INTERTIDAL ECOLOGY OF THE NORTHERNMOST PART OF THE

CHILEAN ARCHIPELAGO
REPORT NO. 50 OF THE LUND UNIVERSITY CHILE EXPEDITION 1948-49 1
HANS BRATTSTROM

BRATISTROM, HANS 199006 20. Intertidal ecology of the northernmost part of the Chilean
SARSIA archipelago. Report no. 50 of the Lund University Chile Expedition 1948-491 . - Sarsia
75:107-160. Bergen. ISSN 0036-4827.

Species composition and zonation patterns at 12 hard-bottom localities are described and
figured. A total of 212 marine species was identified, 135 animals and 77 algae. All spe-
cies, even those not discussed, are enumerated in an appendix with information on their
vertical distribution at each station. The number of algal species was about equal in Seno
Reloncavi and Ancud, whereas the number of animal species was lower in Ancud and
very low in the brackish fjord Estero Reloncavf. Several very common species in Seno
Reloncavi were not found in Ancud and vice versa. Some species had a somewhat higher
relative upper limit in Ancud. These differences are probably mainly due to differences
in exposure and substrate, rocks in Ancud and boulders and stones in sand in Seno Relon-
cavi. .
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The general zonation follows the universal pattern with an infralittoral fringe lowest
down, above which is a littoral region with sub-zones, and upmost a supralittoral region.
Based on the common species, which characterize certain levels, nine distribution types
were distinguished. The vertical distribution at the same stations of some related species
or such with a supposedly similar biology is compared. Most species preferred different
levels but there was much overlapping. The zonation, mainly of algae, at four stations in
Seno Reloncavi, described by Alveal and Romo, is compared with that of the neigh-
bouring stations in the present investigation. The vertical distribution of the few species
treated in both investigations was similar.
The fairly meagre information in the literature on intertidal zonation in Chile has been
compiled for comparison. Though some southern species are not found in central and
northern Chile and some northern species don't reach the Magellanic province or only
occur in its northernmost part, many of the most common and characteristic intertidal
species are distributed along the whole coast or a great part of it. Because of that the
species composition and zonation patterns are fairly uniform from Cabo de Homos to
Arica, a result of the relatively small difference in surface and air temperature between
south and north, caused by the Humboldt Current, which carries cold water far north,
and upwelling in central and northern Chile.

Hans Brattstrom, Department of Marine Biology, University of Bergen, N-5065 Blomster-

dalen, Norway.
1 This report concludes the series.

BRATISTROM, HANS 1990 06 20. Ecologia intermareal de la parte norte del archipielago
SARSIA chileno austral. Informe numero 50 de la Expedici6n de la Universidad de- Lund a Chile
1948-491 . - Sarsia 75:107-160. Bergen. ISSN 0036-4827.

Se describe e i1ustra la composici6n de especies y los patrones de zonaci6n en 12 localida-

des con sustratos duros. Se identific6 un total de 212 especies marinas: 135 animales y
77 algas. En cada estaci6n las especies, aun aquellas no discutidas, se enumeran en un
apendice con informaci6n sobre su distribuci6n vertical. EI numero de ·especies algales fue
casi igual en Seno Reloncavi y Ancud, mientras que el numero de especies animales fue
inferior en Ancud y muy bajo en el fiordo salobre Estero Reloncavi. Varias especies muy
comunes en Seno Reloncavi no se encontraron en Ancud y viceversa. Otras especies te-
nian el limite superior relativo algo mas alto en Ancud. Estas diferencias se deben proba-
blemente a diferencias en exposici6n y sustrato, rocas en Ancud y bloques y piedras en
el Seno Reloncavi.
La zonaci6n general sigue e1 modelo universal con una franja infralitoral mas baja, so-
bre la cual hay una regi6n litoral con subzonas y una regi6n supralitoral arriba. Se distin-
guieron 9 tipos de distribuci6n basados en especies que caracterizan ciertos niveles. En
las mismas estaciones se compara la distribuci6n vertical de algunas especies relacionadas
o aquellas con una supuesta biologia similar. La mayoria de las especies preferia niveles
diferentes pero habia mucha sobreposici6n. Se compara la zonaci6n, principalmente de
algas, en 4 estaciones del Seno de Re10ncavi descrita por Alveal y Romo, con la de esta-

107
 ciones vecinas de la presente investigaci6n. La distrlbuci6n vertical de las pocas especies
tratadas en ambas investigaciones fue similar. '
Se ha compilado para comparaci6n, la informaci6n bastante escasa en la literatura exis-
tente, sobre zonaci6n intermareal en Chile. Aunque algunas especies del sur no se encuen-
tran en el centro y norte de Chile y algunas especies del norte no alcanzan la Provincia
Magallancia u ocurren s6lo en su parte norte, muchas de las especies intermareales comu-
nes y caracteristicas se distinguen a 10 largo de toda la costa 0 sobre gran parte de ella.
A causa de eso, la composici6n de especies y tipos de zonaci6n son bastante uniformes
desde Arlca al Cabo de Homos, un resultado de las diferencias relativamente pequeiias
en temperatura superficial y del aire entre el norte y el sur causada por la corriente de
Humboldt, que transporta aguas frias hacia el norte y las surgencias de Chile Central y
Norte.
Hans Brattstrom, Departamento de Biologia Marina, Universidad de Bergen, N-5065 Bloms-
terdalen, Noruega.

1 Este informe termina la serle.

CONTENTS
Introduction ....... ................................. lOS The Ancud area ................................. 126 Earlier investigations .......................... 143
Material and methods .......... .................. 109 Stn MlO ........................................ 126 Chile south of 48° S ......................... 144
Material ......................................... ,. 109 Stn M55 ........................................ 127 The Chilo. area ....... .............. ......... 145
The stations ...... ................ ..... ...... ..... 109 Stn M56 ........................................ 128 ValdivialMehuin ............................. 146
Choice of base level............................ 110 Stn M57 ........................................ 129 Talcahuanoffumbes ........................ 146
Collecting procedure and Comparison between the two areas ........... 130
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The Montemar area.... ..... ...... ..... ..... 146

its shortcomings... ... ...... ..................
The species, numbers and names ... ... .....
The species to be considered ................
The figures of zonation patterns ... .........
The stations and their zonation patterns .....
The Seno Reloncavi area .....................
StnM22 ........................................
Stn M91 ........................................
Sin M90 ........................................
StnM59 ........................................
Stn M37 ........................................
Stn M139 ...... ........... .....................
Stn M82 ........................................
Stn M31 ........................................
INTRODUCTION
In the beginning of this century PETERSEN in a se-
ries of publications (1913 and others) described a
number of soft-botton 'animal communities' in Dan-
ish waters, each being characteristic of a certain
bottom-type. Similar communities, in which the
dominating species belonged to the same or related
genera as those characterizing the communities of
Petersen, were later described from comparable
bottom-types in other areas of the Northern hemi-
sphere and from tropical West Africa, communities
for which THORSON (1951) introduced the term 'par-
allel communities'. The main purpose of the Lund
University Chile Expedition 1948-1949 (LUCE)
was to find out if bottom communities similar to
those described from northern seas also existed in
an area with comparable topographic, climatologi-
cal, and hydrographic conditions in the Southern
hemisphere and even in another ocean.
The only area in the southern oceans, compar-
able with the Nordic seas, is southern Chile. Both
areas have been formed by ice, have extensive ar-
chipelagos, deep fjords, and fairly similar climate
and hydrography. Puerto Montt at the northern
shore of Seno Reloncavl, the northernmost part of
the Chilean archipelago, was chosen as base for the
110 Number of species.......... ........ ...... ...... 130 Coquimbo ..................................... 147
111 Species composition ........................... 130 Antofagasta ............ ................... .... 147
112 Differences in vertical distribution ...... _.. 132 Iquique .......... .... ......... .................. 147
113 Zonation patterns ................................. 134 Arica ............................................ 147
114 Actual vertical distribution ................... 134 Comparison between the Seno Relon·
115 Types of vertical distribution ................ 135 cavilAncud areas and
115 Vertical distribution ofrelated species .... 137 other parts of Chile ......................... 148
116 Comparison with the zonation patterns Acknowledgements ............................... 149
118 described by Alveal & Romo ..... ...... ..... 139 References .......................................... 150
119 Species composition ........................... 139 Appendices ....................... .................. 151
120 Zonation patterns .............................. 140
123 Species composition and zonation patterns
124 in other parts of Chile ... ....... ..... ..... ..... 143
125
expedition. This town is situated at 41°29' S, a lati-
tude which, in the Northern hemisphere, corre-
sponds to that of the Mediterranean, but the cold
Humboldt Current, running northwards along the
Chilean coast, lowers the temperature of both air
and water to such a degree that the climatological
and hydrographic conditions in the northern part
of the archipelago rather correspond to those in
southwestern Norway some 20 degrees farther from
the Equator. The summer surface temperatures are
about the same in both areas, whereas the winter
temperatures are about 5° C higher in Seno Reclon-
cavl, resulting in a smaller yearly temperature am-
plitude there than in southwestern Norway. It thus
would have been preferable to choose a base a
little farther south, but Puerto Montt was the only
part of the archipelago, where it was possible to
rent a suitable vessel, get laboratory facilities, diffe-
rent kinds of supplies, and accommodation. As it
turned out, the small temperature differences didn't
matter. The same animals and algae were found in
great parts of the northern archipelago.
It was risky, of course, to organize an expedition
to an area, from which no hydrographic survey ex-
isted and where the fauna and flora were little
known. On the other hand, these very facts made
the area more attractive, for they were a guarantee

108
 puer~to
Montt
41° - "37
30'
5 c'il
o OA'

~
90 "Q,
'::, Seno "
" Reloncavf

"",J!~: o, 25 km

Ch itoe ,Golfo de Ancud

Fig, 1. Map of the area, showing the location of the 12 stations investigated and the 4 of
ALVEAL & ROMO (1977), A = Piedra Azul, C = Bahia Chincui, H = Bahia Huelmo, and
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Q =Punta Quillaipe,

that valuable results would be obtained even if the
main object of the expedition couldn't be realized.
In fact the first description of the hydrography of
the area inside the big island of Chiloe, published
in LUCE Report no. 1 (BRATISTROM & DAHL 1952),
is the result of the expedition, and its collections
have been described in LUCE reports 2-49. These
papers are mainly taxonomic and contain descrip-
tions of more than 250 new species and many spe-
cies new to the area.
One other purpose of LUCE was to study rocky-
shore zonation. Since, however, the working area
was chosen with regard to the main purpose of the
expedition, the zonation studies had to be carried
out in that area. The present report no, 50 describes
the results of these studies. Because most species
were unfamiliar to me and because many could be
expected to be new to science, a great many speci-
mens of all species and from all levels on the shore
had to be collected and brought home for identifica-
tion by specialists. Accordingly, the collections be-
came very extensive, and administration, correspon-
dence with specialists, and editing the results natu-
rally was time-consuming. For this reason, many
years passed before specimens of all taxa could be
sent to specialists for identification, and many spe-
cialists needed decades to work up and describe
such large collections, which also comprised the
material from soft bottoms. In fact some animal
taxa, most important of which are the sponges, sed-
entary polychaetes, and siphonarians, have still not
been treated. Of course the description of the zona-
tion couldn't be made before at least most of the
animals and algae, which characterize the different
levels on the shore, had been identified. When fi-
nally it became possible to publish some general
results many other obligations made it necessary to
postpone publishing until now. This does not mean
that the results will be obsolete already at the date
of publication. Since very little has been done in
the area after the expedition, the results represent
new information.
MATERIAL AND METHODS
Material
The field notes, correspondence, and the main part of the
marine invertebrates collected by LUCE are kept in the
Swedish Museum of Natural History, Stockholm, with
duplicates in the Zoological Museum, Lund University.
Some duplicates are also kept in the zoological museums
of the universities of Bergen and Oslo, and duplicates of
certain taxa in the museums, where these taxa have been
worked up. The algae are kept in the Department of
Marine Botany, Goteborg University.
The stations
General collecting was made at many littoral stations, but
since zonation studies only could be carried out around
spring tide, when most of the lower shore emerged at
low-water (LW), and since such studies also were depen-
dent on calm weather and access to transportation, the
number of stations for detailed zonation studies had to
be restricted to 8 in the Seno Reloncavi area and 4 in the
Ancud area, the latter henceforth mostly referred to only
as Ancud (Fig, 1). The former were studied during sum-
mer, autumn, and winter, the latter during early autumn,
Because of the great tidal amplitude, more than 7 m in
Seno Reloncavi and 2 m in Ancud, the water rose very
fast when the tide was in, and the time available for sam-
pling lowest down, making notes, and taking care of the
samples was so short that only a fairly narrow section of
each locality could be investigated,
Of the stations in the Seno Reloncavi area M22, M37,
M59, M90, and M91 were situated in Seno Reloncavi pro-

109
 per, M31 and M82 in the long fjord Estero Reloncavi,
and M139 at the mouth of this fjord. In Ancud the sta-
tions MlO, M55, M56, and M57 were studied. The station
numbers are those used by the expedition but in this
paper only refer to that narrower part of the stations, in
which zonation was studied. Information on the localities,
biotopes, and dates of investigation as well as hydro-
graphic notes and maps are found in BRA'ITSTROM & DAHL
(1952) and zoogeographical results in BRA'ITSTROM & Jo-
HANSSEN (1983).
The choice of exact places for zonation studies was
mainly a question of accessability and working conditions,
such as steepness of the shore and wave action. Most sta-
tions in the Seno Reloncavi area were more or less slop-
ing shores with boulders and stones in sand, those in
Ancud rocks with rock pools and many crevices. An earth-
quake in 1960 (see KILIAN 1961; PARKER 1960) partly
changed the shores at least in Ancud, and it is not possi-
ble now to find the stations exactly as they were before
the earthquake.
Choice of base level
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To be able to compare zonation patterns at different sta-
tions all measurements (in this paper expressed in cm if
not otherwise stated) must be referred to a common and
easily recognizable sea level. For the present investigation
the zero of the Tide tables (ARMADA DE CHILE 1948, 1949)
was chosen as being the most practical one. This level is
the mean of the lowest low-waters at new and full moon
('nivel de las bajamares medias de sicigias'), in Puerto
Montt being 360 and in Ancud 113 cm below mean sea
level (MSL) at the respective places. Because the zero
of the tide gauge in Puerto Montt was 94 cm below that
of the tide tables, 94 cm had to be subtracted from the
readings of the gauge to find the height of the actual sea
levels above or below the zero of the tables.
As the position of these levels at the stations was un-
known at the time of investigation, the measurements,
with few exceptions, started at the day's morning LW,
when the sea level remained stable for a short time, using
this level as a preliminary zero level for the measure-
ments of the vertical distribution of the organisms. For
the stations in Seno Reloncavi the level of this prelimi-
nary zero above or below the zero of the tide tables could
be found by subtracting 94 cm from the height of the sea
level at LW, as registered by the tide gauge in Puerto
Montt. Then all measurements could be recalculated in
relation to the zero of the tide tables. Only for M37 was
it impossible to find the exact level on the shore of this
zero, because the tide gauge in Puerto Montt was out of
function the days of investigation. Since, however, the
vertical zonation at M37 (Fig. 9) seems not to deviate from
that at the other stations, it is believed that the zero level
used at M37 is not significantly different from that used
at the other stations.
The registrations of the sea level in Puerto Montt are
of course only valid for this place. When they neverthe-
less have been considered representative, and used, for the
stations in Seno Reloncavi, this is because the difference
between the level of the sea at LWand HW measured
at these stations proved to be practically the same as that
registered simultaneously by the gauge in Puerto Montt.
However, at M31 innermost in Estero Reloncavi LW oc-
curred about 30 minutes earlier than the predicated hour
for LW in Puerto Montt, but because the sea level is fair-
ly stable during the first half hour after LW, the differ-
ence in time for LW between Puerto Montt and M31 will
probably be almost negligible when it comes to deciding
110
the zero level. This and other levels at the two fjord
stations, M31 and M82, must, however, be considered a
little uncertain and comparisons with the other stations
will be difficult to make.
In Ancud the tidal amplitude is only 226 cm. There was
no tide gauge in Ancud and unfortunately the actual sea
levels could not be compared with those in Puerto Montt,
because the tides in Ancud belong to another tidal re-
gime. Therefore the reference harbour for Ancud is not
Puerto Montt but Valparaiso. Finding the zero of the tide
tables in Ancud by comparing the sea levels there with
the measurements of the tide gauge in Valparaiso, almost
9 latitude degrees farther north, is of course not possible.
It is hardly likely that the actual sea levels should be the
same in two areas so far apart, because air pressure and
wind direction, factors which influence the sea levels, will
certainly not be the same in the 'two places at the time
of LW. However, as in Seno Reloncavi the actual sea
level at LW the days of investigation was used as zero for
the measurements, but the preliminary measurements
could not be recalculated and related exactly to the zero
of the tide tables. This is important to bear in mind when
the stations in the two areas are to be compared. The
exact levels may thus not be quite correct, but the order
in which the species occur on the shore and the extent
of their areas of vertical distribution are correct and com-
parable.
Collecting procedure and its shortcomings
A 2 m high meter with marks for every 5 cm was placed
on the shore with the zero at the morning LW-Ieve1 (Fig.
2). Theoretically it would be possible to find the exact
level on the shore of every organism by reading the me-
ter, when the specimens were in the water-line, but low-
est down where the water rose very fast when the sea
flooded and where the number of species and specimens
was very large, time was too short to permit using this
easy procedure for every single specimen, so another a
little less accurate way of collecting and measuring became
necessary.
By wading to and fro in the section to be studied, col-
lecting was made exactly along the rising water level while
observations on abundance, biology etc. were dictated to
an assistarit on the shore, who made notes and took care
of the samples. Collecting continued in this way until the
meter showed that the water had risen for example 10 cm.
All specimens collected between 0 and 10 cm were placed
in a jar labelled with the station number and 'G-1O cm'.
The width of the levels sampled could vary from station
to station. Collecting then continued as before until the
sea level had reached for instance the 20-cm mark on the
meter, and the specimens collected between 10 and 20 cm
were placed in another jar, labelled '10-20 cm', and so
on. In the higher parts of the shore, where the organisms
were fewer and the water rose slower, the width of the
collecting zones was increased, but the time available now
permitted making detailed notes on the occurrence of the
respective species within each level sampled. A look at
the meter always made it possible to state the exact lev-
els of the specimens collected.
When the water had risen to 2 m, the meter was re-
moved and placed with zero at the 2-m level, and collect-
ing could continue the next 2 m, after which the meter,
in Seno Reloncavi, had to be moved again, first to the
4- and then to the 6-m level. In this way it was possible,
without knowing the species names when collecting, to
state the distribution limits of the respective species in
relation to the preliminary zero, with an error usually not
 Fig. 2. The meter at Stn M56,
Punta Corona.
greater than 5-10 cm, which is insignificant on a shore
where the tidal amplitude may exceed 7 m.
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Owing to the limited time for studies lowest down many
rare species and small animals were certainly overlooked
there, at least such hiding in crevices or under stones and
algae, and because the sections were narrow also some
species common in the area could be missing in a section,
thus giving a wrong impression of the composition of the
fauna and flora at a station. Also small differences in the
biotope can be responsible for the absence of a species
in the section studied or at a certain level there. Usually,
however, species not observed at one station were found
at a nearby one (see Fig. 22), thus proving that they occur-
red in the area even if missing locally.
Since L W often occurred early in the morning there
wasn't always enough light to permit good observations
and sampling lowest down, and since L W on the days of
investigation at some stations could be one metre or more
above the zero of the tide tables, some infralittoral spe-
cies, which normally go a little way up in the littoral,
certainly occurred though they are missing in our collec-
tions, and the lower limit of some littoral species could
not be observed. Because the sea lowest down rose very
fast there was also little time to collect calcareous algae
and such sessile animals, which had to be collected by
using chisel and hammer.
The lichen occurring high up on the shore and the ter-
restrial plants and animals were only rarely collected and
few have been determined. The fact that for instance the
Verrucaria sp. found at three of the four stations in Seno
Reloncavi, studied by ALVEAL & ROMO (1977), was not
recorded at my nearby stations does not mean that. Verru-
caria species were absent there, only that none was col-
lected. By the way, some of my undetermined' lichen may
be a Verruca ria.
In the preceding paragraphs several shortcomings have
been mentioned. When for instance the lowest part of the
shore wasn't always sufficiently studied, when the time
was too short to permit chopping pieces of rocks and
boulders with fixed algae and animals, when specimens
couldn't be sampled because they were submerged, be-
":ause the water was turbid or the light insufficient, and
hen. high-level tithen and terrestrial organisms were"only'
)ccaslOnally sampled, why weren't the studies completed
another day?
First it was" no idea to return another day if the lowest
part of the shore also then would be submerged even at
L W, if L W occurred so early that darkness made detailed
studies and collecting difficult, or if the sea was too chop-
py to permit collecting at the waterline.
Secondly, though some stations could be reached with
our small, open motor boat, its use was dependent on the
weather conditions, and when the sea had reached its high-
est level and time had come to collect lichen and other
high-level organisms, the boat often had to return to the
base before it became too dark.
Thirdly, to reach the stations farthest away we had to
use other means of transportation. For this the only ves-
sel available in the area was a trawler, which could be
rented only a few days each month and not always those,
when conditions for shore studies were favourable. The
days the trawler was at our disposal it also had to be used
for our other studies, often far away from the littoral sta-
tions. It is hoped, however, that the information availa-
ble, though not always complete, will be of value, especial-
ly for comparison with conditions after the earthquake and
with zonation in other areas.
The species, numbers and names
Hitherto 212 marine species, 135 animals and 77 algae,
from the sections studied have been identified, Chaeta-
pterus and Phytia to genus only. One insect and some ter-
restrial plants have also been identified. The 212 marine
species are enumerated in Appendix 1, and Table 1 sum-
marizes the number of marine species of the different taxa
found at the respective stations, the Turbellaria, Nemer-
tini, Sipunculida, Isopoda, Amphipoda, Stomatopoda, and
Pycnogonida excepted. The animals found in rock pools
and the Polychaeta and Bivalvia occasionally found in
sand are outside the scope of this work, but because their
occurrence in the area is little known and any information
about them may be of interest in other connections, they
have been enumerated in Appendix 2A-C. Notes made
in the field on the biology and of colours of living ani-
mals will not be used in this paper, but as they may be
of interest to others, they have been collected in Appen-
dix 3:""
In addition to the determined species at least 30 others
have still not been identified. Such taxa as sponges, spir-
111
 Table 1. Number of identified animals (except Turbellaria, Nemertini, spirorbids Sipunculida, Isopoda, Stomatopoda,
Pycnogonida, and siphonarians) and algae found at the 12 stations. No. = total number of species.
Taxon No. M22 M91 M90 M59 M37 M139 M82 M31 MlO M55 M56 M57
Anthoza 10 4 3 1 1 4 0 0 0 5 3 5 2
Chaetopterus 1 0 1 1 1 0 0 0 0 0 0 0 0
Cirripedia 6 5 6 3 4 4 4 1 1 2 2 1 3
Decapoda 22 9 10 10 11 10 3 2 1 2 2 7 4
Polyplacophora 8 7 4 5 3 5 1 0 0 4 3 6 4
Prosobranchia 24 13 10 10 13 11 5 0 0 7 8 12 7
Opisthobranchia 8 2 2 0 0 2 0 0 0 2 1 3 0
Pulmonata 2 0 0 0 0 0 0 1 1 0 1 0 0
Bivalvia 7 5 5 6 5 5 3 1 0 5 1 5 2
Echinodermata 15 7 12 1 4 4 4 0 0 0 0 1 0
Ascidiacea 7 5 4 2 1 1 0 0 0 6 2 4 1
Sum animal species 110 57 57 39 43 46 20 5 3 33 23 44 23
Chlorophyta 14 2 5 5 4 4 5 4 1 3 5 4 2
Phaeophyta 17 0 4 2 1 4 3 0 0 1 4 6 3
Cyanophyta 4 0 1 0 1 1 0 1 1 0 1 0 0
Rhodophyta 42 9 12 13 11 18 6 2 2 7 13 18 13
Sum algal species 77 11 22 20 17 27 14 7 4 11 23 28 18
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Sum species 187 68 79 59 60 73 34 12 7 44 46 72 41

orbids, amphipods, siphonarians, bryozoans, and lichen
often comprise more than one species, but they are treated
as one species each, when numbers are mentioned, except
where it was obvious that more than one species was pre-
sent. It should also be observed that for instance a crust-
shaped bryozoan found at one station mayor may not
be identical with one so described from another. To the
undetermined species just mentioned come microscopical
algae and early growth-stages of algae and single speci-
mens of animals and algae, which for different reasons
have been determined only to genus and which may be
identical with determined species from the same locality
and therefore have not been counted. The total number
of species found at the 12 stations can thus not be stated
exactly but is not less than about 300.
All the identified animals, the Prosobranchia and Pul-
monata excepted, and the algae have been described in
the LUCE reports, and the species names used in these
are used in this paper, too, except for Nassarius gayii
(Nassa gayii), Nassarius dentiferus (Alectrion dentifera),
and Chitina bulloides (Chitina robustior). There are some
discrepancies between the LUCE reports and this paper
regarding which species were found at the stations, be-
cause the 'stations' in this paper only represent that nar-
rower part of the LUCE stations, in which zonation stud-
ies were carried out. Thus species found outside this sec-
tion and, as mentioned, also in rock pools and sand, are
omitted though they are mentioned from the same station
in the LUCE reports. On the other hand some authors,
when publishing their results, forgot records from some
stations, from which they in fact had material identified
by themselves. These species have been considered here
only if the field notes contain information on their levels
on the shore and show that they have been collected in
the section studied for zonation purposes.
In Appendix 1 and those other tables, in which species
not contained in Appendix 1 are enumerated, the species
have been entered with their full scientific names, fol-
lowed by the author names. In the text, however, author
names to these species have been left out, whereas author
names are mentioned there for species not enumerated in
any table. For two species, cited from other publications,
the author names are not known.
Most genera are represented in the material by a single
species. For the sake of brevity these species will, in the
description of the stations, be referred to only by
the generic name. Appendix 1 shows which these species
are. On the other hand both the generic and species
names will be used, in these descriptions, for all species
belonging to genera represented there by more than one
species. If, however, as is often the case, one of the
species of such genera is common at a station and the
other(s) present rare and only the common species is
mentioned, this will be referred to by the generic name
alone. These species are: Petrolisthes (laevigatus), Acan-
thocyclus (albatrossis), Nacella (magellancia), Patelloida
(cecitiana), Littorina (araucana), Enteromorpha (bulbosa),
Viva (Iactuca), Getidium (pseudointricatum), and Litho-
thamnion (pauciporosum).
Finally, when in a paragraph only one species of a
genus, be it common or rare, is mentioned with full name,
only the generic name will be given if the species, and
no other species of that genus, is mentioned again in the
same or nearest following paragraphs. In other text than
the descriptions of the stations the full species names will
be used except for cases such as mentioned in the preced-
ing sentence.
The species to be considered
In zonation studies the question always arises, which spe-
cies are best suited to characterize the different levels on
the shore. There will always be a subjective component
in the decision of which species to be discussed, but ses-
sile animals and algae are obvious candidates, since they
cannot move. Also hemisessile animals, such as chi tons
and limpets, which move around when the shore is sub-
merged but return to a fixed place, when it is emerged, are
character or key species of this level and of the relatively
narrow area, in which they are moving around grazing,
when the shore is submerged. Also those motile animals,
both grazers and predators which, when submerged, make
restricted foraging excursions are key species of these le-
vels and not only of the narrower levels, between which
they seek shelter at ebb, under algae, at the base of boul-
ders (Fig. 3) or in crevices and holes, places which keep

