theOLiVA OLIVA POPULATIONS

© Maurizio A. Perini
Card n.01-B -  Revision: 01/2011 – Page 1-6

Note

This brief article has an informative purpose only so I apologize to the experts of
population biology if the patterns presented here are highly simplified. I just want to
help the less experienced collectors to understand the intraspecific variability of this
genus and why it’s often so difficult to identify a single species.
Please note that what you can read in these few pages is valid also for many animal
species, not only for the olives.

Introduction

Here I want to show how the oliva populations are present in their areas of diffusion
and how this causes many problems in order to identify these wonderful shells. Even if
sometimes it’s exactly the study of the populations that allows us to individuate the
species within a “complex”. This is what happened to Vassart and Tursch (1999) in
separating Oliva truncata Marrat, 1867 from Oliva polpasta Duclos, 1833 and to Perini
(2000) in separating Oliva samarensis Johnson, 1915 from Oliva oliva (Linnaeus,
1758).

Discussion

We consider a hypothetical oliva species called “Oliva A” (fig.1) and by a white circle
its area of diffusion (fig.2). In this area Oliva A will be present only in suitable habitats
for its survival, here indicated with blue patches (fig.3).

All the specimens living in the area of diffusion (white circle) form the population of
Oliva A. The scientific name of one population living in a place (blue patch) is
“metapopulation” but I prefer to use the word "colony".

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The population of Oliva A can occur in two ways:

1) The specimens of all the colonies are equal to each other (fig. 4). Then the
variability among specimens from a colony is the same as between specimens from
different colonies. In other words, by looking at two specimens of Oliva A we are not
able to understand if they come from the same location (colony) or from different
locations (different colonies). This is the case of Oliva porphyria (Linnaeus, 1758) that
shows the same variability across its distribution area.

2) Each colony shows different specimens (fig.5), usually called “forms”. So, for
example, the specimens of the colony “An” are quite different from the specimens of
the colony “Ab”. It is easy to understand that the specimens of two neighboring
colonies (Ac-Ad or An-Ax) are similar to each other such as the reverse is true for
colonies very far apart (Ah-Ae or Aa-Ag). This is the case of Oliva spicata (Röding,
1798) and many other Oliva species.

Plate 1. Oliva spicata (Röding, 1798) from five different places (colonies) in Baja
California, Mexico.

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Looking closely at fig.5 it is easy to see how it is not possible to identify some
subspecies. My opinion is that it is very difficult if not impossible to name a subspecies
in the Genus Oliva. Indeed an author can name a subspecies only when a situation like
the one shown in fig.6 is present. In this figure we can see Oliva nitidula Duclos, 1835
(Aa) living in the East Africa (coasts and islands) and the subspecies Oliva nitidula
sandwicensis Pease, 1860 (Ab) which lives in the Hawaiian islands.

The fig.7 shows an impossible case. Two colonies, especially if they are very far apart,
cannot have specimens showing the same form (Ak-Ak). If this is the case we have to
think that the olives in Ak do not belong to the species “A” (fig. 8). If the same form,
here called “B”, is present in different places we can think that this is a different
species. (Note: here I use Oliva oliva only as an example)

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But if also the species “B” is present with different forms in each colony (fig.9) there is
no way to separate different species, if they exist. In this case we can only hope to
find two species together (sympatric, better syntopic) in more then one place as we
can see in the fig.10. Here Oliva oliva (A) is separated from Oliva samarensis (B).

Sympatry and syntopy

Since nearly all the Oliva species are based only upon differences in their external
aspect (morphological gaps) we may think that the sympatry is a more correct way to
identify the species. About the concept of sympatry Cain in 1953 wrote “ population
the individuals of which are within cruising range of each other during the breeding
season even though the habitats in which they occur do not overlap in space”. Anyway
Tursch in 1994 thought that the concept of sympatry is too broad for the genus Oliva,
so he wrote “In the case of the Oliva (at least) it is obviously safer to replace our
broad criterion of sympatry by that of syntopy”. About the concept of syntopy Lincoln
et al. in 1982 wrote “pertaining to populations or species that occupy the same
microhabitat, are observable in close proximity and could thus interbreed”.
But also the syntopy is not always the solution, and I will explain why.

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The “circular overlap”

The circular overlap or the “island problem” as I call it, is a distribution pattern of the
colonies well known to the biologists. Even if it may seem quite strange, this pattern
is very often observed in nature. It causes us to question the validity of the use of
syntopy in order to separate two different species.

Thus we see this pattern with the help of two simple figures. In the fig.11 we can see
an hypotethical island called “Happy island”. On this island there are five beaches
separated by five cliffs. One day a colony of Oliva A develops on the first beach (Step
1). This colony “Aa” remains isolated for a long time. Then, some specimens reach the
second beach and form a new colony “Ab” (Step 2). Also this colony remains isolated
for a long time. Then history repeats itself and another colony “Ac” grows on the third
beach (Step 3) … and so on till the fifth beach (fig.12).

After long periods of isolation, depending on how genetic drift has acted, the last
colony may (fig.12-K “Ba”) or may not (fig.12-Y “Ae”) be made up of a new species.
Anyway, when some individuals move from the last beach to the first one meeting the
colony “Aa”, there may be not two but four different situations. In both “K” and “Y”
there are now two syntopic colonies on the first beach: “Ba-Aa” in K and “Ae-Aa” in Y.
Each of these situations has two options, and we will see how.

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K-1 The specimens of the colony “Ba” are now a new species. They are very different
in size, color and shape so we can easily distinguish the species “Ba” from “Aa”.
In this case the syntopy is helpful: we see 2 species, which is correct.
K-2 The specimens of the colony “Ba” are now a new species. The specimens of this
new species only show a different behavior (or have different trace pheromons) but
they are very similar in appearance to the species “Aa”. Then, even if we can not
distinguish them visually, “Ba” and “Aa” do not interbreed. In this case these two
species are called “sibling species”. The sibling species are quite present in nature
then it would be strange not to find them also among the Oliva species.
In this case the syntopy is deceptive: we see 1 species but they are 2, so we are
wrong.
Y-1 The specimens which belong to the colony “Ae” are very different in size, color
and shape but they still interbreed with the specimens of the colony “Aa”. Then “Ae”
and “Aa” belong to the population of the Oliva A (fig.5), but here the difference is
that the two forms live in the same place.
In this case the syntopy is deceptive: we see 2 species but it is only 1, so we are
wrong.
Y-2 The specimens which belong to the colony “Ae” are very similar to the specimens
of the colony “Aa” (see fig.4 “Ae=Aa”, or A=A).
In this case the syntopy is unhelpful: we see 1 species (even if we see only one
colony, not two), which is correct.

By this example we can see that the syntopy helps to us to distinguish two species
only in one case (K-1). But if the syntopy is present in more than one place we can
think that the “circle overlap” pattern is not present so we are sure that we are really
observing two different species.

Plate 2. Oliva sp. from Pangandaran Bay, SW Java, Indonesia.

Conclusion
Having read this far it is clear that the populations study is only a step in trying to
identify a single species. We can not work only with the shell (morphological gaps) but
other studies on genetic, ecology and on their behavior are necessary.

OLiVA
Maurizio A. Perini
Contrada Zausa 4, 36015 St.Caterina di Schio (VI) – ITALY
the e-mail: melapium@yahoo.it

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