Semelparity and Iteroparity

By: Truman P. Young (Department of Plant Sciences, University of California at

Davis) © 2010 Nature Education

Citation: Young, T. P. (2010) Semelparity and Iteroparity. Nature Education

Knowledge 1(9):2
Why do some organisms die immediately after reproducing (some salmon and
bamboos, many insects, and all grain crops), while others live on to reproduce
repeatedly (most plants and vertebrates)

Semelparity

Figure 1: Semelparous plants

Like all grain crops, wheat is
semelparous plant. Annual
plants die after first
reproduction within a year of
germination. (Courtesy of
Simon Howell:
freedigitalphotos.net)
Many plant and animal species
have life histories characterized
by death after first
reproduction. This is called
semelparity, and its alternative
(living to reproduce repeatedly)
is called iteroparity. In plants,
the terms monocarpy and
polycarpy are sometimes used
instead of semelparity and
iteroparity. However, monocarpy can also be used more restrictively to describe
plants in which individual shoots die after reproducing, but not necessarily the entire
plant.

Examples of short-lived semelparous species include annual and biennial plants

(including all grain crops, and many herbaceous vegetables), and certain invertebrate
species, including many spiders (Figure 1). There are even a few semelparous
marsupial mammals in Australia. It is important to note that while all annual plants
are semelparous, not all perennial plants are iteroparous. There are a wide variety of
plant and animal species that live for many years before a single, massive, fatal
reproductive episode (certain species of salmon, bamboo, and century plants, Figure
2).

Semelparity has evolved independently many times, apparently as a derived state

from iteroparous ancestors. This dramatic life history difference offers a model
system for theoretical and empirical studies of adaptive evolution.
Dilemma and Cost of Reproduction
Natural selection maximizes total lifetime reproductive output. So how could
evolution ever favor programmed death after first reproduction? A key clue lies in the
observation that semelparous species typically produce more offspring in their single
reproductive episode than closely related species do in any one of theirs. It appears
that when an organism does not need to withhold some resources to ensure future
survival and reproduction, it can mobilize virtually all available resources to put into a
single, massive reproductive episode. For example, this fecundity advantage is two to
fivefold in plants. So the essential question becomes, "Under what conditions does the
increase in fecundity associated with semelparity more than compensate for the loss
of potential subsequent reproductive episodes?"

Figure 2: Spawning salmon

Salmon at their spawning grounds, far from the ocean. Here, they will reproduce and
die. (Courtesy of Friends of Butte Creek)

Theoretical Approaches
This question has been the subject of a rich theoretical literature. These models fall
into three classes, all of which assume a tradeoff between reproduction and survival:

1. Demographic models predict that when adult survival is low enough (relative
to juvenile survival), evolution should abandon withholding resources for a
future reproduction that is unlikely, and instead favor semelparity (Figure3).
 2. Bet-hedging models predict that when adult survival is highly variable,
evolution should favor iteroparity, because it does not risk putting all
reproductive effort into a single reproductive episode.
3. Models incorporating non-linear patterns of reproductive costs and benefits
predict that semelparity should be more likely to evolve when most of the
costs of reproduction (reduction in future survival or reproduction caused by
increases in current reproduction) happen even at low levels of reproductive
effort, or conversely, when the benefits of reproduction accrue most rapidly at
high levels of reproductive effort.

Figure 3: Schematic of the demographic model of the evolution of semelparity

Imagine semelparous and iteroparous populations. The semelparous (annual)
individuals produce 2.5 times as many seeds as the iteroparous individuals (a
reasonable estimate of relative fecundity, from natural systems). In the first year, the
semelparous individuals out-produce the iteroparous, but the iteroparous have some
probability of living to reproduce again. If the iteroparous population experiences
50% adult mortality each year, the average individual will produce 200 seeds in its
lifetime, fewer than in the semelparous population. If the iteroparous population
experiences 30% adult mortality each year, the average individual will produce 340
seeds in its lifetime, more than in the semelparous population. Therefore, in
populations with sufficiently high mortality, semelparity will be favored over
iteroparity. Demographic models of semelparity explore this more explicitly.
Empirical Evidence
Despite the abundance of theoretical models for the evolution of semelparity, direct
empirical tests for each remain limited. For example, many species of annual plants
are desert species or weedy species that are early arrivals after disturbance in many
ecosystems. Both of these groups are subject to high variation in survivorship, in
opposition to the predictions of the bet-hedging model.

In a particularly elegant test of the non-linearity model, it was demonstrated that the
taller inflorescences of semelparous yucca species produced disproportionately more
seeds than smaller inflorescences, but that this was not true in iteroparous yucca
species. Similarly, pollinators preferred taller inflorescences of semelparous yucca
species, but not iteroparous species. Although this observation fits the non-linear
theory nicely, it has been pointed out that many of these species may not be
pollinator-limited, and that inflorescence height patterns may be physiological
consequences of life history differences, and not their evolutionary causes.

There have been several more successful tests of the demographic model, and they all
show that semelparity is more likely in species (or populations) where adult survival
would be low even if they were not semelparous. These tests come from diverse
systems, including spiders, fish, an alpine mustard, and a giant rosette plant. In
addition, both desert annuals and early successional annuals live in habitats where
survival beyond the growing season might be expected to be low.

Synchronous Semelparity
There is an unusual pattern in semelparous plants characterized by single-aged
populations that live for many years, then synchronously flower and die. Certain
bamboos species are well known to exhibit this pattern, but other examples include
certain palm species, shrubs in the family Acanthaceae, and even a tropical forest
canopy tree. All of these grow in mesic forests that are climatically more moderate
than the extreme environments characteristic of most semelparous species (e.g.,
deserts, alpine habitats, bogs, disturbed sites). Periodic cicadas in the eastern U.S. also
exhibit this pattern. Predator satiation has been invoked to explain both synchrony and
semelparity in these species, but there is no widespread consensus on its causes.

Species "Approaching" Semelparity

Semelparity and iteroparity have been represented here as a simple dichotomy.
Nevertheless, the conceptual framework can be applied more generally. Several
examples have been documented where high levels of adult mortality appear to be
related to iteroparous life histories characterized by early reproduction that is more
frequent and/or more intense. These include subalpine fir trees, patas monkeys, and
several insect species.

Semelparity in Grain Crops

It is probably no coincidence that many herbaceous crops, including virtually all grain
crops, are annuals. Their semelparity results in far higher yields than if they were
iteroparous. It is likely that when selecting grain species for cultivation, early farmers
deliberately chose those species with the highest yields, which were annuals, or
selected strongly enough for high yields that iteroparous species evolved into
semelparous species. The closest relatives of both rice and maize are iteroparous.