112
 Fig. 3. Tegula atra (LESSON) at
the base of a big boulder (M37)
at ebb-tide. When the sea flood-
ed Tegula was moving around
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all over the boulder.
moist or where some water is left, when the shore be-
comes emerged. Such species are, among others, the' pag-
urids, Halicarcinus planatus, several snails, and the echino-
derms, which always are found lowest down, and Cyclo-
grapsus cinereus, mainly found high up. On the other
hand those motile animals which, like Petrolisthes laelli-
gatus, Acanthocyclus albatrossis, and Hemigrapsus crenul-
atus, roam over most of the intertidal area, when the shore
is submerged and which also are found sheltering within
a fairly wide area, when the shore is emerged, cannot be
used as character species for a certain level. They are
characteristic of a great part of the intertidal area. Figs
22 and 24 show examples of both restricted and wide verti-
cal distribution.
Other requirements for character species are that they
are conspicuous, more or less common, and also easy to
determine. Small species difficult to find or identify and
rare species are less suited. Thus only the species most
suitable for characterizing a station or a level there, will
be mentioned in the description of the stations or dis-
cussed in other connections. This of course does not im-
ply that other Doth common or less common species can-
not be equally or even more characteristic, only that they
for different reasons are less suited as character species.
However, even those not mentioned or figured are enu-
merated in Appendix 1 with information on the levels
which they were occupying at the different stations. Unde-
termined species have usually not been considered in the
descriptions and in the discussion, since they cannot be
used for comparison with other areas.
Some similar algae, which grew intertangllid and there-
fore, when collected, were thought to represent one spe-
cies, have later been found to be two or more. This often
makes it difficult to know to which of the species the
notes on for instance abundance refer. Samples of such
species have been left out of the discussion, but often the
same species were found alone in other samples from the
same levels, so that their vertical distribution nevertheless
could be stated.
Some species are probably more common than notes
and collections indicate and as is mentioned in the text
and shown in the figures of vertical distribution. Thus for
example small actinians and Brachidontes purpuratus were
often fixed on the underside of overhanging rocks and
boulders or on stones and rocks, the bases of which now
and then were covered by sand. Such specimens have
certainly been overlooked sometimes. Lasaea petitiana,
which was living well hidden among barnacles and mus-
sels, is also no doubt underrepresented in notes and collec-
tions. In fact many specimens of this small bivalve were
not observed in the field but found later in samples of the
larger species.
The figures of zonation patterns
Figs 4, 5, 7-9, 13-15, 17-19 and 21 illustrate schematically
the vertical distribution at the 12 stations of the more
common, conspicuous, and important animals and plants
occurring there. The choice of species to be figured is of
course somewhat subjective, based not only on field notes
on abundance but also on a general impression of their
importance for characterizing a level. When a species,
which was rare at a station, exceptionally has been inclu-
ded in the figure of that station, it is because it could be
of interest to compare its level there with that at a station
where it is common, or because that species was not found
at any other station and therefore was characteristic of
that where it was found.
Unidentified species, even if characteristic of a level,
have usually not been included in the figures, as for ex-
ample sponges and hyaline ?polychaete tubes. Another
such group well represented at most stations, is the Am-
phipoda, but since they have still not been determined
they cannot be discussed. I will confine myself to mention-
ing that a luminescent species was common lowest down
at M22 and that one or more jumping species were very
common upmost in the littoral at M91 and in the lowest
part of the supralittoral at M22, M59, and M37. Other
species were common at most stations upwards to the
upper limit of the littoral. A systematic study of the
Amphipoda probably will reveal a more intricate pattern.
When other undetermined species exceptionally are fig-
ured, for instance spirorbids, siphonarians, and lichen, and
one finds differences in vertical distribution at different
stations, this can, but need not, be because different spe-
cies are involved.
113
 The vertical distribution of the species is illustrated by
black bars, the width of which shows if the species are
abundant, common, fairly common or rare. Small gaps in
the distribution of a species, probably the results of over-
sight or local differences in the substrate, have not been
shown in the bars. Larger gaps in vertical distribution
have, however, been marked. Ifthe field notes occasional-
ly only state that a species was taken between for instance
50 and 150 or in 'the Elminius belt' without further in-
formation, bars with alternating black and white bands
have been used to emphasize that the exact level at which
the species was living is not known, only that it was taken
somewhere in the area indicated. As for the black bars
the width of the banded ones shows how abundant the
species was. On the other hand, where the levels were
specified but the abundance not recorded, white bars have
been used, all of the same width.
Because the conception of abundance is subjective and
not based on exact counting of the number of specimens
per area unit or on measurements of the areas covered,
it would perhaps have been more natural to give all bars
the same width, thus only indicating at which levels the
species occurred, but then important information on abun-
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dance and preferred levels would have been lost. Even a
subjedve impression of this and thus of the role a speci-
es plays in the zonation pattern is better than a mere indi-
cation of presence.
Where the upper and lower limits, for various reasons,
couldn't be stated exactly, arrows at the ends of the bars
indicate that the species probably occur lower down or
higher up than the bars show. In some cases the lower
limits observed don't represent the real lower limits of
distribution but the level below which observation and
sampling were not possible the day of investigation. In
such cases text on the figures mentions the fact and no
arrows have been used, unless records, without measure-
ments of levels, showed that the species really occurred
lower down.
The bars have horizontal upper and lower ends and a
step-like presentation of abundance, a picture which natu-
rally is artificial. Most species usually begin with few speci-
mens, increasing gradually in abundance upwards, then
decreasing again to the upper limit. The bars have got
this step-like shape because the notes on abundance refer
to the mean abundance within each level sampled. How-
ever, abruptly ending bars or parts of these give a correct
picture of the abundance, 1) when the abundance of a
species was stated to be the same in the whole area of
distribution or throughout a level studied, and 2) if for
example a species occurred all over a boulder but not on
other boulders lower down or higher up at not overlap-
ping levels. In the latter case species might have had a
V':ider, uninterrupted vertical distribution if rocks or over-
lapping boulders had been found at the locality studied.
THE STATIONS AND THEIR ZONATION
PATTERNS
At localities studied by me in Colombia and Pana-
ma three regions were distinguished on the shore,
the sublittoral (= infralittoral), the littoral, and the
supralittoral or maritime zone, each characterized
by certain assemblages of animals and plants
(BRATISTROM 1980, 1985). In both areas the littoral
was divided in the eulittoral, with two or three
often well-defined zones, and the littoral margin
or spray zone, practically exclusively populated by
114
littorinids, especially Littorina ziczac (GMELlN) and
L. angustior (MORCH). These two species were also
found in the eulittoral but were rare there. It must
be remarked in advance that the same three regions
could be distinguished at the Chilean stations, and
that also in southern Chile the littoral could be di-
vided in three more or less distinct zones, the upper-
most one tolerably well corresponding to the litto-
ral margin and being characterized by Littorina
araucana, the common Littorina of the area. This
species was, however, not uncommon also lower
down and it was far from alone in the upper zone
(see the diagrams of zonation and Fig. 22), which
thus had a somewhat other character than the lit-
toral margin in Colombia and Panama. Accordingly
I will not name the uppermost littoral zone the lit-
toral margin but prefer to name the three littoral
zones simply as zones 1 to 3 (see below).
Many subtidaVinfralittoral species continue more
or less high up in the intertidal area, in what STE-
PHENSON (1939) and STEPHENSON & STEPHENSON
(1949) called the sub- or infralittoral fringe, a term
I will use here, or even higher up in the true lit-
toral. Their levels, as other limits on the shore, are
not the results only of the tidal levels but vary from
place to place depending on local conditions such
as substrate, shore topography, light, hiding places,
exposure, and competition. There thus will be a
more or less wide area in which infralittoral and
littoral species occur together. Similar areas of over-
lapping will be found where littoral and terrestrial
species meet and also at the zone limits in the lit-
toral.
The occurrence of these areas of overlapping
makes it difficult to decide where to draw limits
between regions and zones. Sharp limits are thus
rarely found and those decided upon may vary from
station to station and even within areas as small as
the stations studied. When placing the limits, I first
have considered those species, which play an impor-
tant role on the shore of the respective stations or
such which are rare but unique for the station or
level in question.
Based on species of the categories mentioned I
have tried to find out at which levels the greatest
changes in species composition and abundance take
place, but no less important for the decision has
been the visual impression of the zonation pattern
I got at the site. There thus always will be a subjec-
tive element present when to decide where to place
a limit. This one must not forget, especially not
when comparison is made with other places.
As a result of these considerations the following
five regions and zones (see Fig. 23) have been dis-
tinguished at the stations in Chile:
 The infralittoral fringe, the lowest part of the
intertidal area, which is characterized mainly by
infralittoral species.
The littoral region with three zones, the lowest
(1) being a transition zone in which still many in-
fralittoral species occur together with littoral ones,
the middle (2) a zone with mainly littoral species,
and the upper (3), a zone where littoral and supra-
littoral species meet.
The supralittoral region with mainly supralittoral
and some terrestrial species.
However, naming the levels and drawing limits
between them is not the chief concern. It is the
level on the shore at which the species occur that
is of interest, but distinguishing zones and giving
them names makes it easier to describe the zonation
patterns.
For the sake of brevity usually no reference will
be made, in the following description of the sta-
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tions, to Table 1, nor to Appendices 1 and 2. Spe-
cies mentioned as occurring in a region or zone and
marked with an asterisk in front of or after the
name, also occur in the region/zone below or above
or in both, and will be mentioned there, too, if
they are considered characteristic, though not if
their occurrence there is restricted only to a few cm
nearest to the limits of the region/zone.
THE SENO RELONCA vi AREA
The stations (see Fig. 1) will be treated from west
to east, thus M22, M91, M90, M59, M37, M139,
M82, and M31, that is from the supposed most
saline M22 to M31 at the head of Estero Relon-
cav!, where the water is practically fresh.
Stn M22, Golfo de Ancud, Isla Quenu, Punta
Pinto, western side, 41°49'15" S, 73°10'15" W, 16
December 1948 and 11 May 1949 (Fig. 4).
Slightly sloping, fairly exposed sandy shore with scat-
tered stones, above 80 with boulders, stones, and higher
up pebbles, and just below MSL some ledges of conglom-
erate. There was much sand between the boulders and
stones, which often were naked at the base, probably as
a result of sand scouring.
Whilst the fauna was diverse With 68 identified species
and a dozen undetermined ones, only 11 species of algae
were found, all but 2 Rhodophyta. There was especially
many Polyplacophora species. Because the hard substrate
(boulders and stones) was discontinuous the animals and
algae on one boulder were separated from those on other
boulders by areas with pebbles and sand, and one didn't
find the well-defined belts of organisms, which are often
seen on continuous rock surfaces, for even the largest
boulders had relatively small surfaces. Therefore not all
species were present on each or they were represented
by few specimens only, and the pebbles and the conglom-
erate had few species. Seen from a distance, however, the
single boulders merged and appeared as an unbroken rock
surface on which a zonation pattern became visible, espe-
cially a distinct change in colour at about 210, indicating
a major change in species composition. Below 210 the
shore appeared dark, above lighter and more brownish.
The light colour was caused by two species of Chtham-
alus, and the brown by early stages of green algae, prob-
ably Enteromorpha and Viva, growing on many of the
animals.
The infralittoral fringe
It was fairly dark at the time of L Wand the shore then
was submerged up to about 110. Because of that only few
species could be sampled below this level. Some were,
however, observed on stones down to about 80, but even
this level is not their normal lower limit, for most of the
hard-bottom species occurred lower down at other locali-
ties, where hard bottoms were found below 80.
It is not easy to decide where to place the upper limit
of the fringe, for there was a gradual decrease upwards
in the number of species. Fig. 4 shows, however, that
many had common upper limits at about 210, 240/250, and
also at 300/310, indicating changes in living conditions at
these levels. I have chosen to place the limit at 210, not
only because many infralittoral species stopped or became
scarce there but because some littoral species had their
lower limit at about this level, where also the afore-
mentioned colour change was observed. Between 210 and
300/310 there were so many littoral species that it seems
natural to treat this area as part of the littoral region.
Characteristic of the infralittoral fringe, as here delim-
ited, were a yellow crust-shaped sponge, hyaline ?poly-
chaete tubes", big clusters of Balanus psittacus, B. floscu-
Ius", sessile molluscs such as Tonicia atrata', Nacella" ,
Crepipatella" , Aulacomya" , and Ostrea" , crust-shaped
soft, white bryozoans, ascidians as Corella and Pyura" ,
algae as Viva' and Crodelia, and the motile Pagurus ed-
wardsi, Petrolisthes angulosus, Tegula", Nassarius gayii" ,
and Anasterias" and other echinoderms. Pagurus villosus"
and a few less common littoral species had their lower
limit upmost in the fringe.
The littoral region
The lower limit at 210 was less distinct than the upper
at 540, where many of the most common species stopped
or became less abundant, and where yellow lichen began
to appear. The region was populated both by littoral spe-
cies and a great many of those found in the infralittoral
fringe. Zone limits can be placed at 300/310 and 390.
There were also less distinct limits in distribution and
abundance at 240/250 and 440.
Zone 1, 210-310. Chthamalus scabrosus", "Pagurus vil-
losus, "Nucella", "Nassarius gayii, and' Viva" dominated
or were common in the lower part, "Parantheopsis, " Patel-
loida", Monodonta" , and Chondrus in the upper. 'Hyaline
tubes were common and "Balanus flosculus" dominated
throughout. Parantheopsis, Pagurus villosus, *Crepipatel-
la, "Nassarius gayii, Chondrus, and *Anasterias stopped
about halfway or at the upper limit. C. scabrosus, Acantho-
cyclus*, and Littorina" had their lower limit at or near
the lower zone limit, Chthamalus cirratus', Monodonta,
Lasaea" , and Chondrus at different levels above this.
Zone 2, 310-390. The narrow belt of conglomerate be-
gan at the lower limit. The gap in distribution of many
species in this belt indicates that this type of substrate is
avoided. This makes it probable that some of the species
not found above 310 would have occurred higher up and
that some of those beginning to appear above the con-
115

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INVESTIGATIONS BELOW 110, WA}ER_ICOVERED
SAND BELOW 80
...J
~
u.
z
Fig. 4. Zonation at Stn M22, Isla Quenu.
glomerate perhaps would have occurred lower down, had
the substrate been boulders or rock.
The two *Chthamalus species became belt-forming at
the lower limit. Petrolisthes laevigatus, from 340, *Tegula,
and some less common species stopped at the upper limit,
where *Brachidontes*, * Porphyra*, and *Gelidium* became
abundant. * Balanus flosculus' dominated in this zone,
too, together with *Chthamalus cirratus*, *Littorina*,
and, upmost, *Lasaea*. *Monodonta* was common, in
the upper part also Petrolisthes, * Patelloida*, and Tegula.
The less common Mytilus* had its lower limit at 350.
'Mytilus, Brachidontes, Porphyra, and Gelidium were
common on the landward side of larger stones in this and
next zone, whereas Chthamalus cirratus dominated on the
seaward side and the top. Brachidontes also occurred bur-
rowed in the sand at the base of stones.
Zone 3, 39(}"540. *Chthamalus cirratus, *Monodonta*,
*Littorina* , and *Lasaea were abundant or common
throughout, *Balanus flosculus, * Patel/oida, * Brachidon-
tes, 'Ulva and 'Porphyra in the lower part. Balanus stop-
ped low down, the other species between 440 and 450,
where Gelidium became rare. Catenella and Bostrychia
hookeri were common in the upper part, from 490/500.
Siphonarians were rare in the whole zone, Cyclograpsus'
upmost.
The supralittoral region
The area above 540 had a mixture of marine and terrestri-
al species. *Cyclograpsus, *Monodonta, and *Littorina
went up to 580/590. There Enteromorpha appeared, only
116
em
I
I TI ~OT I NVESdATED
SAND
"''lArER-ICO~'ERloD.
to disappear at 610. Yellow lichen had their lower limit
at 530 and bushes became common at 660, thus in the
uppermost part of the intertidal area. They will be
drowned for a short while at EHWS or else by waves.
Stn M91, Seno Reloncavi, Ensenada de Guatral,
41°43'00" S, 73°03'15" W, 13 April 1949 (Fig. 5).
The shore consisted of boulders, stones and, partly,
small rounded pebbles in sand and was fairly well shel-
tered.
The station had 63 identified animal species, 5 less than
at M22, but twice as many algae as there, 22 species,
mostly Rhodophyta, and in addition a dozen undeter-
mined species. Most species were found on the boulders
and larger stones. All littoral cirriped species of the area
were found and the number of echinoderms was much
higher than at any other station, perhaps because investi-
gation was possible down to zero, but at M59, where the
lowest levels could be investigated, only 4 echinoderm
species were taken, whereas M22, where only the area
above 110 could be explored, had the next highest
number (7). The echinoderms didn't, however, play any
important role in the fauna.
The infralittoral fringe
As at M22 many infralittoral species had their upper
limit at 210 and some littoral their lower at 200/220,
wherefore the limit between the two regions has been
placed at 210.
 .....
Ci. em em
o
I--
I--
::::i
<l
g:
iil
6oo+--If-I-H•

.....
<l
gj 500
I--
I--
Downloaded by [UQ Library] at 18:20 01 September 2013

.....
CiLL
Z

Fig. 5. Zonation at Stn M91, Ensenada de Guatral.

Pinnixa was very common and Pseudechinus and Lox-
echinus common below 60. Several sponges, hyaline ?poly-
chaete tubes, and crust-shaped and white, soft bryozoans
were fairly common, and Chaetopterus ?variopedatus,
Pagurus villosus, and corallinaceans dominant in the lower
half. Spirorbids*, Nassarius gayii, and Pyura were abun-
dant and Tegula* common in the whole region, and
Chthamalus scabrosus* and Balanus flosculus" abundant
and Patel/oida", My tilus" , and Aulacomya common in the
upper part. Chthamalus, Balanus, Patel/oida, and Mytilus
had their lower limit at about 150.
The littoral region
The limit to the supralittoral has been placed at 600,
where some common or abundant sl'ecies stopped. Some
mos~ly rare species went up to 250, and several common
species, also found in the infralittoral fringe, had their
upper limit at 350, just below MSL. A few other species
stopped at 400 and several littoral ones had their lower
limit there. A distinct species limit was also seen at about
550. Since the greatest changes appeared at 350 and 400
the zone limits have been placed there.
. Zo~e 1, 210-350. The uppermost specimens of many
mfrahttoral species, among them the "spirorbirds, oc-
curred up to 250, where also the abun9ant 'Chondrus
stopped. Chaetangium was common and restricted to
this zone. 'Chthamalus scabrosus was abundant to 310,
'Balanus flosculus', 'Patel/oida, Monodonta*, Littorina" ,
and Nucella dominated or were common in the whole
zone, and siphonarians*, Enteromorpha', and Porphyra*
highest up. Monodonta, Littorina, Chaetangium, and
Chondrus began to appear at or near the lower limit,
siphonarians at 250, and Enteromorpha at 310.
Zone 2, 350-400, had few species. *Balanus flosculus
was common and stopped at the upper limit, *Mono-
donta" , " Littorina * , 'Enteromorpha*, and *Porphyra*
were dominant or common. Lasaea' had its lower limit
lowest down.
Zone 3, 400-600, was better defined. "Monodonta,
'Littorina, *Enteromorpha" , and Gelidium were abundant
or common throughout. "Mytilus was common lowest
down, *Petrolisthes in the middle part, Chthalamus cirr-
atus between 400 and 550, and Elminius' and Bostrychia
hookeri upmost. Less common but characteristic in part
of the zone were *Lasaea, Cyclograpsus, and Brachidon-
tes. Chthamalus, Brachidontes, and Gelidium began to
appear lowest down, Elminius and Bostrychia at 550.
Chthamalus, Petrolisthes, Mytilus, and Brachidontes
stopped in the upper half, Monodonta, Littorina, Gelid-
ium, and Bostrychia at the upper limit.

