0 5Years
181Espeletia spicata
(Asterales)
0 641Years
182Molinia caerulea
(Poales)
0 9Years
183Viola pumila(Malpighiales)
0 311Years
184Paronychia pulvinata
(Caryophyllales)
0 20Years
185Lobularia maritima
(Brassicales)
0 43Years
186Atriplex canescens
(Caryophyllales)
0 6Years
187Dicerandra frutescens
(Lamiales)
0 3Years
188Ailanthus altissima
(Sapindales)
0 3Years
189Mimulus guttatus
(Lamiales)
0 158Years
190Araucaria hunsteinii
(Pinales)
0 142Years
191Atriplex vesicaria(Caryophyllales)
0 102Years
192Rourea induta
(Oxalidales)
0 11Years
193Pinguicula villosa
(Lamiales)
0 5Years
194Ambrosia deltoidea
(Asterales)
0 33Years
195Tillandsia
macdougallii(Poales)
0 758Years
196Encephalartos
villosus(Cycadales)
0 171Years
197Opuntia rastrera(Caryophyllales)
0 5Years0
0.5
1198
Oenothera deltoides(Myrtales)
0.0
2.5
5.0
0 1276Years
199Acacia bilimekii
(Fabales)
0 15Years
200Astragalus tyghensis
(Fabales)
0 106Years
201Agrimonia eupatoria
(Rosales)
0 29Years
202Andropogon semiberbis
(Poales)
0 43Years
203Trifolium montanum
(Fabales)
0 297Years
204Sclerocarya birrea
(Sapindales)
0 147Years
205Ardisia
escallonioides(Ericales)
0 6Years
206Disporum sessile
(Liliales)
0 6Years
207Festuca gracillima
(Poales)
0 39Years
208Silene regia
(Caryophyllales)
0 59Years
209Geum rivale
(Rosales)
0 16Years
210Syzygium jambos
(Myrtales)
0 5Years
211Eryngium cuneifolium
(Apiales)
0 20Years
212Werauhia
sanguinolenta(Poales)
0 41Years
213Aechmea magdalenae
(Poales)
0 28Years
214Lindera umbellata
(Laurales)
0 41Years
215Achnatherumcalamagrostis
(Poales)
0 12Years
216Haplopappus radiatus
(Asterales)
0 13Years
217Gentiana pneumonanthe
(Gentianales)
0 186Years
218Quercus rugosa
(Fagales)
0 35Years
219Ramonda myconi
(Lamiales)
0 5Years0
0.5
1220
Oxalis acetosella(Oxalidales)
0.0
2.5
5.0
0 28Years
221Scorzonera hispanica
(Asterales)
0 35Years
222Hudsonia montana
(Malvales)
0 3Years
223Aquilaria microcarpa
(Malvales)
0 595Years
224Euterpe edulis
(Arecales)
0 13Years
225Dodonaca angustifolia
(Sapindales)
0 6Years
226Catopsis sessiliflora
(Poales)
0 105Years
227Stryphnodendron
excelsum(Fabales)
0 22Years
228Calocedrus macrolepis
(Pinales)
0 5Years
229Carex humilis
(Poales)
0 6Years
230Neodypsis decaryi
(Arecales)
0 33Years
231Pterocarya rhoifolia
(Fagales)
0 51Years
232Abies magnifica
(Pinales)
0 186Years
233Borassus aethiopum
(Arecales)
0 30Years
234Machaerium cuspidatum
(Fabales)
0 302Years
235Juniperus procera
(Pinales)
0 24Years
236Lepidium davisii
(Brassicales)
0 6Years
237Aquilegia chrysantha
(Ranunculales)
0 11Years
238Hilaria mutica
(Poales)
0 998Years
239Myrsine guianensis
(Ericales)
0 78Years
240Chamaelirium luteum
(Liliales)
0 7Years
241Laminaria digitata
(Laminariales)
0 3Years0
0.5
1242
Limonium carolinianum(Caryophyllales)
0.0
2.5
5.0
0 209Years
243Kosteletzkyapentacarpos
(Malvales)
0 42Years
244Acer rurinerve
(Sapindales)
0 18Years
245Sanicula europaea
(Apiales)
0 499Years
246Roupala montana
(Proteales)
0 38Years
247Pinguicula alpina
(Lamiales)
0 4Years
248Austrostipaaristiglumis
(Poales)
0 59Years
249Pinus lambertiana
(Pinales)
0 38Years
250Miconia prasina
(Myrtales)
0 45Years
251Viburnum furcatum
(Dipsacales)
0 32Years
252Pinus kwangtungensis
(Pinales)
0 4Years
253Eupatoriumperfoliatum(Asterales)
0 65Years
254Magnolia salicifolia
(Magnoliales)
0 4Years
255Phyllanthus emblica
(Malpighiales)
0 61Years
256Geonoma macrostachys
(Arecales)
0 51Years
257Trillium grandiflorum