117
Downloaded by [UQ Library] at 18:20 01 September 2013
The supralittoral region
The uppermost specimens of * Elminius and 'Porphyra
stopped at 620, whereas * Enteromorpha was fairly com-
mon up to 690, from 640 together with a yellow lichen.
Juncus sp., the bamboo Chusquea quila (POlR) (Fig. 6),
and bushes began to appear where Enteromorpha stopped.
Stn M90, Seno Reloncavl, Isla Tenglo, southwestern
point, 41°31'03" S, 73°00'02" W, 12 April 1949
(Fig. 7).
The station was one of boulders and stones in sand. It
was situated near M59 but was a little more exposed.
Only 42 identified marine animals and 20 algae were
found, in addition to one insect, Phidon reticulatus (BLAN-
CHARD), some terrestrial plants, and at least 12 underter-
mined animals.
The infralittoral fringe
The distinct lower species limit at 60 is only the limit of
possible observation. The part below was submerged at
the time of investigation. Changes in species composition
and abundance occur at 160,210, and most conspicuously
at 260/280, where some abundant species have their upper
or lower limit. The limit to the littoral has been placed
at 260.
A few Chaetopterus ?variopedatus were found lowest
down. Crust-shaped yellow sponges', Chthamalus scabro-
sus, Balanus flosculus, Pagurus edwardsi, Patelloida*,
Tegula' , Nucella' ,Aulacomya, Ostrea', bryozoans, Viva'
Chaetangium*, Chondrus*, and Iridaea ciliata' were abun-
dant or common in the whole or part of the fringe.
Chthamalus, Patelloida, Viva, Chaetangium, and Chon-
drus had their lower limit there; for the other species
mentioned the lower limit could not be stated.
118
Fig. 6. Part of Stn M91, Ensenada
de Guatral. Thickets of bamboo,
Chusquea quila (POIR.), and brush-
wood reached down to the upper
limit of Enteromorpha bulbosa
(SUHR.) KUTZ.
The littoral region
The upper limit has been placed at 570, where some abun-
dant littoral species stopped or became scarce and lichen
began to appear. Distinct species and abundance limits
were found at 360, 425, 465, and 520, those at 360 and
465 being most important, and there the zone limits have
been placed.
Zone 1,260-360, had fewer species than the levels near-
est below and above. Only' Tegula*, Mondonta', 'Nucel-
la', and Enteromorpha* were abundant or common
throughout the zone, Littorina', Pyura, 'Chaetangium,
and 'Chondrus' lowest down, and Enteromorpha in trick-
ling freshwater from 280 upwards. The freshwater between
280 and 360 may be the reason why so many species
stopped or became less common at 280, some to reappear
above 360, at which level many other species had their
lower limit.
Zone 2, 360-465, had more abundant and common spe-
cies than zone 1. Chthamalus cirratus' dominated and
'Mytilus' was common in the lower part. * Monodonta' ,
'Gelidium', and Bostrychia hookeri were abundant and
'Petrolisthes and • Enteromorpha* common throughout
the zone and • Littorina' and 'Brachidontes' abundant in
the upper part. Hemigrapsus* and Cyclograpsus' appear-
ed upmost. Siphonarians, Lasaea, Scytothamnus, and Bos-
trychia were restricted to the zone.
Zone 3, 465-570, had relatively few species, but most
were common or abundant, 'Chthamalus cirratus, 'Cyclo-
grapsus*, *Monodonta', and *Gelidium in the lower half,
'Littorina* and 'Enteromorpha in the whole zone.
'Acanthocyclus, 'Mytilus, and *Brachidontes were ~are
and stopped about halfway.
 em em
800

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Downloaded by [UQ Library] at 18:20 01 September 2013

-'
..:
0::
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....
:;

w
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II II
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-65
NOT INVESTIGATED NOT INVESTIGATED

Fig. 7. Zonation at Stn M90, Isla Tenglo, southwestern point.

The supralittoral region The infralittoral fringe

Only four species from zones 2 and 3 went up in this re-
gion, none of them being common. • Hemigrapsus and
'Cyclograpsus were found lowest down, *Monodonta to
640 and • Littorina to 680. Yellow and other lichen oc-
curred from 580 upwards, thus together with the four
marine species. Chusquea quila and Fuchsia sp. were
found from 740, the level of EHWS, only 60 cm above
Littorina.
Stn M59, Seno Reloncavi, Isla Tenglo, western
point, 41°30'45" S, 73°00'13" W, 13- and 15 March
1949 (Fig. 8).
The shore was fairly exposed; slightly sloping, in the
lower part consisting of mud and sand with boulders and
stones, in the upper of hard, slippery clay alternating with
soft sandstone.
Sixty-four identified marine species were found, 47 ani-
mals and 17 algae, besides 9 undertermined species, 1 li-
chen, and 2 land plants, about the same number as at the
nearby Stn M9O. The two stations also had 57 species in
'"ommon, among them the most important ones. Many
'pecles were abundant, especially in the lowest and high-
est parts of the shore. " dance within the region were found at 335 and 410, and
There was a gradual decrease in number of species and
abundance up to 145, where many species had a common
upper limit or became less abundant. Some others continu-
ed to 220, some to 340, but the most distinct change was
that at 145, where the upper limit of the fringe has been
placed. No common littoral species had their lower limit
at 145, but a few began to appear highest up in the litto-
ral fringe.
Chaetopterus ?variopedatus, spirorbids, Balanus floscu-
Ius', Pagurus villosus, Chiton latus", Nacella magellani-
ca*, Tegula*, Crepipatella" , Nucella" , Nassarius gayii,
Aulacomya, Ostrea*, Arbacia*, Loxechinus, Pyura, Li-
thothamnion, and Crodelia were abundant in the whole
fringe or part of it. Some sponges, pinnotherids, rare chi-
tons, bryozoans, Anasterias, and Pseudechinus also charac-
terized the fringe. Lithohamnia have no doubt been over-
looked, for they were not sampled or noted above 45.
Balanus, Nacella, Nucella, and Ostrea had their lower lim-
it high up in the fringe.
The littoral region
The upper limit is at 595, where land plants had their
lower limit. Distinct changes in distribution and abun-

119
 -'
<{
a:
o
>-
>-

em
600

-'
<{
a: 500
o
>-
>-

400

MSL
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300
-'
<{
a:
o>-
>-
:::;

Ii
"-
I-' 100 100
>-
-'
<{
a:
"-
z
---o~------------------------------------ __~------__~-I~~HlI-~~.t~---­ a
Fig. 8. Zonation at Stn M59, Isla Tenglo, western point.

there the zone limits have been placed. A less distinct
change took place at 545.
Zone 1, 145-335, was mainly populated by littoral spe-
cies, three of which were abundant, 'Balanus flosculus
lowest down, Littorina' in the upper two thirds, Chthama-
Ius cirratus' upmost. 'Chiton latus, Patelloida, Monodon-
ta', 'Tegula, 'Nucella, Mytilus, Scytothamnus', Chaetang-
ium', Gelidium*, and Chondrus' were common in differ-
ent parts of the zone.
Zone 2, 335-410, was populated exclusively by species
from one or both of the neighbouring zones. 'Chthamalus
cirratus' and 'Littorina' dominated, as in zone 1, and
'Gelidium' became abundant in the upper part. 'Mono-
donta' , Lasaea', 'Scytothamnus, and Bostrychia hookeri'
were common in the whole zone or part of it.
Zone 3, 41{}-595. Four species were abundant, • Porphy-
ra and 'Gelidium to 545, 'Chthamalus cirratus' and' Lit-
torina' throughout. 'Monodonta and 'Lasaea were com-
mon lowest down, Eliminius', 'Hemigrapsus', Cyclograp-
sus', Enteromorpha*, and 'Bostrychia hookeri* in the
whole zone, and Rhizoclonium tortuosum' and R. ripari-
um' upmost.
The supralittoral region
Juncus maritimus LAM. and Samolus repens PERS. began
to appear at the lower limit. Yellow lichen were not noted
below 630. Bostrychia harveyi was found somewhere be-
tween 595 and 635. The uppermost specimens of the spe-
cies followed by an asterisk in zone 3, went up to 615 or
630/640, among Juncus, the 'algae and' Littorina being
common.
Stn M37, Seno Reioncavi, Punta Pilluco, 41°30'06" S,
72°53'57" W, 20, 29, 31 January, 17 February, 14
and 28 March 1949 (Fig. 9).
The fairly exposed shore consisted of sand, from 110
upwards with big boulders lowest down (Fig. 10) and
small stones higher up and with some beds of hard clay.
With 85 identified species, 58 animals and 27 algae, 1
lichen, j higher plants, and at least 14 undetermined spe-
cies this station was the richest of the twelve. One reason
for this high diversity is that this station was investigated
several times, which reduced the risk of overlooking rare
species. No other station had so many abundant and com-
mon species, and more species than at any other station
had a very wide vertical distribution, some as much as
4-5 m.
The infralittoral fringe
Most infralittoral species had their upper limit somewhere
between 160 and 210, the latter level being used as limit
to the littoral region. Sponges, hyaline ?polychaete tubes,
Pagurus viUosus (in shells of Nassarius dentiferus), Macro-
cystis, and Lithothamnion were found lowest down. Spir-

120
 em
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SAND BELOW -10

Fig. 9. Zonation at Stn M37, Punta Pilluco.

orbids, Pinnotheres, Crepipatella (Fig. 11), Nassarius scabrosus', Balanus flosculus*, Petrolisthes', Patelloida*,
gayii, Ostrea, Rhizoclonium tortuosum, Acrochaetium Chiton latus', Tegula*, Nuceila*, Mytilus*, Aulacomya*,
catenulatum, and Iridaea. ciliatia were found in most of the and Chaetangium fastigiatum* had their lower limit in the
fringe and restricted to this. Bunodactis*, Chthamalus fringe. All species mentioned were common or abundant.

Fig. 10. The boulder· station M37, Punta Pilluco. ."

121

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The littoral region
There was a gradual decrease in number of species up to
410, where many abundant and common ones stopped and
others began to appear. Between 315 and 340 there was
a distinct change in barnacle species. The last marine spe-
cies had their upper limit at 570, some though with stray
specimens a little higher up. An exception was Entero-
morpha, which was abundant up to 680. The upper limit
can thus be placed at 570 and zone limits at 315/340 and
410.
Zone 1, 210-315/340. The abundant *Balanus flosculus
stopped near the upper limit. *Hemigrapsus', 'Chiton
latus, and *Patelloida' were common or abundant in the
lower half, Lasaea' in the middle part, and • Bunodactis',
*Chthamalus scabrosus', • Petrolisthes', Monodonta*,
* Tegula' , • Mytilus', Brachidontes', and 'Chaetangium*
throughout the zone.
122
Fig. 11. Crepipatella dilatata
(LAMARCK) at Stn M37, Punta
Pilluco. Some species are chain-
forming.
Zone 2, 315/340-410. Most of the species in zone 1
were also found in zone 2. *Bunodactis, ·Chthamalus
scabrosus, • Mytilus, and * Chaetangium were as common
as in the zone below and went to the upper limit. Other
abundant or common species from zone 1 or the infralit-
toral fringe, such as *Petrolisthes', *Monodonta', * Tegula*,
and * Brachidontes' passed the limit to zone 3. Chtham-
alus cirratus*, Littorina', and Porphyra' had lower limit
in the zone. A 'peje sapo', Sicyaces? sp., was found on a
boulder at about 385, sheltering its fry (Fig. 12).
Zone 3, 410-570. In spite of the high level the zone
had many abundant species, • Tegula, *Porphyra, and
Bostrychia hookeri in the lower part, * Chthamalus cirra-
tus, • Littorina, and Bostrychia harveyi' in the upper, and
*Petrolisthes, *Cyclograpsus', *Monodonta*, Enteromor-
pha', and Catenella* throughout. • Hemigrapsus* was
fairly common to rare in the whole zone. Lichen' and
Fig. 12. Fry of a 'peje sapo' (Si-
cyaces? sp.) on a boulder at Stn
M37, Punta Pilluco, 25 cm above
MSL (+385 cm).

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NO SAMPliNG. FEW OBSERVATIONS BELOW 145
Fig. 13. Zonation at Stn M139,
EI Milagro, at the mouth of
Estero Reloncavi.
Leptocarpus chilensis (MAST) had their lower limit in the
zone, at 520/530, Samolus repens' and Salicomia peru via-
na* H.B.K. var. doeringii(LoR. & NIED.)at 550, thus
occurring together with the highest specimens of marine
species.
The supralittoral region
The species in zone 3, marked with an asterisk after the
name, only just crossed the limit to the supralittoral.
Only • Enteromorpha went higher and was abundant at
least to 680. Lyngbya aesturaii was taken somewhere be-
tween 570 and 610, perhaps also below and above. Thus
even in this region 'crabs, • Monodonta, and a few "algae
occurred together with land plants.
Stn M139, mouth of Estero Reloncavi, at El Mil-
agro, 41°42'10" S, 72°39'30" W, 14 July 1949 (Fig.
13).
The station was exposed with partly vertical and partly
sloping rocks. The differences in slope seemed to have
little influence on the vertical distribution of the species,
many of which occupied the same levels irrespective of
the degree of slope. It thus seems that the waves rolling
up on the sloping cliffs don't press the limits more up-
wards than do splash and spray on the vertical rocks.
Situated at the mouth of Estero Reloncav! the station
at least now and then must be covered with brackish sur-
face water from the fjord. The day after the shore studies
the surface salinity at Stn H22 just outside M139 was as
high as 26.7 %0, but on 30 March the salinity in the sur-
face at Stn H19, near Isla Poe, 15' inside the mouth, was
only 16 %0 and on 1 April 10.2 %0, which makes it proba-
ble that the salinity at M139 those days must have been
much lower than on 15 July.
em
Only 36 determined species were found, 22 animals and
14 algae, and among others one lichen. These low num-
bers, about half of the mean at the stations in Seno Relon-
cav! proper, are probably not the result of lowered salini-
ty alone, for all stations in the area are exposed to much
rainwater when emerged. The reason for the low num-
bers at M139 is probably partly the working conditions.
We had only a few hours at our disposal and when col-
lecting could start the sea level was at 145. For this
reason species usually living low down couldn't be col-
lected. However, the species found show a distinct zona-
tion pattern with differently coloured belts, and the distri-
bution of the common species does not differ significantly
from that at the other stations.
The only fairly well represented higher taxon was the
Cirripedia with 4 of the 6 species found in the area. No
Anthozoa and Ascidiacea were recorded, and there were
few decapods and only one chiton and one opisthobranch.
The infralittoral fringe
The upper limit at 195 is where typical infralittoral species
had their upper and many littoral their lower limit. Balan-
us flosculus" was abundant, Sphacelaria* and Macrocystis"
common, and Acanthocyclus", Plaxiphora", Nucella",
Brachidontes" , Anasterias" , Arbacia, and Loxechinus fair-
ly common. Lithothamnia gave the fringe a pink or brown-
ish colour.
The littoral region
The uppermost marine species, Elminius, stopped at 515
and the lowest lichen were found at 525. Because of that
the upper limit has been placed at 520. Changes in spe-
cies composition within the littoral were seen at 245, 295,
and about 400, the two last levels being used as zone
limits.
123
 ,E Fig. 14. (left). Zonation at Stn M82, Bahia Sotom6 in
"0"-- Estero Reloncavi.
:C"5t
~ S2 "
,-
:: 0 ;,
"'~o <> E •
E- t Fig. 15 (right). Zonation at Stn M31, Bahia Ralun at the
-<:
~~2 head of Estero Reloncavi.

~~u
~
em
0
em u 0
.:0
~oo.c: :;c
<:" ~ ~.g, ~
_
Only 13 identified species were found, 6 animals and 7
~u CJa.-ti~
b~
o~
~
,-<>
0

.Q-E"f,/)
~ em
600
§ a::::
-,o..COIJJ
~
,
~-
~
~

,,-0
. em
600
algae, plus unidentified amphipods, lichen, and mosses.
This low number is, however, not quite representative.
Because of the sea level being 100 cm above zero, collect-
.~-[~~ ~ ~k

~
ing lowest down was impossible. Also the next lowest
~ t.. til t..
"5Ef?-
~~~-§. ~~~ ~
.;;
metre was difficult to study because it was below the
500 ~ ii"iii.g 500 500 plane of the deck. Our few samples from below 100 were
E - 00
- c: t.._ ,
~
taken by a fisherman without exact information about the

~
LuLuQ..t.J
depth. It is thus not possible to know if species usually

r
400
living low down were missing or if they occurred there
400
but couldn't be sampled. Even above the 2-m level small
MSL - MSL animals and algae can have been overlooked. There is,
however, reason to believe that even if the number of
300 300 species is somewhat higher than that found, the diversity
really is low and that this is the result of low salinity.
The station is situated halfway into the fjord and great
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amounts of freshwater from Rio Petro hue and other riv-

200 200 ers lower the salinity of the upper layers considerably.
! Thus the surface salinity at Stn H20 in Bahia Sotom6,
just outside M82, was only 4.5 %0 on the day of investiga-
100 100 tion (18 %0 at 3 m depth, Table 2). It certainly will be
higher at other occasions but probably always too low to
NO SAMPLING
BELOW 100 permit most true marine animals to live at M82 and too
high to offer freshwater species suitable living conditions.
The only such species found was the pulmonate Chitina,
which was taken also at M31, where the water was al-
most fresh, but at none of the other stations where the
salinity was higher.
Zone 1, 195--295, had a greenish colour and was charac- Most of the few species found were common or abun-
terized by • Balanus flosculus, *Elminius', 'My tilus' , En- dant, probably because of less competition for space than
teromorpha', • Viva', 'Sphacelaria', 'Gelidium*, and at other stations with several times as many species. Be-
lowest down • Macrocystis , all except Enteromorpha hav- cause the species were equally common throughout their
ing their lower limit in the infralittoral. Mytilus was abun- distribution area, their limits were sharp. There were dis-
dant, the others common, and Enteromorpha was the tinct limits at 150/190, where Mytilus and possibly Acan-
most characteristic species. Siphonarians were few and not thocyclus albatrossis had their upper and Chitina and Bos-
observed outside the zone. Several rare species had upper trychia harveyi possibly their lower limit, at 380 where the
limit at 245. three green algae Enteromorpha, Prasiola, and Clado-
phoropsis stopped, at 420 where Elminius had its upper
Zone 2, 295--400, had a blackish colour caused by algae limit, at 560/570 where lichen began to appear, and at
covering Elminius. 'Mytilus was abundant • Elminius' and 625/630 where Bostrychia and probably Calothrix stopped
'Viva'. common in the whole zone, 'Enteromorpha* and
'Sphacelaria lowest down. and the first mosses were found. Chitina was no l taken
above 520 but perhaps occurs higher up.
Zone 3, 4~520, was characterized by 'Elminius, which The areas between these levels had different colours.
here was not covered by algae and formed a distinct belt The visible area below 260 was brown from diatoms and
in the whole zone. 'Viva was found lowest down and young stages of Enteromorpha overgrowing Elminius.
some • Enteromorpha in the middle part.
Table 2. Salinity (%0) and temperature (0C) in the upper-
The supralittoral region most five metres at Stn H20 just outside Stn M82, and
A 10 cm wide naked area between 515 and 525 marked at Stn H6 just outside Stn M31 on the days in 1949 the
the limit between the littoral and supralittoral regions. stations were investigated and, at H6, at an earlier occasi-
The only species in the region were lichen. on (after BRAITSTROM & DAHL 1952).
Stn H20 Station H6
Stn M82, Estero Reioncavi, Bahia Sotomo, Depth Sal. Temp. Salinity Temperature
(m) 31 March 5 Jan. 1 April 5 Jan. 1 April
41°38'30" S, 72°22'47" W, 31 March 1949 (Fig. 14).
Fairly sheltered rocks, which partly were too steep to
o 4.5 5.5 0.6 2.4 16.0 11.8
1 5.9 6.3 0.9 5.0 15.4 12.4
permit working on the shore, so that investigations had 2 8.4 6.5 1.5 8.5 14.8 12.4
to be carried out from the boat, which was moored to the 3 18.0 6.4 16.5 14.5 12.3
rocks. This made measurements less accurate than at sta- 4 16.7 26.4 22.4 12.6 12.3
tions where a meter could be used, but the order in which 5 24.4 28.2 23.1 12.1 12.3
the species occurred is correct.

124
 Fig. 16. Blidingia minima
(NAEG.) KYLlN, Porphyra colum-
bina MONT., and Bostrychia har-
veyi MONT. on Juncus sp. at Stn
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M31, Bahia Ralan.
Above 260 other algae were so common that they hid the
barnacles. Since these algae were not found above 380,
the light colour of the barnacle shells became visible there
and gave the impression of a narrow greyish belt up to
420, the upper limit of Elminius. At a distance this limit
looked horizontal, but at close quarters both this and
other limits were seen to be wave-like, a result of differ-
ences in rock configuration and wash. Thus, all measure-
ments are means. Above 420 there was an upper brown
or yellow-brown belt, getting its colour from Calothrix and
Bostrychia, to 560, above which level white and yellow
lichen and mosses dominated.
Acanthocyclus was with certainty found between 120
and 190, but a sample also containing Hemigrapsus, was
labelled 'in the Elminius belt', without precise levels, so
it is not unlikely that both these crabs also occurred lower
down and above 190. According to the field notes amphi-
pods were found in the 'Elminius and upper algal belts',
which correspond to the grey and upper brown belts, and
Paradynamenopsis was noted from the grey belt. Clado-
phoropsis and Calothrix were not found at any other sta-
tion.
The zonation pattern differed from that of the stations
in Seno Reloncavi proper. The species found both at M82
and in Seno Reloncavi occurred lower down a( M82 than
in the Seno. Thus Mytilus was observed from below zero
up to 190 only, whereas it in Seno Reloncavi was a
littoral species taken from 60 up to 540. Elminius and
Enteromorpha, which occurred from below zen> to a little
above MSL, thus in the lower part of the littoral and also
in the infralittoral, mainly occurred in the upper part of
the littoral in Seno Reloncavi. Bostrychia harveyi, in Seno
Reloncavi only taken upmost in the littoral and in the
supralittoral, was found down to about 160 at M82.
When species which in Seno Reloncavi are littoral and
especially characteristic of the upper part of that region
and even occur in the lower part of the supralittoral, at
M82 occur at levels which in Seno Reloncavi are consid-
c red to belong to the infralittoral and the lower part of
,he littoral, it is not possible to find out where at M82 to
place the border between the two regions. It is, however,
the vertical distribution which is of interest, not if the
species are classified as infralittoral or littoral, and I will
refrain from trying to define the lower limit of the littoral
but only state that this region has its upper limit at
560/565, where white and yellow lichen began to appear
and the highest occurring algae, Calothrix and Bostrychia,
became scarce, and that there is a change in species com-
position between 380 and 420.
The lowest 70 cm of the supralittoral had a mixture of
marine algae and lichen. Then dense forest took over.
Stn M31, Estero Re1oncavi, Bahia Rahin, Cayo
Nahuelguapi, northwestern point, 41°24'30" S,
72°19'05" W, 1 April 1949 (Fig. 15).
The small skerry consisted of hard rock. It was low,
hilly, and very sheltered. Upmost was a freshwater swamp
with Juncus sp. or a similar plant (Fig. 16). Rio Petrohue
and some smaller streams discharge great amounts of
freshwater into Bahia Ralan. Two measurements of salini-
ty and temperature at Stn H6, just off M31, show that
M31, unlike the other 11 stations, is practically a fresh-
water station (Table 2).
Because of the low salinity only a few euryhaline marine
species occurred, competing for space with freshwater and
terrestrial ones. Only 3 animals and 4 algae were found,
together with some undetermined algae, lichen, mosses,
and grasses. A fifth alga, Enteromorpha ramulosa (J.E.
SMITH) HOOK., was found nearby. This species and Blid-
ingia were not taken at any other station, and Chilina
only at this station and M82 farther out.
As at M82 there was no distinct zonation pattern, but
there was a change in species composition at about 300
and 570, these levels being lower and upper limits of Blid-
ingia, Porphyra, and Bostrychia harveyi. Since the last
marine species stopped at 570 this would be a natural
upper limit of the littoral region, but lichen began to ap-
pear already at 405 and Juncus at 460, species not usually
found in the littoral.
Only 4 species were common to M82 and M31. Of
these Elminius and Chilina had similar distribution at
both stations. At M31 the barnacle occurred from at least
-50 to 300, in crevices to 375, and single specimens to
125

450 in rock pools, where the water on the day of investiga-
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tion was fresh. It was extremely luxuriant below 300,
growing in decimetre-thick layers, which because of the
heavy pressure from the crowded specimens had become
loose from the rock and bulged up from this. Thanks to
the pressure many specimens had got tube-shape and
were extremely high. The barnacles were covered by early
stages of green algae, probably Enteromorpha sp. and
Ulva sp. Chilina occurred in the Elminius belt and also
above this to 510 (at M82 to 520). As at M82 Hemi-
grapsus was common in the Elminius belt, and Bostrychia
stopped at 570, the level at which it became rare at M82.
Above about 300, where Elminius ceased to be belt-
forming, there was a belt of Lyngbya with Hemigrapsus
and Chitina to about 440. Lyngbya grew as a greyish-
green felt on the rocks from about 265-420, from about
295 together with Blidingia and Porphyra, and from 315
with Bostrychia. Above 420 these three algae only oc-
curred locally, to about 570. From about 460 they as well
as stray specimens of Elminius grew on turf and among
mosses and grasses but also on the bare rock in trickling
water from the Juncus swamp and even on Juncus (Fig.
16).
THE ANCUD AREA
The stations (Fig. 1) will be treated in the following
order: MlO, M55, M56, and M57. Because the
tidal amplitude is much smaller than in the Seno
Reloncavl area all species have a smaller vertical
distribution and occur lower down than at the sta-
tions in the latter area and the measurements of
levels in the two areas cannot be directly com-
pared, whereas the vertical order in which the spe-
cies occur is comparable.
Stn MiO, ChilOt!, Bahia de Ancud, Punta el Morro,
southwest of Ancud, 41°52'40" S, 73°50'48" W, 2
March 1949 (Fig. 17).
The rocks were rugged with numerous small, sharp-
edged holes and small rock pools and had a steep edge
towards the sea. They were fairly exposed to winds from
126
em
Fig. 17. Zonation at Stn MlO,
Punta El Morro.
the north, but shallow water outside moderated the waves.
Part of the station was sheltered by a rock, separated from
the shore by a few metres of water. There was some pollu-
tion from Ancud.
Only 35 identified animals and 11 algae were found and
also sponges, 3 other undetermined species, and lichen.
No other station had so many sponges and ascidians.
Many species usually found in the area but not at MlO
were found in rock pools there or outside the section stud-
ied.
The infralittoral fringe
The lowest 20 cm couldn't be investigated because of tur-
bid water, so the lower limit at 20 of many littoral species
is probably not their real lower limit but rather one of
possible observation. Though some infralittoral species
stopped or became less common at 70, the natural limit
to the littoral seems to be at 50, since many common
littoral species had their lower limit there. Spirorbids*,
Tegula* , Crepipatella*, Ulva*, Sphacelaria* , and Crodelia*
were common or abundant.
The littoral region
The limit to the supralittoral is found at 220, where two
of the most abundant littoral species, Littorina araucana
and siphonarians, had their upper limit. Zone limits have
been placed at 110, where some species stopped or became
more abundant and one had its lower limit, and at 150,
where there was a more distinct change in species compo-
sition and abundance. Above 150 the species stopped one
by one until 220.
Zone 1, 50-110. Balanus flosculus*, siphonarians", and
Enteromorpha* began at 50, Chthamalus cirratus" and
Gelidium* about midway. 'Spirorbids, • Tegula", siphon-
arians*, *Sphacelaria, and *Crodelia* were common or
abundant in the lower part, Balanus, Patelloida*, 'Crepi-
patella", Enteromorpha, and *Ulva* in the whole zone,
and Chthamalus and Gelidium in the upper part.
Zone 2, 110-150. *Chthamalus cirratus*, 'siphonari-
ans", and "Ulva dominated also in this zone, and *Bala-
nus flosculus, • Patelloida*, "Crepipatella*, * Enteromor-
pha, and *Gelidium* were common.
Zone 3, 150-220, had few species. As in zone 2
*Chthamalus cirratus" and *siphonarians dominated
 Fig. 18. Zonation at Stn M55,
between Punta San Antonio and
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Punta Colorado.
throughout, and Littorina became dominating upmost.
'Chiton granosus was common lowest down, 'Gelidium
up to 200, and Onchidella was fairly common in the up-
per part of the zone.
The supralittoral region
Only four species were found. 'Chthamalus cirratus, abun-
dant from 90 in zone 1, was still abundant up to 240, in
depressions with wash. 'Porphyra was taken somewhere
between 200 in zone 3 and 265. Prasiola, the only marine
species not found lower down, was abundant between 225
and 275, and yellow lichen from 245.
Stn M55, Chiloe, Canal Chacao, between Punta San
Antonio and Pu~ta Colorado at the southern side
of the mouth of Bahia de Ancud, 41°51'30" S,
73°49'50" W, 27 February 1949 (supplementary
studies other days) (Fig. 18).
The rocks were hard, partly vertical, partly sloping with
many holes and rock pools and exposed to winds from
northwest to northeast, but kelp beds outside gave some
shelter.
Only 25 identified animals and 23 algae were found,
10 animals less and 12 algae more than at MIO. In addi-
tion undetermined sponges and some other animals were
found. No higher taxon was represen,ted by many species.
Waves, splash, and spray wetted the rocks high up and
kept holes and crevices filled with water. Because of that
some species were found higher up and had a wider verti-
cal distribution than at the more sheltered MlO, and ac-
cordingly the limits between the regions and zones were
found higher up than there, the difference being about
30 cm lowest down and about 60 upmost (see Fig. 23).
The infralittoral fringe
The limit to the littoral region could be placed at 60, 80,
or 100, but since several common littoral species had their
lower limit at or near 80 or became more common there,
80 seems to be the best choice. . dominated and 'Rhizoclonium" was common throughout.
The rocks were low down covered by crustose coral-
lines', which higher up were concealed by other algae.
Viva' was the only truly abundant species, lowest down
as tufts on mollusc shells, higher up forming a green mat.
Phymactis', spirorbids', Balanus flosculus', a crust-
shaped bryozoan, Pyura, Enteromorpha bulbosa', and
Crodelia' were common.
Few oLservations were possible below O. For this reason
the lower limit of many species at 0 is not the real lower
limit. Three species of sea-anemones were mostly found
in water-filled holes. On smooth rocks their upper limit
was lower than in the holes. Balanus was mainly found
on the exposed, seaward side of the rocks.
The littoral region
At the upper limit at 275/280 the two dominating algae,
Porphyra and Gelidium stopped. Three zones with fairly
distinct limits at 140 and 190 could be distinguished.
Zone 1, 80-140, had four abundant species, 'Viva' in
the whole zone and 'Gelidium', Hildenbrandtia', and Iri-
daea boryana" in the upper part. Many other species
were common, lowest down at least two species of
sponges, "Phymactis, Fissurella costata, and" Lithothamni-
on, highest up Chthamalus cirratus" and Chiton granosus,
and in the whole zone • Patelloida, Brachidontes, "Entero-
morpha", "Crodelia', and "Corallina officinalis. Viva and
Crodelia characterized the zone, upmost together with
Iridaea. Siphonarians' had their lower limit low down.
Zone 2, 140-190, had few but fairly common to abun-
dant species. "Viva, " Gelidium" , and 'Iridaea were as
abundant as in zone 1, • Hildenbrandtia only lowest down.
'Chthamalus cirratus" was common throughout, • Entero-
morpha and 'Crodelia lowest down, and Rhizoclonium
riparium* upmost. The upper zone limit was very dis-
tinct. Viva and Iridaea stopped there and Littorina and
Porphyra began to appear.
Zone 3, 190-280, had still fewer but mostly common
or abundant species. Littorina', • Gelidium, and Porphyra
127
Downloaded by [UQ Library] at 18:20 01 September 2013
Fig. 19. Zonation at Stn M56, Punta Corona.
*Chthamalus cirratus in the lower half. Chthamalus
formed a distinct belt, taking over the dominant role of
Iridaea boryana. *Siphonarians stopped low down, where
Phytia* sp. began to appear. The limit to the supralittoral
was very distinct because the abundant Gelidium and
Porphyra stopped there. The limit of Gelidium at 280 is,
however, not absolute for single specimens occurred in a
crevice at 400.
The supralittoral region
In addition to the *Gelidium species just mentioned, only
three species occurred, namely • Littorina which was abun-
dant to 350 and common to 380, • Phytia sp. which was
found to 310, and *Rhizoclonium, common to 300.
Stn M56, Chiloe, Golfo Coronado, Punta Corona,
northern point, 40°47' S, 73°53'07" W, 26 and 28
February 1949 (Fig. 19).
The shore consisted of gently sloping, step-like and soft,
rough sandstone with numerous small holes (Fig. 20) and
crevices and highest up many small, shallow rock pools.
It was more exposed than the other stations in the area,
especially to winds from northwest over north to east. In
spite of this the vertical distribution of the species was
less wide than at M55 and M57 and more like that of the
most sheltered MI0. The reason can be shelter from some
kelp outside the station and that the steps in the rock had
a braking effect on the waves.
The number of species was considerably higher than at
any of the other three stations, perhaps a result both of
the exposure and the many holes and crevices, which gave
shelter and remained water-filled or wet even when the
shore was emerged. There were also many more abun-
dant species than at the other stations. In all 55 animals
and 28 algae were found, in addition to a 'peje sapo', a
lichen, and at least 9 other undetermined species. Algae
dominated and in the field notes the station was charac-
terized as an algal station. No less than 8 isopod species
were found, but this does not mean that Isopoda have
been overlooked elsewhere. Their presence at M55 may
be due to the many algae. Small animals were often only
found when algal samples were washed.
128
The infralittoral fringe
Many infralittoral species stopped at 10, 30/35, 50, and
60, but three of the dominant algae disappeared at 60,
and because of that the limit to the littoral has been
placed there.
The fringe was characterized by the algae, among them
5 species of corallines, but also by sponges*, sea-anemo-
nes, spirorbids, chitons', siphonarians*, and ascidians.
Antholoba was abundant lowest down, Phymactis*, Patel-
loida*, and Tegula* common throughout the fringe, spir-
orbids* highest up, and a crust-shaped and a soft, white
bryozoan low down. The most common algae were Macro-
cystis, Lithothamnion, and Corallina chilensis lowest
down, and Ulva lactuca*, U. nematoidea*, Crodelia*,
Corallina officinalis*, Trematocarpus, Iridaea boryana*,
l. ciliata with its epiphyte Ceramium rubrum, Centro-
ceras*, Heterosiphonia, and Herposiphonia in practically
the whole fringe.
The littoral region
Many of the most common species had their upper limit
at 195/200 and the upper limit of the region has been
placed there. Distinct zone limits were seen at 100 and
140.
Zone 1, 60-100. *Phymactis, *Patelloida*, the two *Ulva*
species, 'Crodelia*, 'Iridaea boryana', and 'Centroceras
were about as common or abundant as in the infralittoral
fringe. Also Porphyra* and Bostrychia hookeri*, with
lower limit in the zone, were abundant. Phymactis and
Centroceras stopped at or near the upper limit.
Zone 2, 100-140. The two last-named species excepted,
all abundant or common species from zone 1 were also
found in zone 2, where also Chthamalus cirratus' , Brachi-
dontes* , Lasaea*, and Enteromorpha*, having lower limit
in the zone, were common or abundant, and • Bunodactis
and Onchidella*, fairly common to common. Some
sponges, but not the same as lower down, were found.
*Ulva lactuca stopped upmost.
Zone 3, 140-195/200. Here, too, some sponges were
found. All important species were the same as in zone 2
and were abundant or common. 'Patelloida, *Ulva nema-
 Fig. 20. Stn M56, Punta Coro-
na, detail of the rocks.
toidea" and • Iridaea boryana stopped near the upper
limit. *Chthamalus cirratus, • Littorina (rare in zone 2),
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'Onchidella, *Brachidontes, *Lasaea, *Porphyra, *Crode-
lia, and *Bostrychia stopped at 195/200, and since most
species were very abundant (Onchidella and Brachidontes
common) all the way to the upper limit, this was very
distinct. *Enteromorpha* crossed this limit.
The supralittoral region
*Enteromorpha was found up to 240 together with some
dwarfed specimens of • Porphyra. Yellow lichen had their
lower limit at 230.
NOT INVESTIGATED
Fig. 21. Zonation at Stn. M57, Punta Auhi.
Stn M57, Chiloe, Canal Chacao, Punta Auhi, at the
northern side of the mouth of Bahia de Ancud,
41°49'47" S, 73°51'46" W, 1 March 1949 (Fig. 21).
The rocks were a soft sandstone with harder outcrops.
Outer border steep with crevicies and single small rock
pools, inner part gently sloping with few, shallow rock
pools. The station was exposed to winds from north and
northwest.
Only 41 identified species were found, less than at any
other station in the area. As at M55 the algae with 18
species were relatively better represented than the ani-
129
 mals with 23. In addition amphipods and one lichen were
found. The number of species gradually decreased up to
100, above which level many species had common upper
limits at 120, 150, and 245. Based on the distribution of
all species, and not only those in Fig. 21, the upper limit
of the infralittoral fringe has been placed at 90, that of
the littoral at 245 with zone limits at 150 and 180.
The infralittoral fringe
No investigations could be made below zero, so the lower
limit of most littoral species is unknown. Ulva* was abun-
dant in the whole fringe, Lithothamnion' and Corallina
chilensis' in the lower part and common in the upper.
Balanus flosculus', Fissurella costata', Tegula', Ralfsia
australis', Gelidium lingulatum', Crodelia', Ceramium
dozei' , Heterosiphonia*, and Polysiphonia' were common
in the whole fringe or part of it. The less common Balan-
us psittacus only went to 60 and spirorbids to the fringe
limit.
The littoral region
Zone 1, 90-150. The species abundant or common in
the infralittoral fringe were also found, with the same
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abundance, in the whole zone 1, except for • Gelidium,
'Crodelia, 'Corallina, and *Heterosiphonia, which
stopped at 110/120.
Zone 2,150-180, was very narrow and mainly populated
by the uppermost specimens from zone 1. They stopped
at the upper limit, *Ralfsia though already at 170. *Ulva'
was still abundant, and Enteromorpha*, which began at
150, was common.
Zone 3, 180-245, was wider and more characteristic
than zone 2. Chthamalus cirratus*, Brachidontes, Lasaea,
Porphyra', and Bostrychia hookeri' began to appear at
the lower limit or a little above. Chthamalus and *Ulva
dominated, the others and *Enteromorpha' were more
or less common. Littorina was fairly common lowest down.
The supralittoral region
Only four species crossed the border from zone 3,
'Chthamalus, abundant to 280, • Enteromorpha and • Bos-
trychia, common to 255/265, and • Porphyra fairly common
to 280. Yellow lichen were found from 320, on exposed
parts.
COMPARISON BETWEEN THE TWO AREAS
Number of species
Only 1 sipunculid, stomatopod, and pycnogonid
species were found. The Turbellaria (4) and Isopo-
da (9), small and hiding under stones or among
fronds of smaller algae or in holdfasts of kelp spe-
cies, have no doubt often been overlooked, and
Nemertini (9) were only casually seen, since they
lived hidden in sand, in polychaete tubes, in shells
or under stones. The material of these 25 species
is too scanty and casual to be of any value in a
discussion of zonation. Of the 212 identified marine
species thus 187 remain (see Table 1) to be dis-
cussed, but also among them many are represented
by one or few specimens only and therefore of
little interest in such discussions. The number of
species to be discussed or figured is therefore relati-
130
vely low, but as mentioned earlier all available
information also on those, which not or only occasi-
onally will be treated, has been gathered in Appen-
dix 1.
Table 1 shows that 110 of the 187 species are
animals and 77 algae. It also shows how many spe-
cies of each taxon were found at the respective sta-
tions. Of the animals about 1/3-1/2 were found at
each station in Seno Reloncavi proper but only
about 1/4-113 at those in Ancud, whereas the algae
were about equally well represented at all, except
at M22 and MI0. M91 and M37 in Seno Reloncavi
had 79 and 73 identified marine species respective-
ly, and no other station in this area had less than
59. Lower numbers were found at the mouth of
Estero Reloncavi (M139) and especially in that
fjord. Table 1 also shows a great difference in An-
cud between M56 with 72 species, only 1 less than
at M37 in Seno Reloncavi, and the other three sta-
tions with only between 41 and 46 species. Since,
however, sampling was easier at M56 several of the
smaller species found there may have been over-
looked at the other three stations. There is thus
reason to believe that the differences in number of
species at the four Ancud stations are smaller than
the present figures seem to indicate, but even if the
number of species at three of these stations may
be underestimated, there nevertheless is a clear dif-
ference in numbers between the two areas.
Species composition
Of the 187 marine species considered 138 were
found in the Seno Reloncavi area and 108 in An-
cud, and only 41 animals and 18 algae were com-
mon to both areas. Thus there not only is a differ-
ence in number of species in the two areas but also
in species composition. The difference is most dis-
tinct and probably most real concerning larger and
conspicuous species, which are not easily over-
looked and which were especially searched for as
being suitable character species in zonation studies.
Table 3A shows that no less than 29 of the most
widespread species (a few of them not common)
found at 3-6 stations in the Seno Reloncavi area,
were not taken at any station in Ancud, and Table
3B that 8 such species found at 3 or all 4 Ancud
stations were not found at any of the stations in the
Seno Reloncavi area, where investigations were
easier to carry out and the risk of overlooking a
species less. This does not imply that the species
not found in one of the areas are totally absent
there, only that they were absent or not observed
at the stations studied. In fact a dozen species
(marked with an asterisk in Table 3A) found at the
Seno Reloncavi stations have been taken in the
 Table 3. Differences in species composition at the stations in the Seno Reloncavi and Ancud areas. Species marked
with an asterisk are known from the other area though not at the stations investigated. The remarks on distribution
are based on material from LUCE if reference to a publication is lacking.
A. Twenty nine species found at 3-6 stations in Seno and
Estero Reloncavi but not at any of the four Ancud sta-
tions.
Chaetopterus ?variopedatus 3.
Chthamalus scabrosus 6.
• Elminius kingi 6. Also known from Chiloe (NILSSON-
CANTELL 1957).
Pagurus edwardsi 3.
'Pagurus villosus 5. Also recorded from Chiloe (RETAMAL
1981).
'Cancer edwardsi 3. Also two records from Chiloe, 8
fathoms, and Ancud without further information
(GARTH 1957).
'Pinnotheres politus 4. Also known from Castro, east
coast of Chiloe (GARTH 1957; RETAMAL 1981).
Pinnixa bahamondei 3.
• Hemigrapsus crenulatus 6. Also intertidally near Ancud
and in a rock pool at MI0.
(*)Cyclograpsus cinereus 5. Known from Peru to Calbuco;
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Chiloe not mentioned (GARTH 1957; RETAMAL 1981).
*Ischnochiton pusio 3. Also taken at 8-25 m depth in
Canal Chacao.
Tonicia atrata 5.
Fissurella oriens 4.
Nacella magellanica 6.
Littorina peruviana 3.
*Nassarius gayii 5. Also in sand near Ancud.
• Anasterias varium 4. Also in 6-40 m depth in Canal
Chacao.
Ancud area, too, but only in rock pools, at other
littoral stations or in deeper water in Canal Chacao
or, by other authors at some place in eastern Chi-
loe, mostly without details on localities and depths,
and Taliepus dentatus, found at all 4 Ancud stations
(Table 3B) but at none in the Seno Reloncavi area,
is known from this bay, though not intertidally.
Then 17 species remain in the Seno Re10ncavi area,
which are not hitherto known from the Ancud sta-
tions or from Chiloe altogether, and 7 in Ancud,
which have not been recorded from anywhere in
Seno or Estero Reloncavi.
The less common and less wide-spread species
show a similar difference in species composition.
However, also some of these rarer species, lacking
at stations in one of the areas, have been taken at
other localities in that area, mostly.in deeper wa-
ter, and if the lowest part of the shore could have
been investigated at all stations, more species, both
common and rare, would probably have been found
there, increasing the number of species common to
the two areas. Future studies will no doubt reduce
the difference between the areas both in numbers
and composition, especially of the less common
species.
Some differences, hardly due to oversight, will
be commented upon here. As has been shown the
Ophiactis asperula 3.
*Arbacia dufresnii 5. Also taken at 8-40 m depth in Ca-
nal Chacao and recorded from east coast of Chiloe
(LARRAIN 1975)
'Chaetomorpha aerea 3. Also found in rock pools at MID
and M55.
Rhizoclonium tortuosum 3.
Scytothamnus fasciculatus 3.
Lyngbya aestuarii 3.
Acrochaetium catenulatum 3.
Chaetangium fastigiatum 4.
Catenella fusiformis 3.
'Chondrus canaliculatus 5. Also in rock pool at M56.
Iridaea crispata 3.
Bostrychia harveyi 4.
B. Eight species found at (2) 3-4 Ancud stations but not
at any of the stations in the Seno Reloncavi area.
Phymactis clematis 4.
'Taliepus dentatus 4. Also in Seno Reloncavi but not inter-
tidally.
Chaetopleura peruviana 2.
Fissurella costata 4.
Patelloida zebrina 3.
Onchidella marginata 3.
Leathesia difformis 3.
Trematocarpus dichotomus 3.
number of species in Estero Reloncavi is extremely
low, only 36 at M139 at the mouth, where the salini-
ty is somewhat lowered, 13 at M82 midway, where
the salinity is low, and only 7 at M31 at the head,
where the water is practically fresh. Not astonish-
ingly the lastnamed two stations were the only ones
where the freshwater pulmonate Chilina bulloides
occurred. • Anthozoa, 'Chaetopterus ?variopedatus,
Polyplacophora, Prosobranchia, *Opisthobranchia,
siphonarians, Echinodermata, *Ascidiacea, and
Phaeophyta were totally missing in the fjord, those
marked with an asterisk also at the mouth, at M139.
It is evident that the low diversity and the presence
of Chitina at the fjord stations is a result of the low
salinity. Most of the few species found in Estero
Re10ncavi were very common, probably because of
less competition for space and food.
Most unexpected was the absence at the Ancud
stations of Chaetopterus ?variopedatus, Chthamalus
scabrosus, Elminius kingi, the decapods Pagurus
edwardsi, P. villosus, pinnotherids, Hemigrapsus
crenulatus, and Cyclograpsus cinereus, Ischnochiton
pusio, Tonicia atrata, Nacella magellanica, Nassari-
us gay ii, Chaetangium fastigiatum, and Chondrus
canaliculatus, which all were common and found
at many stations in Seno Reloncavi. Also unex-
pected was that Loxechinus albus was the only echi-
131
 noderm species taken in Ancud (at one station on-
ly) whereas 9 of the 15 species taken by LUCE in
Seno Reloncavi, were found in Bahia de Ancud,
outside my stations. Equally unexpected was that
Mytilus edulis chilensis played an insignificant role
at the Ancud stations and that Phymactis clematis,
Patelloida zebrina, Onchidella marginata, and Lea-
thesia difformis, which were common at 3 or 4 sta-
tions in Ancud, were absent at the Seno Reloncavi
stations. Again it must be remembered that these
differences in species composition are valid only for
the stations/sections at which zonation studies were
carried out. Some of the species missing at '"just
these stations were found at others in the same area.
The absence of Pagurus villosus and Pinnixa ba-