(Liliales)
0 22Years
258Actaea spicata(Ranunculales)
0 43Years
259Tachigali vasquezii
(Fabales)
0 33Years
260Cassia nemophila
(Fabales)
0 12Years
261Gardenia actinocarpa
(Gentianales)
0 25Years
262Ulex minor(Fabales)
0 158Years
263Sarcocapnos baetica
(Ranunculales)
0 40Years0
0.5
1264
Pentaclethramacroloba(Fabales)
0.0
2.5
5.0
0 40Years
265Parashorea chinensis
(Magnoliales)
0 150Years
266Sarracenia purpurea
(Ericales)
0 3Years
267Rhopalostylis sapida
(Arecales)
0 277Years
268Ardisia elliptica
(Ericales)
0 215Years
269Hyparrhenia diplandra
(Poales)
0 76Years
270Eremospathamacrocarpa
(Arecales)
0 40Years
271Calochortuspulchellus
(Liliales)
0 19Years
272Cleome droserifolia
(Brassicales)
0 12Years
273Arisaema serratum
(Alismatales)
0 13Years
274Arisaema triphyllum
(Alismatales)
0 21Years
275Pinus ponderosa
(Pinales)
0 12Years
276Tillandsiamulticaulis
(Poales)
0 92Years
277Rhododendron ponticum
(Ericales)
0 409Years
278Silene acaulis(Caryophyllales)
0 56Years
279Scorzonera humilis
(Asterales)
0 36Years
280Pinguicula vulgaris
(Lamiales)
0 17Years
281Acer mono(Sapindales)
0 4Years
282Chlorocardium rodiei
(Laurales)
0 198Years
283Minuartia obtusiloba
(Caryophyllales)
0 550Years
284Encephalartos
cycadifolius(Cycadales)
0 3Years
285Cirsium palustre
(Asterales)
0 81Years0
0.5
1286
Sarcocapnospulcherrima
(Ranunculales)
0.0
2.5
5.0
0 386Years
287Calocedrus decurrens
(Pinales)
0 18Years
288Armeria maritima
(Caryophyllales)
0 10Years
289Mimulus lewisii
(Lamiales)
0 4Years
290Mimulus cardinalis
(Lamiales)
0 30Years
291Harrisia fragrans(Caryophyllales)
0 31Years
292Mauritaria flexuosa
(Arecales)
0 70Years
293Tsuga canadensis
(Pinales)
0 239Years
294Dicymbe altsonii
(Fabales)
0 30Years
295Sabal yapa(Arecales)
0 69Years
296Pterocarpusangolensis(Fabales)
0 36Years
297Silene
douglasii(Caryophyllales)
0 45Years
298Primula elatior
(Ericales)
0 52Years
299Phytelephas seemannii
(Arecales)
0 9Years
300Styrax obassia
(Ericales)
0 16Years
301Acer amoenum
(Sapindales)
0 22Years
302Lepanthes rupestris
(Asparagales)
0 294Years
303Rumex obtusifolius
(Caryophyllales)
0 84Years
304Pinus nigra
(Pinales)
0 41Years
305Dacrydium elatum
(Pinales)
0 16Years
306Neobuxbaumiamacrocephala(Caryophyllales)
0 20Years
307Pterocereus gaumeri
(Caryophyllales)
0 3Years0
0.5
1308
Mammillariapectinifera
(Caryophyllales)
0.0
2.5
5.0
0 23Years
309Mammillaria
magnimamma(Caryophyllales)
0 48Years
310Avicennia germinans
(Lamiales)
0 5Years
311Cynoglossumvirginianum(Lamiales)
0 20Years
312Tillandsia juncea
(Poales)
0 3Years
313Eupatorium resinosum
(Asterales)
0 5Years
314Bouteloua rigidiseta
(Poales)
0 178Years0
0.5
1315
Miconia albicans(Myrtales)
0.0
2.5
5.0
0 6Years
316Zamia amblyphyllidia
(Cycadales)
0 101Years
317Avicennia marina
(Lamiales)
0 8Years
318Astrocaryum mexicanum
(Arecales)
0 137Years
319Bertholletia excelsa
(Ericales)
0 16Years
320Neobuxbaumia tetetzo
(Caryophyllales)
0 100Years
321Nothofagus fusca
(Fagales)
0 320Years0
0.5
1322
Aesculus turbinata(Sapindales)
0.0
2.5
5.0
0 9Years
323Araucaria
cunninghamii(Pinales)
0 6Years
324Lindera benzoin
(Laurales)
0 3Years
325Neobuxbaumiamezcalaensis(Caryophyllales)
0 4Years
326Calathea ovandensis
(Caryophyllales)
0 25Years
327Prunus africana
(Rosales)
0 4Years
328Plantago media
(Lamiales)
0 9Years0
0.5
1329
Trilliumcamschatcense
(Liliales)
Weakly positive senescence
‘Negative’ senescence
• Great deal of variation among species.