hamondei at the Ancud stations is natural. P. villo-
sus lives in shells of Nassarius gayii and sometimes
in Nassarius dentiferus, none of which was found
at my Ancud stations, which all are rocky. These
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snails seem to prefer sand-bottom, and Pagurus has
in fact been recorded from Chiloe (RETAMAL 1981).
Pinnixa lives in tubes of Chaetopterus ?variopeda-
tus, which was not taken at any Ancud station.
Difficult to explain is the opposite situation for
Pagurus edwardsi, Pinnotheres politus, and
Pinnaxodes chilensis, which also were missing at the
Ancud stations. The hermit crab lives in shells of
Tegula atra, found at all four stations, and also in
Nucella calcar, which was taken at MlO, Pinnothe-
res lives under shells of Crepipatella dilatata, which
was found at all four stations, and Pinnaxodes in
Loxechinus albus, which was taken at M56. The two
pinnotherids have, however, been recorded from
Chiloe (RETAMAL 1981).
Turbellaria, Nemertini, Paranthus niveus, and
Chaetopterus ?variopedatus were only found in Seno
Reloncavi, mostly on the underside of stones. The
lack of samples of these taxa from Ancud is proba-
bly due to the fact that this type of biotope was
not found there. Differences in the biotopes, in
Seno Reloncavi boulders and stones in sand, in
Ancud rocks, can perhaps also explain" why more
animal species were found in the littoral of Seno
Reloncavi than in Ancud. Though the rocks had
many depressions, crevices, and holes where ani-
mals could find shadow and shelter against the
waves and a wet or moist place when the tide is
out, boulder shores have more niches and places
where animals can avoid predators and the force
of the sea and find hiding places, when the shore
is emerged (ct. VIVIANI 1975:178).
Also unknown competition conditions and local
human interference (removal of edible species) may
have been different in the two areas. Balanus psitta-
cus, many crabs, molluscs, Loxechinus albus, Pyura
132
chilensis, Porphyra columbina, and other species
are sampled for consumption by the local popula-
tion, in many places to such a degree that they are
now rare there or have even been eradicated, as for
example Loxechinus in Calbuco (LARRAINE 1975).
The species assemblages found at such places thus
are not typical of an undisturbed shore. Removal
of one species can also have influence on others.
Thus JARA & MORENO (1984) and MORENO & al.
(1984) have shown that when Fissurella picta and
other herbivores were removed, Iridaea boryana
increased in density, and also that areas nearest to
large fishing villages had a higher cover of Iridaea
and a lower abundance of F. picta, the latter having
been removed for consumption. Also the absence
of some snails in Ancud and of Aulacomya ater,
Ostraea chilensis, and Loxechinus at three of the
stations and the scarcity of Mytilus edulis chilensis
there perhaps can be the result of similar activity.
Hydrographical conditions may also play a role.
The Ancud stations are on the open coast, those
in Seno Reloncavi in an archipelago, and many
species prefer one or the other type of environ-
ment. Chaetopleura peruviana was described from
Peru, where there is open coast. As this species
was found at three Ancud stations but not in Seno
Reloncavi, the first idea that comes to hand is that
this is an open-coast species, which does not thrive
under conditions in an archipelago (less exposure,
lower salinity) or has not been able to reach the
latter, but Littorina peruviana, also with type locali-
ty Peru, was found at three stations in Seno Relon-
cavi but at none in Ancud, in spite of the fact that
it, according to KNOX (1960:609), is dominant in the
lower supralittoral in Chiloe.
To be able to confirm if the differences found
are real, and if so is the case, to find out if they
are general, investigations of larger areas over a
longer period and at different seasons must be car-
ried out. Such studies perhaps also will help to find
the causes of possible differences.
Differences in vertical distribution
Fig. 22 (see also Appendix 1) seems to indicate
that some species and higher taxa richly represented
in both areas and therefore comparable have a
slightly higher relative upper limit at the Ancud
stations than at those in Seno Reloncavi proper.
Some are also more abundant higher up in Ancud.
Local differences such as type of substrate, direction
of the localities, slope, degree of exposure, fresh-
water runoff from rivers and swamps, and even
waves from boat traffic cause differences in the
vertical distribution of organisms even between
nearby stations, but unless one or more factors are
 '055 5657 229' 905937'39 '0555657

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Lichen HIgher plants Onchidella marginata Porphyra columbino Bostrychio hooker/ + harvey;

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Rhizoclonlum forfuosum • Hemigrapsus crenulatus Cirripedia Gelidium pseudointricofum Mocruro • Anomura

ripor/um

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11
Downloaded by [UQ Library] at 18:20 01 September 2013

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Aconthocyc/us albafrossis Iridea, 4 species Nucella calcar

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Crodelia accedens
IiIJ Various Rhodophyta
11

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Sponges Anthozoa U{va lacfuca Various Prosobranchia

Polyplocophoro Nacelfa magellanico Phaeophyta Crepipoteflo di/atoto Lithothomnion. 2 spec las

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Aulacomyo afer Pyura chilensls Ostreo chilensis
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Chondrus cano/icufotus Hormomyo granulata

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Asteroidea Ophiuroidea HolothuriOldea Flssurella. 5 species Holicarelnus planofus

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Concer spp. + pinnotherids Ech i noidea Spirorbids Brachiopoda + Bryozoa Various Ascldiacea

. . Abundant til Common ~ Fairly common ~ Rare 1:8] Frequency not noted

Fig. 22. The occurrence and abundance of 50 species or higher taxa at the five levels with comparable ecological condi-
tions.

identical at all stations in an area and differ from In the chapter 'Choice of base level' was men-
those in another, can they be the cause of general tioned that the zero for the measurements at the
differences in vertical distribution. Ancud stations may differ somewhat from that used

133
 in Seno Reloncavl, but since the amplitude of the
tides in· Ancud is only about one third of that in
the other area, a possible difference in height of the
base levels cannot be great. This is supported by
the fact that the relative height of the lower limits
of the species above the respective base levels don't
differ much in the two areas. A possible difference
in zero levels is thus not great enough to explain
the observed higher upper limits of some species in
Ancud.
One reason for the differences in upper limits,
if these are real, can be the different nature of the
shores in the two areas. In Seno Reloncavi they are
gently sloping with boulders and stones. The larg-
est boulders were found in the lower part of the
shore and acted as breakwater, so the upper part
becomes less wetted by waves than it would have
been without the sheltering boulders. This contri-
butes to lowering the upper limit of many species.
Downloaded by [UQ Library] at 18:20 01 September 2013
Furthermore, many rocky-shore species only occur
if the substrate is continuous and not broken up in
such isolated smaller areas as boulders and stones.
This, too, prevents many species from living as high
up as they would have, if also the upper part of the
shore had had big boulders or been rocky.
Though shallow water or kelp beds outside the
stations in Ancud gave some shelter, the stations
were more exposed than those in Seno Reloncavi,
and since they were rocky the waves will break and
wet the shore high up, and where the rocks were
sloping, the waves rolJed high up. The rocks were
also full of crevices and small holes (Fig. 20) in
which water was left, when the sea receded, or
which were kept wet by waves, splash, and spray.
This, too, could permit many species to occur high-
er up than in Seno Reloncav!.
On the same type of shore a wave, let us say 1 m
high, will probably, even at LW, wash the whole
intertidal area in places with small tides, as in
Ancud, whereas it where the tidal amplitude is as
great as in Seno Reloncavl, at LW only will wash
the lower part of the intertidal area, leaving that
above more or less dry. This may be one more rea-
son why species in Ancud sometimes occur higher
up in relation to the tidal levels than in Seno Relon-
cav!.
Though the upper limits of the species may vary
from station to station and even between the Ancud
and Seno Reloncavl areas, the vertical order in
which the species occur on the shore usually, as
could be expected, is the same in both areas. The
small differences in level of the upper limits of some
species thus have no influence on the zonation pat-
terns at the different stations, discussed in next
chapter.
134
ZONATION PATIERNS
Actual vertical distribution
Figs 4,5,7-9, 13-15, 17-19, and 21 show the verti-
cal distribution at the 12 stations of the more impor-
tant species and also that, though some have a wide
vertical distribution and some also are common
within their whole area of occurrence, most are
restricted to a certain narrower level or are common
only there. There is a great variation in the distribu-
tion patterns (cf. Figs 22 and 24), but when all speci-
es of some importance are considered, the afore-
mentioned five more or less distinct levels with dif-
ferent species assemblages and abundances could
be distinguished; the infralittoral region/fringe; the
littoral region with three zones; and the supralitto-
ral region.
The 12 figures of vertical distribution show the
actual levels, expressed in cm above or below the
zeroes used at the 12 stations, at which the respec-
tive species were found, but because the tidal ampli-
tudes, exposure, and other factors in the two areas
are so different, a comparison of the zonation pat-
terns at the various stations cannot be based on the
actual levels of occurrence. Thus for example a
species found at 360 in Seno Reloncavl and at 113
in Ancud, the mean sea levels at the respective
areas, in fact has the same relative position on the
shore in relation to the tide levels. In both areas
the time the species is submerged and emerged
should be the same at these levels, provided other
factors than the tides don't differ, which they, how-
ever, always do. Accordingly the five regions and
zones distinguished reflect the sum of these dif-
ferent conditions at the respective stations and be-
cause of that have different widths and position on
the shore (Fig. 23), though they in fact have the
same relative widths and position as their equal
numbers at other stations. Therefore a comparison
between the zonation patterns at the stations must
be based on levels with equal environmental condi-
tions and not on tidal levels, for as STEPHENSON &
STEPHENSON (1949:303) stress: 'A mistaken idea,
which is commonly held, is that intertidal zonation
is a result of tidal action. This is not so'.
The 12 figures mentioned above are less suited
for such a comparison. Each of them shows the
zonation patterns of a selection of species occurring
at one station and thus easily permits a comparison
between the species at this very station, but be-
cause the figures are spread out on many pages, a
comparison between the stations is difficult. Fig.
22 has been constructed_to make such a comparison
easier. Each of the 50 sections in the figure shows
the vertical distribution and abundance of 0 n e spe-
 cies or a higher taxon at a II stations and therefore
makes possible a direct comparison of the distribu-
tions of these taxa. Since this figure compares the
five levels with equal environmental conditions, all
levels could be figured equally wide at all stations
irrespective of their real width and vertical position.
Thus each level is represented by a square, and the
degree of blackness of the squares reflects the mean
frequency of the species at the different levels. Spe-
cies hardly entering a level have not been figured
as occurring there, and rare and little known spe-
cies have not been figured. The two stations in
Estero Reloncavf are not included, because the spe-
cies in that fjord were so few and the environ-
mental conditions so different from those at the
other stations that a comparison would be futile.
In the previous chapter was shown that the spe-
cies composition was different at the different sta-
tions and in the two areas. This of course highly
Downloaded by [UQ Library] at 18:20 01 September 2013
influences the zonation patterns, not least from a
visual point of view. One only needs to think how
different a station with barnacles will look com-
pared to one without. This aspect of the zonation
pattern will, however, not be discussed further. The
discussion will be concentrated upon the patterns
of the species present at the respective stations.
Types of vertical distribution
Fig. 22 shows that the same species and some high-
er taxa at all stations, in both areas, for the most
part occur at ecologically corresponding levels and
there occur with about the same abundance. The
differences found are not greater than could be
expected taking local differences in environment
and sampling conditions into consideration. Fig. 22
also shows that at least nine more or less distinct
types of vertical distribution can be distinguished.
In cases when a partial figure shows the vertical
distribution of a higher taxon, information on the
distribution of the species of that taxon is given in
the description of the respective stations and in
Appendix 1.
1. Species exclusively or mainly living in the in-
fralittoral fringe with only single specimens in the
littoral zone 1 at some stations. Examples are Chae-
topterus ?variopedatus, Balanus psittacus, B. laevis,
Alpheus chilensis, Betaeus truncatus, crabs as Piso-
ides edwardsi, Cancer edwardsi, C. plebeius, C.
polyodon, Gaudichaudia gaudichaudi, Homalaspis
plana, Pinnotheres politus, Pinnixa bahamondei,
and Pinnaxodes chilensis, luminous Amphipoda,
Lysiosquilla chilensis, Opistobranchia less Onchidel-
1([ marginata, several Bryozoa, the Echinoidea, and
\scidiacea except Pyura chilensis. These species are
character species of the infralittoral ~ringe.
em
700
600
500
400
MSL
300
200
100
a
Fig. 23. The five regions/zones at the stations in
Seno Reloncavl proper and in the Ancud area.
The infralittoral fringe is black, the littoral region
with three zones dotted, and the supralittoral
region white.
2. Infralittoral and littoral species which are al-
most as common and wide-spread in the infralit-
toral fringe as in zone 1, the lowest part of the lit-
toral which doesn't emerge at all low-waters and
therefore has practically the same environmental
conditions as the infralittoral fringe. Single speci-
mens of a few species in this group were occasional-
ly found also in zone 2 of the littoral. The spir-
orbids, hyaline tubes of another ?polychaete,
Chthamalus scabrosus, Halicarcinus planatus, the
five Fissurella species, Aulacomya ater, Hormomya
granulata, Ostrea chilensis, Asteroidea, Ophiuro-
idea, Holothurioidea, Pyura chilensis, and many
algae like Sphacelaria furcigera, Ralfsia australis,
Leathesia difformis, Lessonia nigrescens, Macrocys-
tis pyrifera, Chaetangium fastigiatum, Chondrus ca-
naliculatus, Iridaea ciliata, and Polysiphonia abscis-
sa have this type of distribution. For many species
in the first two groups it could be difficult to decide
to which of the two to refer them.
3. A mixture of infralittoral and littoral species
which are about equally common in the infralittoral
fringe as in the two lower zones of the littoral re-
gion, as for instance Balanus flosculus, Pagurus
edwardsi, P. villosus, Nacella magellanica, Crepipa-
tella dilatata, Nucella calcar, Chaetomorpha aerea,
several Phaeophyta, Lithothamnion pauciporosum,
L. caroli, Iridaea boryana, and I. ciliata.
4. Species about equally well represented in the
three lower levels but in addition at some stations
135
 occur more or less commonly in zone 3 of the litto-
ral. The sponges were mainly infralittoral but at
some stations in Ancud also occurred in the whole
littoral. Other species are the Anthozoa, Petrolis-
thes laevigatus, Polyplacophora, Tegula atra, Littori-
na peruviana and some other Prosobranchia, Ulva
lactuca, Crodelia accedens (in Ancud), Iridaea cris-
pata, and some more Rhodophyta.
5. Species with the same vertical distribution as
group 4 but which in addition are almost as com-
mon in the uppermost littoral zone as at the three
lower levels, like Acanthocyclus albatross is, Patel-
loida ceciliana, the siphonarians, Mytilus edulis chil-
ensis, Brachidontes purpuratus, and Gelidium pseu-
dointricatum. The siphonarians and Brachidontes
were rare in the infralittotal fringe and Gelidium
most common in the two upper littoral zones.
6. Lasaea petitiana, the only common animal spe-
cies found exclusively in the littoral, where it was'
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most common in the upper two zones, but because
this small species probably has been overlooked in
some places, the observed distribution may be too
restricted. Of algae Scytosiphon lomentaria and Scy-
tothamnus fasciculatus were found only in the upper
two littoral zones, and Anabaena variabilis only in
the uppermost one.
7. Species with the greatest vertical distribution,
occurring from the infralittoral fringe to the supra-
littoral, as Hemigrapsus crenulatus, Cyclograpsus
cinereus, various Amphipoda, Monodonta nigerri-
ma, Littorina araucana, Enteromorpha bulbosa,
Rhizoclonium tortuosum, R. riparium, Bostrychia
hookeri and B. harveyi. Cyclograpsus and Littorina
were occasional in the infralittoral fringe and the
Rhizoclonium species most common in the upper
parts.
8. Chthamalus cirratus, Elminius kingi, Parady-
namenopsis lundae, Onchidella marginata, and
Porphyra columbina were not found in the infralit-
toral fringe at any station but were about equally
common in the whole littoral and, unlike Lasea at
some stations also in the supralittoral. Also Lyng-
bya aestuarii, Microcoleus chtonoplastes, Cladopho-
ra subsimplex, and Catenella fusiformis were not
found below the littoral nor in the lowest zones of
this region.
9. Lichen and land plants, which were found
mainly in the supralittoral but at a few stations also
upmost in the littoral. In this uppermost part jump-
ing amphipods were common.
The above survey shows that not all of the in-
fralittoral species found intertidally were restricted
there to the infralittoral fringe. On the contrary
many infralittoral species, which endure being
emerged for shorter or longer periods, went more
136
or less high up in the littoral region, where they
occurred together with littoral species, many of
which could stand being submerged for a shorter
or longer time and therefore also went down into
the infralittoral fringe. Some littoral species also
went up into the supralittoral region, where they
lived on or between terrestrial plants. At many
places bushes and forest were found down to or
even below the extreme high-water line, an observa-
tion also made by ALVEAL & RaMO (1977) at two
stations in Seno Reloncavi.
It was, however, often difficult to know for sure
whether a species should be characterized as in-
fralittoral or littoral, because the area below zero
as well as the lowest part of the intertidal area of-
ten could not be investigated so that the limits there
are unknown. Perhaps therefore some supposedly
littoral species, the lower limit of which was not
observed, rather are infralittoral ones, going high
up in the littoral. Only those intertidal species,
which had there lower limit well above zero, could
with certainty be characterized as littoral. This
doesn't exclude that even these at some places and
under special circumstances could be found lower
down, for instance at M139, where Chthamalus cir-
ratus was taken in the infralittoral fringe, and in
Estero Reloncavl, where the littoral Elminius kingi,
Acanthocyclus albatrossis, Mytilus edulis chilensis,
and Enteromorpha bulbosa were found also in the
infralittoral, probably as a result of the low surface
salinity in the fjord.
On the other hand some species, supposed to be
infralittoral and the lower limits of which are not
known, could be littoral ones going down into the
infralittoral fringe. What matters is, however, not
how the species are classified. It is their distribution
on the shore which is of interest from an ecological
point of view and for comparison of the vertical
distribution of different species or the distribution
of a certain species at different localities.
The infralittoral fringe was the most diverse area.
There the shore became emerged only for a short
time at extreme low-water, the food supply was
good, and infralittoral and littoral species oc-
curred together. It is the area of soft-skinned ani-
mals as sponges, sea-anemones, echinoderms, and
ascidians, which cannot stand being exposed to the
air, though some sponges, sea-anemones, Pyura
chilensis, and many algae also went a little higher
up.
The decreasing numbe:' of species at higher lev-
els was not only a result o! longer periods of emer-
gence and exposure to sun, wind, and air tempera-
ture but also of less supply of nourishment, influ-
ence of trickling freshwater or rain, and differences
 in substrate. At the boulder stations the size of the
boulders decreased upwards, and small boulders
and pebbles are less attractive to animals than
larger ones.
At the higher levels, which are often emerged for
hours or even days, were found animals, which, as
acorn barnacles, limpets, and mussels, can close
their shells or press them against the substrate, in
this way trapping an amount of water between or
under the shells. Also some motile animals like
porcellanids, crabs, and snails, which can seek shel-
ter actively when the sea is out, were common high
up. Specimens of several marine species, especially
those living higest up, were found above their nor-
mal upper limit in depressions, crevices, water-filled
holes, under boulders etc., where the waves rolled
higher up than on the sides or where some water
was left or the substrate kept wet when the shore
emerged, or they were found under a sheltering
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cover of algae.
The uppermost part of the intertidal area was one
of Cyanophyta, lichen, and the lowest specimens
of terrestrial plants.
Vertical distribution of related species
When related species or such with similar biology
occur at the same locality and level on the shore
the reason may be that they occupy different
niches, have different day-rhythms, depend on dif-
ferent food, etc. It was therefore of interest to find
out if and to what degree such species co-existed.
For this parts of Fig. 22 can be used, but since its
partial figures are arranged according to the type
of vertical distribution of the respective species,
those to be compared arc~ not always figured near
each other, which makes it difficult to compare the
distributions. Moreover some of the partial figures
show the distribution not of single species but of
higher taxa. This drawback has been eliminated in
Fig. 24. There also the stations in Seno Reloncavf
and those in Ancud are figured separately, and
because of that the differences in tidal amplitude
in the two areas can be neglected. Since the species
to be compared live at the same station, thus under
the same environmental conditions, their levels on
the shore have been expressed in em above or be-
low zero. Each bar in the figure shows the vertical
distribution of a species at one station. Stations
M82 and M31 in, and M139 at the mouth of Estero
Reloncavi, have not been figured.
The 12 figures, which show the vertical distribu-
tion and abundance of the species of some impor-
tance occurring at each of the 12 stations, make it
easy to find which species do and which don't over-
lap at the different stations but make comparison
between the stations difficult. In Fig. 24 the verti-
cal bars showing the vertical distribution (but not
abundance) of each of the related species at all sta-
tions, are placed together. By comparing the bars
with the same station numbers one will see which
species overlap at a certain station and find the
degree of overlapping, if any.
Eleven species groups have been considered in
Fig. 24. For all these the zonation patterns seem
to be fairly similar in Seno Reloncavi and Ancud.
1. Cirripedia. Balanus psittacus occurred only in
the infralittoral fringe, B. flosculus in the lower
half of the shore, Chthamalus scabrosus in the
middle part, and C. cirratus and Elminius kingi in
the upper half. There thus was a distinct difference
in vertical distribution but much overlapping. The
only species which didn't overlap at any station
were B. psittacus on the one side and C. cirratus
and E. kingi on the other. Balanus laevis was only
taken on shells of B. psittacus.
2. Pagurus edwardsi and P. villosus were both
restricted to the lower part of the shore and over-
lapped at all stations, but the former had its main
occurrence a little higher up than the latter.
3. Smaller crabs and Petrolisthes laevigatus
seemed to have a similar biology. Halicarcinus plan-
atus was found in the infralittoral fringe, at some
stations also in the lower part of the littoral, where-
as Cyclograpsus cinereus occurred in the upper part
and didn't overlap Halicarcinus except at M37,
where a few specimens of Cyclograpsus occurred
together with the uppermost ones of Halicarcinus.
Acanthocyclus albatrossis, Petrolisthes laevigatus,
and Hemigrapsus crenulatus were found at all lev-
els, except lowest down, and overlapped at all sta-
tions, at some also with the two first-named spe-
cies, but Hemigrapsus went a little higher up than
Acanthocyclus and Petrolisthes. Petrolisthes mitra
and P. angulosus occurred lower down than P. lae-
vigatus, and Acanthocyclus gayi higher up than A.
albatrossis.
4. Polyplacophora did not occur in the upper
part of the shore except for stray specimens of a
few species. Tonicia elegans was only found lowest
down, Plaxiphora aurata, Tonicia atrata, and Chiton
latus in the lower third of the shore, where also
Ischnochiton pusio and Chiton granosus were found,
except lowest down. At some stations the three
last-named were found also in the middle part.
Ischnochiton imitator was taken within the distribu-
tion area of I. pusio, and Chaetopleura peruviana
had almost the same distribution as Tonicia ele-
gans. There thus was much overlapping among the
Polyplacophora species, but species overlapping at
one station did not always overlap at others.
137
 em em
700 700

600 600

500 500

400 400

300 300

200 200

100 100

em 57 em
10

r-lli-
300 300
5
200 200

100 ---- ----- 10---

56 100

a
X X X X X X • •
56
•
Downloaded by [UQ Library] at 18:20 01 September 2013

55
em em
300
10
S
55
56 10 57 57

1
5756
I 10
300

-~i i-----~- j---

57 5Ij0

Ir!i
200
200

lOa
X
-II
X X
_16 5_ _ 1_-_ 10
!~6__ i---I---
10
100

em Broken horizontal tines = MSL

700
59
X = The species was not found

il 'p ~ w~
91 91

It---I
37 at the localities studied
600 p 1 59
600

500
37
I 37
500
90

~:-I

~i
400 400
-

Ifj
300 300

200
200

100 -
22
100

Enterom Ulva Bostrychia Porphyr. Gelid. Chandr. Iridaea

bulbosa lact. har.lhookeri columb. pseudo. canal. cris.1 cil iat.

em em
300 300

200 200

100 100
X x Fig.24. Vertical distribution of some
related species.

138
 5. Of the hemisessile Gastropoda Crepipatella
dilatata and Nacella magellanica occurred in the
iower half of the shore, but Nacella was missing
lowest down and also went a little up in the upper
half. The only record of Nacella aenea was found
below N. magellanica, and Crepidula plana occurred
in the lower part of the distribution area of Crepi-
patella. Patelloida zebrina occurred in the middle
part, where also P. ceciliana had its main occur-
rence.
The siphonarians, which may comprise more than
one species, were taken in the upper part, at some
stations also in the middle. Crepipatella and the si-
phonarians didn't overlap in Seno Reloncavi, except
at M91, but had a wide overlapping area at all
Ancud stations. Crepipatel/a, Nacel/a, and Patel/oi-
da ceciliana overlapped at all stations in Seno Re-
loncavi, and Crepipatel/a and P. ceciliana at all in
Ancud. P. zebrina overlapped P. ceciliana at two
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and siphonarians at three Ancud stations. The Fis-
surel/a species (not figured) were mainly living low
down, but the specimens were too few to permit
reliable comparison of the distribution at different
stations.
6. Of motile Prosobranchia Nassarius gayii and
Nassarius dentiferus occurred together in the lower
third of the shore and overlapped Tegula atra and
Nucella calcar and at one station Littorina arauca-
na. Nucella and Tegula were found in the lower and
middle parts, Monodonta nigerrima and Littorina
in the middle and upper parts, but all four species
overlapped. However, below 200 the first two (in
Seno Reloncavi) were alone, above 550 the last-
named two. The few specimens of Littorina peruvia-
na were taken in the m,iddle part within the area
occupied by L. araucana, and Prisogaster niger
overlapped Tegula.
7. Bivalvia. Aulacomya ater and Ostrea chilensis
lived up to about MSL, and Mytilus edulis chilensis
in the lower and middle parts of the shore, at some
stations though reaching the upper part, where
Brachidontes purpuratus and Lasaea petitiana had
their main occurrence. Hormomya granulata (not
figured) was taken in the lower and middle parts
of the shore. Of the mytilids Mytilus. overlapped
Brachidontes at all stations and Aulacomya at four.
Aulacomya also overlapped Brachidontes at three
stations. Ostrea overlapped all three mytilids at four
stations, Aulacomya at six, and Brachidontes
at three. Though Lasaea probably has been over-
looked at some stations and levels, the many sam-
ples at hand indicate that overlapping with Aulaco-
my a and Ostrea is not common, whereas overlap-
ping with Brachidontes was observed at all stations
and with Mytilus at all but one.
8. Of the two common Chlorophyta Ulva lactuca
occupied the lower and middle parts and went a
little up in the upper, whereas Enteromorpha bul-
bosa in Seno Reloncavi was found in the upper
half of the shore, in the Ancud area at all five lev-
els, though not at all stations. Overlapping was
found in both areas except at M22. Ulva nemato-
idea was found in the lower part.
9. Bostrychia hookeri and B. harveyi at one sta-
tion occurred together in the upper half of the
shore, the former though mainly below the latter.
At another station B. hookeri had its upper limit
where B. harveyi had its lower.
10. Rhodophyta (except Idridaea and Bostry-
chia). Chondrus canaliculatus was only found in the
middle part, Gelidium pseudointricatum and Por-
phyra columbina in the middle and upper parts.
The other Gelidium species occurred together with
G. pseudointricatum. Chondrus overlapped Gelidi-
um at two and Porphyra at four stations, and Gelidi-
um overlapped Porhyra at six.
11. Iridaea ciliata was restricted to the lower and
middle parts, I. crispata to the middle. I. boryana,
membranacea, undulosa, and dichotoma occurred
at the same level as I. ciliata, I. dichotoma also
together with I. crispata. The samples of the Iridaea
species are few and no species occurred at more
than three stations. All that can be said at present
is that I. ciliata only overlapped I. crispata at one
station and that I. ciliata overlapped I. boryana at
one station but not at the other one, where both
occurred.
From the above is seen that most of the related
species considered overlapped and that the overlap-
ping areas often were considerable, but if one looks
at the main area of occurrence related species most-
ly preferred somewhat different levels. There were,
however, not many distinct belts where one species
dominated.
COMPARISON WITH THE ZONATION PATTERNS
DESCRIBED BY ALVEAL & ROMO
Species composition
In their description of the zonation patterns in 1971
at four littoral stations in Seno Reloncavi, ALVEAL
& ROMO (1977), in this chapter cited as A & R,
only mention 6 animals, Chthamalus cirratus, Balan-
us psittacus, Tegula atra, Littorina araucana, Mytilus
chilensis (my M. edulis chilensis) , and Perumytilus
(Brachidontes) purpuratus, but 22 algae, 2 of which
with 2 and 1 with 3 'forms'. In addition are mentio-
ned 3 algae determined to genus only, 2 lichen, and
about 10 land plants, 5 of which determined to spe-
cies.
139
 At those four of my stations in Sen~ Reloncavi,
which are situated near those of A & R, namely
M91, M59, M90, and M37 (see Fig. 1), about 80
large animals, which hardly can be overlooked, and
unidentified spirorbids, amphipods, siphonarians, a
'peje sapo', and a great number of smaller, both
identified and undertermined animals were found.
Of algae about 40 species and some unidentified
ones were taken, thus about twice as many as men-
tioned by A & R. The number of land plants is
about the same in both investigations.
Since it is not likely that so many species should
have disappeared during the 23 years between my
and their investigations, the reason for this great
difference in number of species may, at least part-
ly, be that my studies were devoted to both ani-
mals and plants, whereas A & R say that their stud-
ies concerned the marine flora. However, since they
nevertheless mention and figure the vertical distri-
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bution of 6 animals, which are as important for
characterizing the different levels on the shore as
are the algae, one may wonder why they didn't
consider any of the many other animals, which are
equally important in this respect, as for example
Nacella magellanica, Patelloida ceciliana, Crepipa-
tella dilatata, siphonarians, Aulacomya ater, Ostrea
chilensis, and Pyura chilensis. Perhaps the two au-
thors did find them, without discussing them, for
one of course need not mention all species found
if some suffice to give a good picture of the zona-
tion.
All 6 animals and 10 of the 22 algae mentioned
by A & R were also found at my stations in Seno
Reloncavi. Regarding the species only found in one
of the investigations, there are some records, or
missing records, which are difficult to explain. For
example I didn't find Enteromorpha compressa
GREV. and E. intestinalis LINK., which A & R found
at 3 and 4 stations respectively, whereas they don't
mention E. bulbosa, found at 11 of my 12 stations.
They mention Gelidium pusillum from all 4 sta-
tions, a species which I only found in Ancud, but
they don't mention G. pseudointricatum, which was
found at all my stations in Seno Reloncavi and at
2 in Ancud. Similarly they found Bostrychia mixta
HK. f. & HARV. at 2 stations but don't mention B.
hookeri and B. harveyi, which were common at 4
respectively 2 of my nearby stations. In these and
some other cases one may wonder if some species
can have been incorrectly identified.
Zonation patterns
Fig. 25 shows that the identified algae and animals
common to the stations of A & R and my nearest
stations had fairly similar vertical distribution with
140
especially good agreeement between the boulder
stations Punta Quillaipe and M37 (Fig. 25D). In
some cases, however, the common species went
higher up or lower down at the stations of A & R
than at my neighbouring stations, in other cases the
opposite was found, and in some cases their, in
others my species had a wider or narrower vertical
distribution. There thus was no general trend in the
differences found and these were not greater than
between my stations (cf. Figs 22 and 24) and than
can be expected as a natural result of local differ-
ences in environment, collecting conditions and pro-
cedures etc.
A & R distinguish three main zones at all their
stations, Zona Marina, characterized by infralittoral
species, which also occur in that lowest part of the
intertidal area, which is submerged most of the
time, Zona Hidrolitoral, characterized by intertidal
species, and Zona Terrestre, characterized by ter-
restrial plants. At Bahia Huelmo and Punta Quil-
laipe A & R (figs 4 and 2) also distinguishh a Zona
Geolitoral, characterized by the lichen Verrucaria
sp. and Caloplaca sp.
A & R don't present any exact measurements
of the zone limits and the distribution of the spe-
cies, but I have calculated all limits for my Fig. 25
from their figs 2-5, in which the limits are marked.
No levels are exact, for the figures of A & R have
so small scales that an error in the calculations of
10-20 cm may well occur.
As seen from their figures but most clearly from
my Fig. 26 there is a considerable overlap between
the four 'zones' of A & R, in some cases 1-2 m
or more. Zones are belts on the shore, which can
have different situations and widths at different
places, because of different environmental condi-
tions, but two 'zones' cannot exist at the same level
at the same place but, as STEPHENSON & STEPHEN-
SON (1949:290) say 'one above the other'. They thus
must follow each other from below upwards, with-
out overlapping. If overlapping occurs it is the ques-
tion of wrongly placed zone limits or of transition
areas. Thus when A & R describe zones with differ-
ent species composition as occurring at the same
level, they in fact are not describing zones but sim-
ply assemblages of species of different origin, which
coexist in a zone, at least for part of their vertical
distribution.
Fig. 2 of A & R of the zonation at Punta Quillai-
pe (see also Fig. 25D) may serve as an example.
In this Rhizoclonium tortuosum is said to belong to
Zona Hidrolitoral and Tegula atra to Zona Marina,
though both for a couple of metres occur at the
same level. Their fig. 2 also shows 17 species in
Zona Hidrolitoral, but within that zone are also fig-
 E
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Downloaded by [UQ Library] at 18:20 01 September 2013

o 0"
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700 .-:: Q.. 700

600 600

500 500

400

300 300

200

100
C o

Fig. 25. A. Vertical distribution, in cm above or below zero, of species common to Bahia Huelmo (black bars) and
Stn M91 (open bars), from fig. 4 in ALVEAL & RaMo (1971) and my Fig. 5. - B. The same for Bahia Chincui (black
bars) and Stns M59 (open bars) and M90 (hatched bars), from fig. 3 in A & R and my Figs 8 and 7. - The same for
Piedra Azul (black bars) and Stn M37 (open bars) from fig. 5 in A & R and my Fig. 9. - D. The same for Punta
Quillaipe (black bars) and M37 (open bars) from fig. 2 in A & R and my Fig. 9.

Fig. 26. Comparison between the

'zones' described by ALVEAL &
RaMo (1977) and my corresponding
regions and zones. A. Bahia Huel-
rna + M91. B. Bahia Chincui + 0
M90 and M59. C. Piedra Azul + em
000
M37. D. Punta Quillaipe + M37.
The four (in Band C three) left 900
bars show the four/three overlapping 800
zones of A & R, Zona Marina
(black), Zona Hidrolitoral (dotted), 700
Zona Geolitoral (hatched), and 600
Zona Terrestre (white). The bars
(marked with an asterisk) with not 500
overlapping zones show how I would 400
have delimited the zones at the four
stations of A & R, and the last (in 300
B two last) bars show the regions/ 200
zones at my nearest stations, infralit-
toral fringe black, littoral region 100
dotted, and supralittoral region
white.

141
 ured 4 species referred to Zona Marina, 2 to ZO!1a
Geolitoral, and 1 to Zona Terrestre. Those speci-
mens of these species, which occur at the level of
Zona Hidrolitoral, of course also belong to this
zone, whereas the remaining specimens of these
species belong to the neighbouring zones.
However, let's for the comparison with my zones
follow the terminology of A & R and accept their
Zona Marina, Zona Hidrolitoral, and Zona Terres-
tre, though considering all species found within the
limits of a zone as belonging to that zone whether
they are confined to the zone or also occur in other
zones. Zona Geolitoral is disregarded here because
its lower part overlaps with the upper part of Zona
Hidrolitoral and the upper part with the lower part
of Zona Terrestre (except between 650 and 680 cm
at Bahia Huelmo; see Fig. 26A). The overlapping
thus is practically total and I see no reason why the
two lichen cannot be referred to their Zona Terres-
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tre, the more so as A & R themselves in fig. 3 from
Bahia Chincui have referred them to that zone.
This may, however, be due to an oversight, for the
bars representing the two lichen are closely hat-
ched, as in figs 2 and 4, where the lichen are refer-
red to Zona Geolitoral, and not black as for all
species from Zona Terrestre. However, such a pos-
sible oversight doesn't change my reason for consi-
dering Zona Geolitoral as superfluous.
Since A & R are not treating zones but overlap-
ping assemblages of plants and animals, the limits
of these assemblages have to be replaced by new
zone limits to permit a comparison with my zones.
As mentioned before such limits have to be placed
at the levels at which the greatest changes in spe-
cies composition and abundance are found. To ex-
emplify this we can return to fig. 2 in A & R on
the zonation pattern at Punta Quillaipe. Without
having seen the station it is difficult to know exact-
ly where to place the zone limits. However, judging
from the figure the limit between Zona Marina and
Zona Hidrolitoral could be placed at 80 (cm), whe-
re A & R placed the lower limit of Zona Hidrolito-
ral, but the species composition rather points to 130
or perhaps 200 as a more natural limit. In the first
case Zona Marina would be characterized by Mac-
rocystis pyrifera, Iridaea ciliata, Viva lactuca f. laci-
nulata?, Lithothamnion sp., Polysiphonia abscissa,
and the lowest specimens of Tegula atra. If the limit
is placed at 130, Zona Marina in addition would
be populated by the lowest specimens of Pylaiella
litoralis from Zona Hidrolitoral of A & R. With the
limit at 200 the lowest specimens of 3 more species
from Zona Hidrolitoral would belong to Zona Mari-
na.
If we treat Zona Geolitoral as part of Zona Ter-
142
restre, the limit between the latter zone and Zona
Hidrolitoral at Punta Quillaipe (fig. 2) could be
drawn at 700 or 720. At 700 A & R placed the
lower limit of Zona Terrestre, at 720 the upper limit
of Zona Hidrolitoral. Since one cannot permit over-
lapping, 700 seems to be the best choice as limit.
There Chthamalus cirratus had its upper and
Caloplaca sp. and Sphagnum sp. their lower limit.
Zona Terrestre will then be characterized by higher
plants, ferns, mosses, the lichen Verrucaria sp. and
Caloplaca sp., and in addition the uppermost speci-
mens of Rhizoclonium sp., Blidingia minima, and
Porphyra columbina, which three species according
to A & R belong to Zona Hidrolitoral. On the
other hand the lowest specimens of Verruca ria sp.
will be found in Zona Hidrolitoral.
Similar considerations can be applied to the other
three stations, only that, as A & R themselves state,
there it would not be possible to suggest exactly
where to place the limit between the two lowest
zones, for between about 100 and 200 there is a
50 to 100 cm wide interruption in the bars (A & R
figs 3-5), which illustrate the vertical distribution
of the species. This gap is probably also the reason
why some other species have their upper or lower
limits at this level, limits which probably are too low
or too high. In their fig 3 (Bahia Chincui) is stated
that no samples were taken at this level. This ex-
plains the interruption in the bars in this figure.
Such a remark is lacking in figs 4 and 5, but it is
likely that the explanation in all three cases is the
same, for the figures show that all gaps are found
at a level at which there was a belt of sand bottom.
No gap is shown in fig. 2 from Bahia Quillaipe,
where the bottom throughout consisted of boulders
and stones.
Both investigations thus show that one can distin-
guish three main zones/regions, and within these
many of the same species occur. The Zona Marina,
Hidrolitoral, and Terrestre of A & R correspond
to my infralittoral fringe and littoral and supralitto-
ral regions. In Zona Hidrolitoral A & R distinguish
three subzones, Hidrolitoral inferior, medio, and
superior. In my investigation also three subzones..
were observed in the littoral region, named, from
below, zones 1, 2, and 3. They could equally well
have been named lower, middle, and upper zone.
In my field notes was often remarked that a spe-
cies was 'association-forming', with which was
meant that it formed a distinct belt on the shore,
for example a Chthamalus or a Mytilus belt. It
was, however, also clear that within the area cov-
ered by one such association often were other 'asso-
ciations', occupying the whole or parts of the area.
The associations thus overlapped instead of forming
 distinctly separated belts on the shore, one above
the other. This made me refrain from trying to
describe associations. Much more detailed studies
are necessary before this will be possible, at least
where one species isn't alone. A & R have, how-
ever, described a dozen associations, an attempt
that seems premature.
With associations A & R mean 'populations
which always occur in similar conditions and in
which a few species are distinctly dominant' [my
translation]. Their associations have been named
after one to three dominant algae. The other spe-
cies mentioned as occurring in the associations are
also algae, except in one case where Chthamalus
cirratus and Littorina araucana are said to occur in
the Porphyra columbina association. At Bahia
Huelmo this association is only 60 cm wide, at Pied-
ra Azul 600 cm. Is it possible that the conditions
in these two cases are 'similar' and that it is the
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question of the same association, which moreover
in the first place has its upper limit 150 cm below
Zona Terrestre, in the other comprises this zone?
Another association of A & R is the Rhizoclon-
ium-Blidingia association, characterized by Rhizo-
clonium sp. and Blidingia minima and found at
Punta Quillaipe and in Bahia Chincui. At the latter
station Bostrychia mixta is substitute for Blidingia.
Can one then speak of a Rhizoclonium-Blidingia
association at that station, and is it advisable to
name an association after an unidentified species?
In their Bostrychia mixta-Gelidium association
the characteristic species are B. mixta and G. pusil-
lum. It is found at Piedra Azul, but in the figure
of the zonation there Bostrychia is not found, and
Gelidium occupies a narrow belt within the Porphy-
ra association.
These three examples suffice to emphasize my
opinion that it is premature to describe associations
and that these have to be defined in another way.
At least one must expect that the species after
which the associations are named really occur there
or play an important role in the whole association,
not only in a minimal part of it.
SPECIES COMPOSITION AND ZONATION
PATTERNS IN OTHER PARTS OF CHILE
Earlier investigations
In addition to my investigations in the Seno Relon-
cavf and Ancud areas and those of ALVEAL & ROMO
(1977) in the former, zonation studies have been
'arried out in southernmost Chile by SKOTISBERG
1941) and KNOX (1960), in Estrecho de Magalla-
nes, Isla Takan, Valdivia, and Talcahuano by AL-
VARES STRAEHL (1964), in the Montemar area by
GUILER (1959b), ALVEAL (1970, 1971), ROMO &
ALVEAL (1977), and SANTELICES & aI., (1977), and
in Coquimbo, Antofagasta, Iquique, and Arica by
GUILLER (1959a).Though the number of places thus
is fairly high, the statement by VIVIANI (1975) that
little has been done and much remains to be done
in Chile, still holds good.
The investigations have been carried out in diffe-
rent ways and during different years and seasons,
and the results have been presented in different
ways and in more or less detail. Some papers only
or mainly concern algae, few give measurements
of the levels between which the species occur, and
the names and limits of regions/zonesibelts differ.
Often only the generic names are given, some spe-
cies names are synonymous, some determinations
are tentative, and the authors of the species are not
always mentioned. Because of these deficiencies
and since many species occupy different levels as
juveniles and adults, since others show daily or sea-
sonal differences in levels and some even don't oc-
cur yearly, it is evident that investigations even in
the same area can give different results, which may
be difficult to compare. The most useful informa-
tion is that referring not to the levels on the shore
but to presence and on the vertical order in which
the species occur.
VIVIANI'S paper (1975), based on studies mainly
in Iquique and Valdivia, is not a standard-type
description of zonation but a study of interrela-
tions between species mainly in the Lessonia and
Chthamalus belts. He also calls attention to the
often neglected fact that most intertidal investiga-
tions throughout the world have been made during
the day and when the shores are emerged, thus
when investigations are easiest to carry out and
when hemisessile and motile animals are more or
less inactive and found at restricted levels, often low
down in their much wider activity areas. If a shore
is studied only when it is emerged, one thus will
get the wrong impression that the species are con-
fined to the narrow levels where they are found
when the tide is out. The conditions are the same
as when the biotope of for instance some terrestrial
animals is said to be 'under stones', while the fact
is that they only spend their inactivity period there.
Our way of collecting debarred such mistakes.
Since all specimens were collected when in the
water-line and since collecting continued until the
day's high-water was reached, we could observe the
animals leaving their resting-places, when the water
reached them, and follow their upwards wandering,
with the rising sea, to the upper limit of their activi-
ty area. The bars in the figures of zonation thus
143
 show the total area of the respective species.
VMANI also emphasizes that many intertidal spe-
cies are active only during the dark hours, for which
reason their nightly and often extensive wanderings
and real vertical distribution are difficult to study.
The total distribution area of many such species
thus must be wider than observed during the day.
Moreover some night-active species, which may be
as important as character species as those studied
during the day, live hidden during the day so one
isn't even aware of their existence. Without night
studies the picture of the zonation is thus often frag-
mentary, and one cannot exclude the possibility
that the differences between day and night can be
as striking on the shore as on coral reefs, which
present themselves as two completely different
worlds during day and night. Since, however, the
zonation studies in Chile seem to have been carried
out during the day, the results are comparable but
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only show the day conditions. Many more studies
around the clock, at different seasons, and in diffe-
rent years are needed before one can get a true
picture of the zonation.
Chile south of 48° S
SKOTISBERG (1941) described the algae at several localities
south of 48° S. He distinguished a sublittoral and a litto-
ral region, at one locality also a supralittoral belt characte-
rized by land plants, Cyanophyta, and a few other algae.
He gave no levels for the species in the respective regions!
belts and no information on algae on rocks under the
heading 'sublittoral region', but when describing the 'litto-
ral region' he at some localities mentions algae as occur-
ring in the 'uppermost sublittoral', in the 'lower littoral-
uppermost sublittoral', or as taken 'at the low-water
line', 'at the waters edge' or 'at about low-tide mark'. In
all these cases it seems to be the question of a level more
or less corresponding to the infralittoral fringe. At this
level he describes associations of Adenocystis utricularis*,
Lessonia nigrescens, Macrocystis pyrifera, Durvillea ant-
arctica* (CHAM.) HARlOT, D. harveyi HK. f., Lithophyl-
lum* sp., and Lithothamnion* sp., and in addition menti-
ons Scytothamnus fasciculatus' and Ceramium involutum
Kfrrz. The species marked with an asterisk also occurred
in the lower belt of the littoral.
SKOTISBERG divided the littoral region in a lower and
an upper belt, at one locality also distinguishing a middle
belt, or he distinguished a 'middle lower' and an upper
belt, in which case one doesn't know which of the species
mentioned were found in the middle and which in the
lower part.
Keeping to the rock localities and omitting rock pools
the following species were most often recorded in the
lower belt, in addition to the afore-mentioned sublittoral
ones, which went a little up in the littoral: Cladophora
pacifica (MONT.) Kfrrz., Scytosiphon lomentaria, Chaetang-
ium fastigiatum, and Catenella fusiformis, the last two also
typical of the upper belt. Among others Cladophoropsis
brachyartra (SVEO.) BORG., several Cladophora species,
Pylaiella litoralis, species of Iridaea, Ceramium, and Poly-
siphonia were also taken.
144
In. the middle belt was an association of Porphyra um-
bilicalis (L.) J.AG., and Chaetangium fastigiatum (also
typical of the other two belts) and Iridaea caespitipes (S.
& G.) were also found there, in addition to possible spe-
cies at this level recorded from the 'middle-lower' belt.
The upper belt was characterized by Ulva lactuca, Cla-
dophoropsis brachyartra, Rhizoclonium riparium (domi-
nant), Cladophora subsimplex KUTZ., Porphyra umbilica-
lis, Chaetangium fastigiatum, Hildenbrandtia lecannellieri
(dominant), Catenella fusiformis, and Bostrychia mixta
(dominant) .
KNOX (1960) briefly described the vertical order in
which the species occur in the Magellan province. The
only locality mentioned is Isla Hermite farthest south in
Chile, about 56° S, so one must suppose that his descrip-
tion at least partly refers to this island. He distinguishes
an upper sublittoral zone between ELWS and MLWS, a
littoral zone between MLWS and MHWS, divided in lower,
middle, and upper littoral, and a supralittoral zone above
MHWS, divided in lower and upper supralittoral. He thus
does not use species limits but tidal levels as limits be-
tween the zones, irrespective of the width of the tidal
zone at different places, exposure conditions, etc.
His upper sublittoral is characterized by a crust of Lith-
ophyllum and Lithothamnion, and among others Desma-
restia species, Macrocystis pyrifera, Lessonia, and Cerami-
um rubrum.
The algal crust extends to the lower littoral, where
Durvillea antarctica dominates on exposed coasts. Also
characteristic are chitons and species of Fissurella and
Nacella, with Aulacomya ater forming'a dense cover local-
ly.
The midlittoral is characterized by Balanus sp., Elmini-
us kingi and, upmost, Chthamalus scabrosus. Siphonarians
are the dominant limpet-like species and Mytilus edulis
chilensis the characteristic mussel species. Ulva lactuca,
Adimocystis utricularis, Scytosiphon lomentaria, Scyto-
thamnus fasciculatus, Porphyra umbilicalis, Ceramium,
rubrum and some southern species were also seen. In this
zone and the two below there is a turf of mixed algae
with Iridaea species dominating in the midlittoral.
In the upper littoral Chthamalus scabrosus dominates
together with Siphonaria lateralis GOULD and Brachidontes
purpuratus replaces Mytilus. Most conspicuous is a carpet
of Bostrychia mixta on exposed and B. scorpioides MONT.
on more sheltered shores, both species extending up in
the lower supralittoral. Associated are species of Rhizoclo-
nium, Cladophoropsis, and Cladophora (these three also
found in the midlittoral), Chaetangium fastigiatum, and
Catenella fusiformis.
The lower supralittoral is blackened by Verrucaria and
blue-green algae on which patches of Hildenbrandtia le-
cannellieri occur. This species often extends down in the
upper littoral. According to KNOX the most conspicuous
features of the Magellanic region are pink, orange (Calo-
placa sublobulata) and, upmost, white bands of lichen in
the upper supralittoral.
ALVAREZ STRAEHL (1964) studied four hard-bottom loca-
lities at 53° S in Estrecho Magallanes, only 3° north of
Isla Hermite. She distinguishes the same three main zones
as KNOX, but in the two cases she divides 'mesolitoral' she
distinguishes two portions only, 'inferior' and 'superior'.
In her 'franja sublitoral', below about 100 cm, she re-
cords among others Parantheopsis cruentata*, Phragmato-
poma* sp., Chthamalus scabrosus, Balanus flosculus',
Halicarcinus planatus* , Acanthocyclus albatrossis*, Plaxi-
phora aurata, Fissurella oriens, F. picta, Nacella magellani-
ca*, Patelloida ceciliana*, Nucella calcar, Brachidontes
 purpuratus, Anasterias antarctica (LfrrKEN), Pseudechinus
magellanicus, Ulva" sp., Macrocystis pyrifera, Adenocystis
utricularis" , Lithothamnion sp., Lithophyllum sp., and
Iridaea sp. Species marked with an asterisk were also
found in the next zone.
In the mesolitoral inferior, about 100-130/140 according
to her figures, she mentions Antholoba achates, Tonicia
lebruni ROCHEBRUNE, Porphyra sp., Ceramium rubrum,
and the species marked above.
In the mesolitoral superior were found Nacella mag-
ellanica, Laevilittorina caliginosa (GOULD), Siphonaria les-
soni (BLAINVILLE), S. lateralis (GOULD), Mytilus patagoni-
cus (ORBIGNY), Brachidontes purpuratus, Lasaea petitiana,
Adenocystis utricularis, Porphyra sp. Ceramium rubrum,
and Polysiphonia sp.
In her 'franja supralitoral' were found Laevilittorina
caliginosa, Siphonaria lateralis, Lasaea petitiana, and in
the summer Porphyra sp.
Species mentioned in her text as occurring in a special
zone are not always illustrated in the figun,s as present
in that zone, and the vertical distribution shown in the
figures representing the four localities is not always the
same as in the figure summarizing the conditions at the
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stations. This lack of clarity and the fact that the limits
used between the zones by the three authors are not quite
comparable makes comparison difficult. The animal
species enumerated by ALVAREZ STRAEHL are only partly
the same as those recorded by KNOX, and the few algae
she mentions are only partly the same as those recorded
by SKOTISBERG and KNOX, but in spite of the many uncer-
tainties the species mentioned with full names by the three
authors seem to occupy approximately the same levels at
all localities. There is thus nothing in the descriptions by
the three authors, which indicates any principal differ-
ences in species composition and zonation between the
localities in southernmost Chile, which isn't unexpected
because of the relatively small distance between them and
the uniform environment in the area.
One difference is, however, worth mentioning. While
KNOX stated that littorinids were completely absent, ALVA-
REZ STRAEHL recorded Laevilittorina caliginosa from the
mesolitoral superior and the franja supralitoral. Perhaps
this difference is local and a result of different exposure
conditions. Also in other places in Chile littorinids are
lacking locally. In her fig. 18, summarizing the results from
her four stations, Littorina araucana is figured, probably
by a slip of the pen, for she doesn't mention this species
in the text or those figures on which fig. 18 is based, and
it is not known south of 44° S (see BRAITSTROM & JOHANS-
SEN 1983). In fig. 18 she also figures Mytilus chilensis,
which might be another slip of the pen, for the. only speci-
es mentioned in the text or in the other figures is Mylilus
patagonicus ORBIGNY.
The Chiloe area
According to SKOITSBERG (1941), the lower belt of the lit-
toral region at Isla Huafo, south of Chilbe, was domina-
ted by Iridaea boryana, the upper by Bostrychia sp., Cla-
dophora sp., and Scytosiphon lomentaria. At nearby Isla
San Pedro the lower belt was characterized by Lithophyl-
lum skottsbergi LEM., the upper by Hildenbrandtia lecan-
nellieri with Catenella fusiformis.
In Bahia de Ancud at the north coast of Chiloe the
lower belt was dominated by Entermorpha bulbosa and
the middle and upper belts by a form of E. bulbosa,
Rhizoclonium riparium, and Bostrychia mixta. Other spe-
cies at the two lastnamed levels were Cladophora sp.,
Scytosiphon lomentaria, Scytothamnus australis HK. f. &
flARV., Catenella fusiformis, Gigartina glomerata HowE,
and Bostrychia scorpioides. Only few of these were taken
at my localities in Bahia de Ancud.
KNOX (1960) also gave some information from Chiloe,
without mentioning localities. In the upper sublittoral
Macrocystis pyrifera dominated with an undergrowth of
Lithothamnion, Halopteris, and red algae.
The lower littoral was characterized by Durvillea antarc-
tica and Lessonia nigrescens with Lithothamnion, Corall-
ina, and red algae as undergrowth. Large chi tons, Fissurel-
la species, and Concholepas concholepas were the domi-
nant animals.
Iridaea laminarioides BORY dominated in the midlittoral
superimposed on barnacles, Mytilus edulis chilensis, and
Brachidontes. Lowest down was an algal turf of Ulva,
Adenocystis, Scytosiphon, and Colpomenia. Chitons and
species of Fissurella were abundant in crevices.
In the upper littoral a Chthamalus sp. dominated and
limpets and siphonarians were abundant. Bostrychia was
more scattered than in the south. Littorina peruviana do-
minated in the lower supralittoral with a band of lichen
above.
ALVAREZ STRAEHL (1964) investigated three localities at
Isla Talcan, east of Chiloe, about 42°45' S. In her franja
sublitoral Chaetopterus variopedatus* at one locality for-
med dense 'colonias'. Among other species at this level
she mentions Spirorbis sp., Betaeus truncatus, Pagurus
forceps H. MILNE EDWARDS, Halicarcinus planatus", Plax-
iphora aurata*, Ischnochiton pusio", Fissurella costata*
and F. oriens*, Nacella magellanica*, Crepipatella dilata-
ta*, Trophon geversianus (PALLAS), Ostrea chilensis*,
Bryozoa, Anasterias antarctica, Ophiactis asperula, Ar-
bacia dufresnii, Loxechinus albus", Enteromorpha sp.,
Ulva sp., and upmost, Ceramium rubrum. Species marked
with an asterisk also occurred in the lower mesolitoral.
In the lower mesolitoral or part of it she also mentions
Bunodactis hermafroditica*, Chthamalus cirratus*, Bala-
nus psittacus, Acanthocyclus albatrossis, Chiton granosus,
Patelloida ceciliana*, Monodonta nigerrima", Tegula atra,
Littorina araucana, Nucella calcar, Mytilus chilensis*, Bra-
chidontes purpuratus, Ulva sp., and Iridaea laminario-
ides", those marked with an asterisk also occurring in the
upper part of the mesolitoral, where the most characteris-
tic species were Chthamalus cirratus, Balanus psittacus
(see next subchapter), and Brachidontes purpuratus. Other
species were Halicarcinus planatus and Hemigrapsus cre-
nulatus.
It is difficult to compare the zonation of the algae at
the localities in the Chiloe area from the meagre informa-
tion given by SKOITSBERG, KNOX, and ALVAREZ STRAEHL,
for whereas KNOX distinguishes three littoral zones the
other two only distinguish two, and KNOX and ALVAREZ
STRAEHL refer the distribution to tidal levels, but not exact-
ly the same. The levels of the three authors are thus not
fully comparable. Since moreover many algae were deter-
mined only to genus, one doesn't know if it is the questi-
on of the same species at the different localities. In fact
only one alga determined to species was recorded by
more than one author. The species composition at the
different localities therefore seems to be much more diffe-
rent than it probably is, for the same genera are mentio-
ned by two or all three authors. Thus the Scytosiphon
mentioned by KNOX in the midlittoral could be identical
with S. lomentaria recorded by SKOITSBERG, and the Bos-
trychia, which KNOX mentions in the upper littoral, could
be B. mixla or B. scorpioides, which SKOITSBERG found
in the upper belt. The most one can get out of the in-
145
 formation at hand is the vertical order in which the fully
named species appear at the respective localities.
Nor is it easy to compare the vertical distribution of the
animals as described by KNOX and ALVAREZ STRAEHL.
Both mention large chi tons and Fissurella species in the
lower littoral, but it is not possible to find out if the le-
vels are quite comparable and the species the same. ALVA-
REZ STRAEHL recorded Chthamalus cirratus and Brachidon-
tes purpuratus in the upper mesolitoral and KNox a Chtha-
malus sp. there, which can be cirratus.
ValdivialMehuin
The tidal amplitude in Valdivia at 39°48' Sis 119 cm (AR-
MADA DE CHILE 1949). The earthquake in 1960 caused the
shore to subside about 1.6 m (PARKER 1960). The whole
intertidal area and the area above, earlier washed by
waves, thus became inundated (ALVAREZ STRAEHL 1964).
Her studies and those by KILIAN (1961) of the same locali-
ty in Mehuin, after the earthquake, made it possible to
study the colonization of the new intertidal area by ma-
rine organisms. . ties was as follows:
In 1961 KILIAN found a mass occurrence of a Sabellaria
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in the infralittoral fringe, according to ALVAREZ STRAEHL
probably being Phragmatopoma peruensis HARTMAN,
which she found to be abundant in that zone both in 1961
and 1963. Already seven months after the earthquake
Fissurella sp., Concholepas concholepas, Nucella calcar,
Semimytilus algosus (GOULD), and Iridaea sp. were found
in the new franja sublitoral.
In January and August 1961 a few Brachidontes purpur-
atus were found in the upper part of the mesolitoral, at
which time Semimytilus dominated also in this zone. The
two mytilids compete for space, and when in 1963 juveni-
les of Brachidontes dominated in the upper part of the
littoral, Semimytilus had been restricted to the franja subli-
toral. Littorina araucana was fairly abundant in the whole
franja supralitoral in January 1961.
The zonation pattern in 1963, about 2! years after the
earthquake, showed a franja sublitoral with Phragmatopo-
ma peruensis, Fissurella sp., Patelloida ceciliana, P. zebri-
na, Concholepas concholepas, Nucella calcar (lowest
down), Semimytilus algosus, Iridaea sp. and, in exposed
parts, Chthamalus scabrosus and Elminius kingi.
In the mesolitoral Phragmatopoma and Iridaea sp. oc-
curred lowest down and Brachidontes purpuratus in the