• Huge majority incongruent with classical senescence theory.
• No pattern in relation to growth form, nor taxonomy.
Demographic trajectories in plants
Replication in plant demographySpatial replication in COMPADRE
Number of replicates
Freq
uenc
y
0 5 10 15 20 25 30 35
0
100
200
300
400
500
Temporal replication in COMPADRE
Length of study (yrs)
Freq
uenc
y
0 20 40 60
0
50
100
150
Spatial replication
Study length
• Most species are only known (demographically) from a single site.
• Most studies are about as long as a PhD.
• How representative are the data?
Demographic plasticity
• Response to ecological factors (population density, temperature, rainfall, nutrients, pH,...)
• Genetic differences.
• Will study-site selection affect life-history inferences?
• Will study-site selection affect estimates of phylogenetic signal?
• Can we do meaningful research with low replication?
How much does this variation really matter?
Within-species variation in senescence trajectories
Age (yrs)
Stan
dard
ised
mor
talit
y
0 5 10 15
01
23
45
Mortality
Age (yrs)
Stan
dard
ised
recr
uitm
ent
0 5 10 15
01
23
45
RecruitmentSt. Johns wort (Hypericum cumulicola)
Rock babyroot orchid (Lepanthes rupestris)
Age (yrs)
Stan
dard
ised
mor
talit
y
0 5 10 15 20
01
23
45
Mortality
Age (yrs)
Stan
dard
ised
recr
uitm
ent
0 5 10 15 20
01
23
45
Recruitment
Extract measures
• Mean age at first reproduction
• Mean life span
• Life expectancy at maturity
• Keyfitz’s entropy (survival)
• Demetrius’ entropy (degree of iteroparity)
• Phylogenetic signal
EntropyKeyfitz’s entropy
(survival)
Demetrius’ entropy(degree of iteroparity)
• High repeatability for age at maturity and Keyfitz entropy.
• Low repeatability for other traits.
Repeatability
Age atMaturity
Keyfitzentropy
Demetriusentropy
LifeExpectancy Lifespan
Trait
Rep
eata
bilit
y
0.0
0.2
0.4
0.6
0.8
1.0
Phylogenetic signalDemetrius
entropyKeyfitz’sentropy
Age atmaturity
Life expectancyat maturity
Life span
K
Frequency
0.0 0.2 0.4 0.6 0.8 1.0
050
100
150
200
p−value
Frequency
0.0 0.2 0.4 0.6 0.8 1.0
0200
400
600
800
K0.0 0.2 0.4 0.6 0.8 1.0
p−value0.0 0.2 0.4 0.6 0.8 1.0
K0.0 0.2 0.4 0.6 0.8 1.0
p−value0.0 0.2 0.4 0.6 0.8 1.0
K0.0 0.2 0.4 0.6 0.8 1.0
p−value0.0 0.2 0.4 0.6 0.8 1.0
K0.0 0.2 0.4 0.6 0.8 1.0
p−value0.0 0.2 0.4 0.6 0.8 1.0
Using methods from Ives et al. 2007
Conclusions
• Plant demography is incongruent with classical senescence theory.
• There’s plenty of data available to address plant comparative demography.
• ... but we lack replication.
• Trait selection may be critical.
• Opportunity for research into demographic plasticity.
Acknowledgements
✴ Jonathan Silvertown and Miguel Franco for starting COMPADRE.
✴ “Compadrinos” - for data entry.
✴ Max Planck Institute for Demographic Research
✴ Max Planck Institute Independent Research Group for Modeling the Evolution of Aging
Demographic trajectories in plantsArisaema triphillum
Cypripedium acaule
Pinus palustris
Silvertown et al. (2001)