middle part. Chthamalus scabrosus, Elminius kingi, Or-
chestia sp., Dynamenella eatoni, Patelloida ceciliana, P.
zebrina, Littorina araucana, Siphonaria laeviuscula (So-
WERBY), and Porphyra sp. were found in the whole zone.
In the franja supralitoral the same species were found
as in the mesolitoral, Phragmatopoma, Brachidontes, and
Iridaea sp. excepted.
Also VIVIANI (1975) studIed the intertidal zone in Me-
huin, between 1958 and 1973. I have not found any refe-
rence in his paper to the earthquake in 1960, so I don't
know if his description refers to the conditions before or
after the coast subsided, but as he mentions the same
species as the other authors from approximately the same
levels, it seems likely that he has studied well-established
conditions.
Though VIVIANI mainly discusses the ecology of the
Chthamalus belt he also gives some information on the
levels at which the species occur. Balanus flosculus occurs
lowest down. Upmost it overlaps Chthamalus scabrosus,
which occurs in the middle part of the belt and in its turn
merges into a belt of C. cirratus, which only occurs in the
highest part of the intertidal area. Contrary to the condi-
tions in Talcahuano (see below), C. scabrosus at Mehuin
146
dominates over C. cirratus, because it is less sensitive to
lowered salinily than C. cirratus, which is less sensitive to
drought. The heavy rainfall in Valdivia thus favors C.
scabrosus. Other important species in the Chthamalus
belt are 'Acanthocyclus sp., 'Chiton granosus, various
Acmaea and Fissurella species, Littorina araucana, 'Con-
cholepas concholepas, 'Nucella calcar, Semimytilus algo-
sus, Brachidontes purpuratus, 'Sicyaces sanguineus (MUL-
LER & TROSCHEL), and sea birds (the species marked with
an asterisk were by VIVIANI considered key species).
Mytilys chilensis is said to be characteristic of the fjords
and 'canales' in the south. That single specimens were
taken in Mehuin can be explained by the low salinity
there, caused by much rain (VIVIANI 1975:101).
Talcahuanoffumbes
The two northernmost localities studied by ALVAREZ
STRAEHL (1964) were one on each side of Peninsula
Tumbes near Talcahuano, at 36°40' S, about 3° north of
Valdivia. In great outlines the zonation at the two locali-
In the franja sublitoral were recorded Phragmatopoma
moerchi KINBERG, Chaetopleura peruviana, Tonicia ele-
gans, Fissurella costata and F. crassa, LAMARCK, Tegula
atra, Prisogaster niger, Concholepas concholepas, Nucella
calcar, Hormomya granulata, Corella eumyota, Pyura
chilensis, Viva nematoidea, Codium difforme KliTz. ,
Lithothamnion sp., Lithophyllum sp., and Grateloupia
cutleriae KUTz.
Chthamalus cirratus and C. scabrosus, Brachidontes
purpuratus (not upmost), and Iridaea laminarioides were
found in the whole mesolitoral zone, the mussel being the
most characteristic species there.
In addition the following were recorded at different le-
vels in the narrow lower mesolitoral: Phymactis clematis,
Bunodactis hermafroditica, Phragmatopoma moerchi, Pet-
rolisthes laevigatus, Acanthocyclus albatrossis, Chiton gran-
osus, Fissurella crassa and F. costata, Patelloida ceciliana
and P. zebrina, Tegula atra and T. tridentata (POTIEZ &
MICHAUD), Prisogaster niger, Concholepas concholepas,
Nucella calcar, Hormomya granulata, Viva nematoidea,
Porphyra sp., Gymnogongrus sp., and Gelidium filicinum
(BORY).
Acanthocyclus albatrossis, Orchestia sp., Petrolisthes lae-
vigatus, Chiton grancsus, and Patelloida ceciliana were
found lowest down, Ulva nematoidea in the lower part,
Patelloida orbignyi (DALL) and Littorina araucana in the
upper part, and Onchidella marginata upmost in the up-
per mesolitoral zone. Siphonaria laeviuscula occurred in
the whole upper mesolitoral.
In the franja supralitoral Patelloida orbignyi and Littori-
na araucana dominated. The uppermost specimens of
Chthamalus cirratus and C. scabrosus, Chiton granos us,
Patel/oida ceciliana, P. orbignyi and P. zebrina, Siphona-
ria laeviuscula, and Iridaea laminarioides went high up in
the zone.
The Montemar area
According to GUILER (1959b) the infralittoral fringe at
exposed localities in Montemar, at about 33° S, is charac-
terized by Lessonia nigrescens, a few Durvillea antarctica,
and Lithothamnion. The fauna in clefts and Lessonia hold-
fasts is rich with, among others, Phymactis clematis, Bala-
nus psittacus, small decapod crustaceans, Chiton cumingsi
FREMBLY, C. granos us, Enoplochiton niger (BARNES),
Acanthopleura echinata (BARNES), Fissurella crassa, Con-
cholepas conchoiepas, Loxechinus albus, and Pyura chilen-
 sis. Because of the extreme exposure the infralittoral
fringe extends above MSL.
Lowest down in the midlittoral is a belt of mixed algae,
characterized by Centroeeras clavulatum, Gelidium filici-
num, and Corallina ehilensis. This belt merges into one
characterized by Chthamalus ?seabrosus, Aeanthocyclus
gayi, Pilumnoides perlatus (POEPPIG), Braehidontes purpur-
atus, and Iridaea laminarioides with Pyura ehilensis and
Ulva laetuea, the latter sometimes forming a belt above
the Braehidontes-lridaea belt. Colpomenia sinuosa and
Porphyra eolumbina occur in both belts.
Most of the midlittoral is occupied by a barnacle belt.
Lowest down in this Balanus laevis is the common species
and occurs together with B. floseulus. Higher up' follow
Chthamalus ?seabrosus and upmost C. cirratus. Within the
barnacle belt are two sub-belts, the lower one of Porphy-
ra eolumbina, the upper one of Littorina peru viana. Other
species are few and mainly found in clefts, for example
Leptograpsus variegatus (POEPPIG), Chaetopleura peruvia-
na, and Chiton eumingsi.
There is no supralittoral fringe and the supralittoral
most often is bare. Lichen are missing, probably as a re-
sult of bird excreta and too much spray on the low rocks.
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Leptograpsus may be found moving around in this high
part, too.
ALVEAL (1970, 1971) studied the same rocks in detail
and his results in great outlines harmonize with those of
GUiLER. Of interest is that ALVEAL mentions Littorina
araueana as occurring together with the uppermost speci-
mens of L. peruviana but mainly above that species.
GUiLER (1959b) and ROMO & ALVEAL (1977) found the
same zonation pattern in Bahia de Quintero, just north
of Montemar, but they also found Maeroeystis integrifolia
BORY in the infralittoral fringe in localities with less than
maximum wave exposure, and ROMO & ALVEAL mention
three species of lichen in their 'Zona Geolitoral'.
SANTELICES & al. (1977) have published a valuable re-
port on the organisms on one exposed and one sheltered
rocky-shore locality at Caleta Horc6n at 32°40' S, thus
just north of Bahia de Quintero and only 15' north of
Montemar. To my knowledge this is the first really quanti-
tative study of intertidal communities on rocky shores in
Chile. The pattern of vertical distribution at the two locali-
ties was, in general,. similar to that of the nearby areas,
described above.
Coquimbo
GUiLER (1959a) also studied the zonation at Coquimbo,
about 30° S, where the conditions were similar to those
in Montemar. The infralittoral fringe is characterized by
Lessonia nigreseens and lithothamnia. Balanus psittaeus,
Acanthopleura eehinata, Coneholepas eoncholepas, and
Tetrapygus niger (MOLINA) are found in this belt.
The midlittoral is characterized by a barnacle belt with
the same species, in the same ordl;r, as in Montemar.
Porcellanid crabs, chitons, Acmaea and Siphonoria spe-
cies, and mussels are common, among the latter Mytilus
chorus MOLINA and Brachidontes purpuratus. There is no
Littorina belt above that of barnacles but L. peruviana
formed a discontinuous belt within the barnacle belt.
Above this the rock was bare.
Antofagasta
Principally the species composition and zonation pattern
in Antofagasta at 23°40' S, studied by GUiLER (1959a),
are the same as in Coquimbo with only small local differ-
ences. GUiLER thus doesn't mention any large algae in the
infralittoral fringe, which is characterized by Litho-
thamnion sp., Enoplochiton niger, Aeanthopleura echinata,
Fissurella costata, F. limbata SOWERBY, and Concholepas
concholepas.
Above the Lithothamnion belt follows a belt of Pyura
ehilensis with Balanus psittacus, both overgrown by Ulva
lactuca in the lower and Corallina chilensis in the upper
part. Many small animals as Petrolisthes angulosus, Acantho-
eyclus gayi, Fissurella limbata, and Brachidontes pur-
puratus live between the tests of Pyura.
The barnacle belt here is one of Chthamalus cirratus,
in the lower part with Acmaea viridula (LAMARCK), Siphona-
ria lessoni, and other limpets, though no large ones.
Algae as Enteromorpha sp. and Colpomenia sinuosa play
a great role in the lower part and a strip of Littorina peru-
viana in the upper. Above the Chthamalus belt the rock
is bare.
Iquique
At Playa Blanca, Iquique, 20015' S, the rocks lowest
down, according to GUiLER (1959a), are covered' by
at least three lithotharnnia species, Lessonia nigra, Coralli-
na ehilensis, and other algae, for instance Glossophora
kunthii. Other species in this part of the shore are a.o.
Pachycheles grossimanus (GU¥R1N) Acanthopleura echina-
ta, Enoplochiton niger, Fissurella costata, F. limbata, and
Tegula atra.
Higher up Mytilus chorus and Brachidontes purpuratus
with Balanus flosculus and Acmaea sp. on their shells form
a belt together with Colpomenia sinuosa.
The barnacle belt is characterized by Chthamalus cirr-
atus with Balanus laevis in the lower part. At the exposed
Primeros Rocas nearby the barnacles are fewer but Ac-
maea viridula, Patelloida ceeiliana, Siphonaria lessoni, and
Ulva lactuca are common, as are some chitons in clefts.
At Playa Blanca Littorina peruviana formed a belt
above the barnacle belt, but at Primeros Rocas it occurred
in the barnacle belt, 'in what is considered to be the no[-
mal condition for the Chilean coast' (GUILER 1959a; cf.
Coquimbo).
Arica
At Arica, farthest north in Chile, 18"20' S, GUiLER (1959a)
found the lowest part of the shore being dominated by
lithothamnia. Here Phragmatopoma moerehi occurs. The
Lessonia belt is well developed with Petrolisthes angulo-
sus, Pachycheles grossimanus, and Gaudichaudia gaudi-
chaudi in the holdfasts. Other species in this belt are
Enoplochiton niger, Acanthopleura echinata, Fissurella
costata, Tegula atra, Concholepas concholepas, and Gigar-
tina lessoni (BERG) J.AG.
The midlittoral is dominated by barnacles. Lowest down
Chthamalus cirratus occurs together with Balanus laevis
and B. f/osculus, higher up with the latter species, and
highest up with Littorina peruviana, which in some places
occurs above Chthamalus. In the lower part of the barnac-
le belt Brachidontes purpuratus is common. Other species
there are Chiion cumingsi and C. latus, Acmaea sp., Si-
phonaria lessoni, Enteromorpha sp., and Ulva lactuca. The
conditions in Arica thus are fairly similar to those in Iqui-
que.
147

Comparison between the Seno ReloncavilAncud
areas and other parts of Chile
Since no surveyor rocky-shore zonation in Chile
seems to exist, and in order to facilitate a compari-
son between the zonation patterns in the Seno Re-
loncavi and Ancud areas and those in other parts
of Chile, the above summary of earlier zonation
studies (some mimeographed and not easy to find)
in Chile has been made extensive, enumerating the
more important species recorded in these studies
and which also were found in the Seno Reloncavi
and Ancud areas and which play a leading role in
characterizing the different levels at all localities.
In addition some few southern and northern species
have been mentioned, which were missing in the
Seno Reloncavi and Ancud areas and which contri-
but,e to characterizing the different levels on the
shore-in the south and in the north, giving the zona-
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tion'patterns there a somewhat different appearance
than in Seno Reloncavi and Ancud.
The few algae mentioned with full name by
SKOTfSBERG (1941), KNOX (1960), and ALVAREZ
STRAEHL (1964) in the Chiloe area and which also
were .found at my stations in the Seno Reloncavi
and Ancud areas, mostly occurred at corresponding
levels everywhere. Though these levels are not ex-
actly comparable, the relative levels and vertical
order of the same algae at all localities from Boca
del Guafo to Seno ReloncavilAncud seem to be
practically the same. This was expected since the
environmental conditions are almost identical in the
whole area.
There is, however, a possible difference between
Isla Talcan and Seno ReloncavilAncud regarding
the level of some green algae. According to ALVA-
REZ STRAEHL the 'franja que corresponde el sublito-
ral' at one of her stations was easy to identify by
the presence of Ulva and Enteromorpha species (at
one station Ulva also was taken in her lower mesoli-
toral). At my stations in Seno Reloncavi and Ancud
Ulva lactuca and Enterom()rpha bulbosa, the com-
mon species in the area of these genera, were found
in the infralittoral fringe and in the whole littoral
region, in which SKOTISBERG, too, found E. bulbosa
in Bahia de Ancud. Thus, if the Ulvaand Entero-
morpha species at Isla Talcan are the same as those
in Seno Reloncavi and Ancud, their very low situa-
tion at Isla Talcan could be a local phenomenon
(cf. below).
Regarding the animals practically all those men-
tioned from Isla Ta1can by ALVAREZ STRAEHL were
found in Seno Reloncavi, too, but in spite of the
short_distance between Ta1can' and Seno Reloncavi
many species, among them Bunodactis hermafroditi-
148
ca, Acanthocyclus albatross~, Nacella magellanica,
Monodonta nigerrima, Tegula atra, Littorina arau-
cana, and Ostrea chilensis had from 1-2 m lower
upper limits at Talcan than in Seno Reloncavi.
This probably is a result of the different tidal condi-
tions. The exact tidal amplitude at Isla Talcan is not
known, but the figures of ALVAREZ STRAEHL and the
amplitude at other places in the same area (ARMADA
DE CHILE 1949) indicate that it perhaps could be
as much as H-2 m less than in Seno Reloncavi, and
as the localities at Ta1can were in the small Estero
Talcan, which has a very narrow and sheltered en-
trance, they probably are very sheltered, so that the
upper limits of the animals there, like the algae
mentioned above, are determined more by the tidal
levels than by waves, splash, and spray as at the
more exposed stations in Sena Reloncavi.
A difference of another kind is that at Ta1can
Balanus psittacus, according to figs 19-22 in the
paper by ALVAREZ STRAEHL, was found between 1.5
and 4.8 m in the lower and upper mesolitoral, thus
much higher up than in the more exposed Seno
Reloncavi, where it was observed only between 0
and 2.1 m, thus in the infralittoral (fringe). Since
this species in other places, too, is a character spe-
cies of the lowest part of the shore, the levels at
Ta1can are questionable. In three of her figures the
same bar is used to demonstrate the distribution
of both Chthamalus cirratus and Balanus psittacus.
Because of that the upper part of the bar could
represent Chthamalus and only the lower part Bala-
nus psittacus, but in a fourth figure, no. 19, with
one bar for each species, both bars are equally long
and placed at the same level in the middle and
upper mesolitoral. It seems hardly likely that this
is correct. Perhaps the explanation simply is a dra-
wing error.
Appendix 1 and my zonation figures show that
practically all species recorded with full names by
SKOTISBERG, KNOX, and ALVAREZ STRAEHL in the
Magallanes area, and in the Chiloe area as well,
were found at my stations in Seno ReloncavilAncud
and there occupy about comparative levels as at the
localities farther south. Some subantarctic and
cold-temperate species in the south don't reach
north to Seno ReloncavilAncud, and some warm-
temperate species there stop north of Tierra del
Fuego, but apart from this there seems to be no
principal difference in species composition and zo-
nation patterns within the vast area, about 15 latitu-
de degrees long, from Isla Hermite in the south to
Seno Reloncavi in the north.
A comparison between Valdivia and Talcahuano
shows some differences which can be local and not
valid for the whole areas. For instance Elminius
 kingi, Mytilys chilensis, and Semimytilus algosus
were recorded at the Valdivia station but not at the
two at Tumbes, and Patelloida orbignyi, Ulva nem-
atoidea, Gelidium filicinum, Gymnogongrus sp.,
and Iridaea laminarioides at Tumbes but not in
Valdivia. Phragmatopoma peruensis in Valdivia was
replaced by P. moerchi at Tumbes. In Valdivia
Chthamalus scabrosus dominated over C. cirratus,
at Talcahuano the opposite was the case. For many
other species a comparison is impossible because
only their generic names are given. The general
impression is, however, that the zonation in the two
places is fairly similar and that the conditions are
not principally different from those in Seno Relon-
cavi and Ancud. Practically all species mentioned
by ALVAREZ STRAEHL, KILIAN, and VIVIANI from
Mehuin (Valdivia) and by the former from Tumbes
were found at my stations in Seno ReloncavilAncud
and also at corresponding levels, not surprisingly
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since Mehuin and Tumbes only are about 2 respecti-
vely 5° north of Seno Relonacvi/Ancud.
ALVEAL & ROMO (1977) state that 95 % of the
species they studied in the intertidal area in Seno
Reloncavi also occur in the intertidal in Montemar,
and GUILER (1959a) stresses the uniformity of zona-
tion at the places he investigated from Valparaiso
(Monte mar) to Arica, a distance of about 15 de-
grees latitude. The differences found between the
localities are slight, some even of a local character.
In the infralittoral fringe Lessonia nigrescens is the
dominating species, at Antofagasta Pyura chilensis
(which species also occurs in the fringe and low
down in the littoral in the Seno Reloncavi and
Ancud areas). The midlittoral is characterized by a
barnacle belt with Balanus psittacus lowest down,
sometimes accompanied by B. laevis and B. floscu-
Ius, and with Chthamalus cirratus higher up (as in
Seno Reloncavi). The character of the barnacle
belt changes, however, from place to place and
from below upwards dependent on which other spe-
cies occur in the belt, such as Mytilus chorus, Bra-
chidontes purpuratus, Colpomenia sinuosa, and
Corallina chilensis. Littorina peruviana, usually
found within the upper part of the barnacle belt,
at some localities in the north formed a separate
belt above the barnacles, in the supral1ttoral fringe.
From the scarce and partly fragmentary informa-
tion referred to above, there seems to be little dif-
ference in species composition between the shores
of southernmost Chile and the Seno Reloncavil
Ancud areas, between these and the localities north
to Montemar, and between Montemar and the
coast to Arica in the north. The conclusion thus is
that the species composition along the whole Chile-
an coast is fairly uniform. It is true that several
species found on the northern shores don't reach
farther south than between Valparaiso and Chiloe
or a little beyond, and that several southern species
stop south of Chiloe or between this island and
Valparaiso, and zoogeographical papers also con-
clude that there is a limit at about 42° S between
the Cold- and Warm-temperate regions or, in an-
other terminology, between the Magellanic and
Central Chilean provinces, with a transition area
between 42 and 33° S (see BRATTSTROM & JOHANS-
SEN 1983, figs 7 and 18). This is supported by what
has been mentioned above.
Since it is among the wide-spread species one
finds most of those charact .:rizing the different le-
vels on the shore, not only the species composition
but also the zonation patterns show great similari-
ties along the whole Chilean coast. The differences
observed both in species composition and zonation
pattern mainly depend on the occurrence of the
accompanying less widely distributed species and
are not much greater than can be found between
localities in the same area because of local differ-
ences in the environment, different ways of col-
lecting, etc.
It may seem remarkable with such great uniformi-
ty along a coast, about 38 latitude degrees long,
but considering the hydrographical and climatical
conditions in the whole area it is not unexpected.
North-flowing currents along the shore carry cold
water far north (see BRATTSTROM & JOHANSSEN
1983, fig. 6), and upwelling in central and northern
Chile also contributes to lowering both sea and air
temperature. During the winter the mean surface
temperature is about 5° C in the south and 14° C
in the north, in the summer 8 and 20° C respective-
ly. The corresponding air temperatures are about
3° C in the south and 17° C in the north in the win-
ter and 8 and 22° C respectively in the summer. It
is these relatively small differences in temperature
between south and north and between winter and
summer which permit so many shore species to
have a wide distribution and therefore to play a
role in the zonation in the whole country, resulting
in little variation in zonation patterns from Cabo
de Homos to Arica.
ACKNOWLEDGEMENTS
I thank the members of the expedition for their help both
in the field and in the laboratory, especially Professor
Nibaldo Bahamonde, Santiago, who assisted me in the
field work at all localities, and Mrs Ingrid Brattstrom,
Bergen, who typed the field notes and correspondence
and wrote all labels.
149
 During many years after the expedition, Mrs Inger
Toppe Haugsb0, Bergen, with a grant from Norges almen-
vitenskapelige forskningsrad, assisted me with the collec-
tions, correspondence etc. The Director of the Instituto
Hidrognifico de la Armada, Valparaiso, Captain Fernando
Espinosa, and the Acting Directors of that institute, com-
manders Patricio Figuera and Carlos Bari, have given me
important information on tidal conditions in Puerto Montt
and sent me copies of the tide-gauge registrations there
during 1948 and 1949.
Mrs Elin Holm, Bergen, made the drawings for this
article, Mr Goran Brattstrom, Askim, corrected the Eng-
lish, and Professor Jose Stuardo, Concepci6n, translated
the English abstract to Spanish and corrected some speci-
es names.
I thank all these persons and the many other unmen-
tioned ones, who ha·:e assisted me in one way or other,
for their kind and valuable help.
REFERENCES
Alvarez Straehl, Amada 1964. Aspectos eco16gicos de al-
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gunas ('reas intercotidales de la costa Chilena entre
los paralelos 36° y 51{' lat. sur. - Mimeographed
thesis, Univ. de Concepci6n. 144 pp.
A1veal, Krisler 1970. Estudios ficoecol6gicos en la regi6n
costera de Valparaiso. - Revista de Biologia Marina
14(1):7-88.
- 1971. El ambiente costero de Montemar y su ex-
presi6n biol6gica. - Revista de Biologia Marina
14(3):85-119.
Alveal, Krisler & Hector Romo 1977. Estudios de distri-
buci6n vertical de la biota costera en el Seno Relon-
cavi - Chile. - Gayana. Miscelanea (7):1-28.
Armada de Chile 1948, 1949. Tablas de mareas de la cos-
ta de Chile 194811949. - Departamento de la Nave-
gaci6n e Hidrografia.
Brattstrom, Hans 1980. Rocky-shore zonation in the San-
ta Marta area, Colombia. - Sarsia 65:163-226.
- 1985. Rocky-shore zonation on the Atlantic coast
of Panama. - Sarsia 70:179-216.
Brattstrom, Hans & Erik Dahl 1952. General account,
lists of stations, hydrography. Reports of the Lund,
University Chile Expedition 1948-49. 1. - Acta
Universitatis Lundensis N.F. Avd. 2. Bd 46(8):1-88.
Brattstrom, Hans & Arild Johanssen 1983. Ecological and
regional zoogeography of the marine benthic fauna
of Chile. Report no. 49 of the Lund University
Chile Expedition 1948-49. - Sarsia 68:289-339.
Garth, John S. 1957. The Crustacea Decapoda Brachyura
of Chile. Reports of the Lund University Chile
Expedition 1948-49. 29. - Acta Universitatis Lund-
ensis. N.F. Avd. 2 Bd 53(7):1-130.
Guiler, E.R. 1959a. Intertidal belt-forming species on the
rocky coasts of northern Chile. - The Papers and
Proceedings of the Royal Society of Tasmania
93:33-58.
1959b. The intertidal ecology of the Montemar
area, Chile. - The Papers and Proceedings of the
Royal Society of Tasmania 93:165-183.
150
Jara, H. Fernando & Carlos A. Moreno 1984. Herbivory
and structure in a inidlittoral rocky cdmmunity: a
case in southern Chile. - Ecology 65:28-38.
Kilian, E. 1961. Die Entwicklung der Gezeitenfauna nach
einem Seebeben an der pazifischen Kiiste. - Deut-
sche Zoologische Gesellschaft, Artikel 45, 6 pp, 2
figs, 1 map.
Knox, G.A. 1960. Littoral ecology and biogeography of
the southern oceans. - Proceedings of the Royal
Society B 152:577-624.
Larrain, Alberto P. 1975. Los equinidos regulares f6siles
y recientes de Chile. - Gayana, Zoologia
(35):1-189.
Moreno, C.A., John P. Sutherland & Fernando Jara 1984.
Man as a predator in the intertidal zone of south-
ern Chile. - Oikos 42:155-160.
Nilsson-Cantell, Carl-Aug. 1957. Thoracic cirripeds from
Chile. Reports of the Lund University Chile Ex-
pedition 1948-49. 31. - Acta Universitatis Lunden-
sis. N.F. Avd. 2. Bd 53(9):1-25.
Parker, L. Hernandez 1960. Catflstrofe en el Paraiso. -
Editorial del Pacifico S.A., Santiago de Chile. 198
pp. + 61 figs.
Petersen, e.G.Joh. 1913. Valuation of the sea. II. The
animal communities of the sea-bottom and their
importance for marine zoogeography. - Report of
the Danish Biological Station 21:42 + 68 pp, 6 pla-
tes, 3 maps.
Retamal, Marco Antonio 1981. Catalogo ilustrado de los
crustaceos decapodos de Chile. - Gayana. Zoologia
(44):1-110.
Romo, Hector & Krisler Alveal 1977. Las comunidades
del litoral rocoso de Punta Ventanilla Bahia de
Quintero - Chile. - Gayana. Miscelanea (6)
1977:1-41.
Santelices, B., J. Cancino, S. Montalva, R. Pinto & E.
Gonzalez 1977. Estudios ecol6gicos en la zona cos-
tera afectada por contaminaci6n del "Northern
Breeze". II. Comunidades de playas de rocas. -
Media Ambiente 2:65-83.
Skottsberg, C. 1941. Communities of marine algae in sub-
antarctic and antarctic waters. - Kungliga Svenska
Vetenskapsakademiens Handlingar. Tredje Serien
19(4):1-92, 3 plates.
Stephenson, T.A. 1939. The constitution of the intertidal
fauna and flora of South Africa. Part I. - The
Linnean Society's Journal- Zoology 40:487-536, 4
plates.
Stephenson T.A. & A. Stephenson 1949. The universal
features of zonation between tidemarks on rocky
coasts. - Journal of Ecology 38:289-305.
Thorson, Gunnar .. 1951. Zur jetzigen Lage der marinen
Bodentier-Okologie. - Verhandlungen der Deut-
schen Zoologischen Gesellschaft in Wilhelmshaven
1951:271-327.
Viviani, Carlos Antonio 1975. Comunidades marinas
litorales. - Publicacion Ocacional, Laboratorio de
Ecologia Marina, Iquique. 196 pp. (Mimeographed,
cited with permission of the author).
Accepted 5 April 1990.
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APJk,.JIX 1. Vertical distribution of all identified marine species at the 12 stations, except species found in sand and rock pools. Numbers in italics = distribution below zero. x =
level not noted. A ? before or after the levels = lowest or highest occurrences not noticed. Brackets around depths show that the species was taken somewhere between the two levels.

Seno Reloncavf area, tidal amplitude 720 em Ancud area, tidal amplitude 226 cm
M22 M91 M90 M59 M37 M139 M82 M31 MIO M55 M56 M57
ANTHOZOA (10 species)
Cac/osoma chi/emis (Mc MUIIIIICH) 1~210
Phymactis clematis(DIlAYrON) 50-150 0-130 25-100 90-180
Anthopll'ura hermafroditica
CAIILGItEN 20-440
Bunodactis hermafroditica
(McMUltIIICH) 110-490 180-210 260-280 20-410 70-90 60-130 25-140 0-90
Parantheopsis cruentata
(DIlAYrON) 180-310 10-30
Paranthus niveus (LESSON) 250-310 20-45
Antholoba achates (CoUTONV) 210 25-150 60-130 25-30
Cereus? herpetodes (Mc MURRlCH) 170-200 105-22S
An/hothoe chi/emis (LESSON) 26-130 30-50
Aiptasiomorpha elongata CAIILGItEN 70-240 130-150
TURBELLARIA (4 species)
AprostalUm stiliferum BOCK 180-210
NotopltllUl australis SCHMAIIDA
f. huina MARCUS 350-540
N%plana chierchilli (PLEHN) 150-180
Notoplana palta MAllcus 90-265
NEMERTINI (9 species)
Cariniflll pacifica FIuEDIIICH 350-410
Baseodiscus aureus? (BiiIlGEJI) (90-265)
Poliopsis lacazei JOUBIN 350-410
Euborlasill nigrocincta COE 110-130
Parborlasill corrugatus McINTOSH 0-50
Nemertopsis gracilis COE 510-580 350-410
Amphiporus iwatai FltlEDIIICH 265-410
Dichonemenes coensis FIuEDIIICH 510-580 350-410
Atyponemertes chilemis FIIIEDIIICH 210-240 85-110

POLYCHAETA (1 species + ?)
Chaetopterus ?variopedatus
(RENIEIt) 0-130 85-110 30-70
Spirorbids 110-180 10-250 60-260 10-145 11)...210 195-245 20-70 0-110 30-70 60-90
SIPUNCULIDA (1 species)
Golfingill margaritacea (SAltS) 190-250 701 90-105 1
CIRRIPEDIA (6 species)
.... Chthamalus cirratus DARWIN 240-540 400-550 360-520 295-615 340-570 105-195 90-240 115-230 90-195 180-280
VI
.... Chthamalus scabrosus DAltWIN 210-480 160-310 210-260 255-295 70-410 (195-295)
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......
Vl Seno Reloncavi area, tidal amplitude 720 cm Ancud area, tidal amplitude 226 em
N
M22 M91 M90 M59 M37 M139 M82 M31 M10 M55 M56 M57
- - - - - - - - - - - - - - - - - - - - - - - - - -----

BalanuS {aevis BRUGUIERE 130-150 70-90

Balanus psittacus (MOLINA) 110-150 0-210 0-60
Balanus flosculus DARWIN 110-410 150-400 60-260 105-295 70-315 105-295 50-150 0-140 0-180
Elminius kingi GRAY 550--620 410--615 265-540 145-515 ?100-420 ?-375
DECAPODA (22 species)
Betaeus truncatus (DANA) 10
Alpheus chilensis COUTIERE 100-155
Pagurus edwardsi (DANA) 110-210 30-70 60-210
Pagurus villosus NICOLET 190-250 0-210 85-110 10-125 10-140
Petrolisthes laevigatus (GUERIN) 340-390 230-530 85-465 255-515 90-570 145-195 50-70
Petrolisthes angulosus (GUERIN) 80-210 10-20
Petrolisthes mitra (DANA) 45-70
Taliepus dentatus
(H. MILNE-EDWARDS) 20-90 x 10-50 100-150
Pisoides edwardsi BELL 10-30
Halicarcinus planatus (FABRICIUS) 110-210 70-250 250 10-220 10-265 195-245 10-30 x
Acantocyclus gayi (H. MILNE-
EDWARDS & LUCAS) 150-180
Acanthocyc/us albatrossis
RATHBUN 210-390 230-450 250-520 190-515 90-540 145-295 ?120-19O? 30-50
Cancer edwardsi BELL 80-310 110-130 10-105
Cancer plebejus POEPPIG 90-105
Cancer polyodon POEPPIG x
Gaudichaudia gaudichaudi
(H. MILNE-EDWARDS) 20 10-30
Homalaspis plana
(H. MILNE-EDWARDS) 60-210 10 40
Pinnotheres politus (SMITH) 190-250 60-160 10-15 10-160
Pinnixa bahamondei GARTH 0-50 85-110 30-70
Pinnaxodes chilensis
(R. MILNE-EDWARDS) 0-70
Hemigrapsus crenulatus
(H. MILNE-EDWARDS) 250-600 445-590 105-635 160-590 X2 x2
Cyclograpsus cinereus DANA 500-580 450-600 445-610 410--635 200--620
ISOPODA (9 species)
Jaeropsis bidens MENZIES 50-20
1socladus calcarea (DANA) 340-390 150-180 x 70-90 x 100-120
Exosphaeroma lanceolata (WHITE) 70-100
Dynamenella eatoni (MIERS) 110-130 30-90 0-90
Dynamenella tuberculata MENZIES 50-70
Dynamenella acuticauda MENZIES X
Cymodocella foveolata MENZIES X
Amphoroidea typa
H. MILNE-EDWARDS 20-20
Paradynamenopsis lundae MENZIES 490-590 400-550 x 340-410 x2 190
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Seno Reloncavi area, tidal amplitude 720 cm Ancud area, tidal amplitude 226 cm
M22 M91 M90 M59 M37 M139 M82 M31 MlO M55 M56 M57
STOMATOPODA (1 species)
Lysiosquilla chilensis DAHL 70-100

PYCNOGONIDA (1 species)
Tanystylum cavidorsum STOCK
var. steatopygidium HEDGPETH 50-30

POLYPLACOPHORA (8 species)
Plaxiphora aurata (SPALOWSKY) 40-240 80-250 120-160 100-170 145-295 60-80 10-50 0-30
Chaetopleura peruviana (LAMARCK) 20-110 30-50
Ischnochiton imitator (SMITH) 170-300 110
Ischnochiton pusio (SOWERBY) 180-250 70-145 90-410
Chiton granosus FREMBLY 210-240 160-360 90-410 110-220 80-140 50-140 90-180
Chiton latus SOWERBr 80-390 70-250 60-210 10-220 130-265 20-35 10-50 100-150
Tonicia atrata (SOWERBY) 80-300 0-210 85-160 10-145 10-180
Tonicia elegans (FREMBLy)3 110-130 50-70 60-85 20-70 0-100 20-70 60-150

PROSOBRANCHIA (24 species)

Fissurella costata (LESSON) 20-35 20-130 20-50 0-150
Fissurella maxima SOWERBY 20-10
Fissurella nigra (LESSON) (280-360) 190-220 130 145-245 30-50
Fissurella oriens (SOWERBY) 110-250? 0-130 10-90 145-195
Fissurella picta (GMELIN) 110-270
Nacella aenea (MARTYN) 10-15
Nacella magellanica (GMELIN) 110-390 70-360 60-360 70-410 90-410 195-295
Patel/oida ceciliana (ORBIGNY) 150-440 160-350 90-425 190-335 90-470 20-170 0-140 20-180 60-150
Patel/oida zebrina (LESSON) 180-190 100-180 90-180
Monodonta nigerrima (GMELIN) 270-590 200-600 270-640 190-480 210-580 100-140
Tegula atra (LESSON) 110-390 30-350 90-425 10-335 70-440 20-180 130-140 20-140 0-180
Prisogaster niger (WOOD) 80-190
Liltorina araucana ORBIGNY 190-590 220-600 260-680 210-630 400-570 190-220 190-350 100-195 180-195
Liltorina peruviana LAMARCK 430 425-465 (190-255)
Trochita trochiformis GMEUN 10-30
Crepidula plana SAY 50-70 50-70
Cripipatella dUatata (LAMARCK) 120-310 0-210 160-310 10-335 20-235 20-170 85-110 20-190 150-180
Argobuccinum argus (GMELIN) 180-110 0-50 20-10
Trophon xanthostoma BRODERIP 105-315
Concholepas concholepas
(BRUGUIERE) 0-60 90-180
Nucella calcar MARTYN 110-420 110-350 60-455 80-375 80-350 145-245 20-145
Euthria magellanica PHIL. 150-250 210-260
Nassarius dentiferus POWIS 90-105' 10-70'
Nassarius gay;; KIENER 80-250 0-210 10-145 10-160 195-245

......
Ul
W
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...
VI Seno RelclDcavf area, tidal amplitude 720 cm Aricud area, tidal 'ajnplitude 226 cm
-f'- M91 MI0 M55 'M56 M57
M22 ·M90 MS9 M37 M139 M82 M31
~~~
OPISTHOBRANCHIA (8 species) ;~~~- '.\:.'\.;;;~ ~~II'

Onchidella marginata (GOULD) ~'f!!J ,.. 190-220 l:ID-lSO 100-195

Berlhella plalei (BERGH) 140
Rrulanga pulchra MAC FARLAND 30-120
Neodoris erinacea MARCUS 0
Neodoris caTVi MARCUS 20-10
Anisodoris lesselala BERGH SO 12G-155
Phidiana inca ORBIGNY ~30-150
Aeolidia papillosa
var. serolina BERGH 200 SO

PULMONATA (2 species + ?)
Phyliasp. 225-310
Siphonarians 390-540 15G-400 360-465 33S-515 19S-295 5G-220 90-210 3G-140 15G-ISO
Chi/ina bulloides SOWERBY ?1SG-520 ?22G-510
BIVALVIA (7 species)
Mytilus edulis chi/emis Hupt 33G-540 1SG-520 6G-520 14S-335 7G-41O 14S-395 1G-190 130-150 100-120
Aulacomya aler MOLINA 11G-240 G-ISO 30-280 10-145 7G-410 14S-295 2G-150
Hormomya granulata HANLEY 8S-46S IG-70
Brachidonles purpuratus LAMARCK 21G-540 4OG-520 290-540 29s-410 18G-540 14S-245 7G-ISO 8G-140 100-190 18G-245
Oslrea chi/emis (PHILIPPI) 130-300 5G-220 6G-360 90-220 7G-240 4G-110
Lasaea petiliana RECLUZ 24G-540 3SG-55O 36O-46S 33s-45O ~70 17G-190 100-195 18G-245
Rocellaria denticulala DESHAYES 1G-50
ASTEROIDEA (4 species)
Cycelhra verrucosa (PHILlPPPI) 18G-210
Patiriella fimbriala (PERRlEII) G-110
Anaslerias varium (PHILIPPI) 8G-31O G-230 10-125 1-205
Crumaslerias lurido (PHILIPPI) G-5O 1-205
OPHIUROIDEA (5 species)
Ophiomilrella chi/emis MORTENSEN 140
Ophiactis asperula (PHILIPPI) ISO 90-110 1~
Amphiura magellanica LJUNGMAN 90-110
Amphiodia chilemis
(MOLLEII & TIIOSCHEL) SG-90
Amphipholis squamata
(DELLE CHWE) 190-250 90-265
ECHINOIDEA (3 species)
Arbacia dufresnii (BLAINVILLE) 8G-15O G-210 6G-85 10-190 9S-195
Pseudechinus mage/lanicus (PHILIPPI) G-70 1s-45
Loxechinus albus (MOLINA) 15G-21O G-150 10-105 :ID-220 ?9S-145 20-10
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Seno Reloncavf area, tidal amplitude 720 em Ancud area, tidal amplitude 226 em
M22 M91 M90 M59 M37 M139 M82 M31 MI0 M55 MS6 M57
HOLOTHURIOIDEA (3 species)
Cucumaria godeffroyi SEMPER 1»-310 0-90
Pseudocnus dubiosus leoninus
(SEMPER) 0-70
Chiridota pisanii LUDWIG 70
ASCIDIACEA (7 species)
Amauroucium [uegiense
(CUNNINGHAM) 180-190 25-90 10-30
Didemnum chilense ARNBACK nO-130 180-190 10-30
Lissoclinum caulleryi
(RmER & FoISYI'H) UO-130 20-35 0
Distaplia orientalis BANCROFT nO-150 20-70
Corella eumyota TIAUSTEDT UO-200 90-130 65-110 25-70
Styela paessleri MICHAELSEN 20-35 10-30
Pyura chi/ensis MOLINA 80-2607 10-210 260-280 1tJ.-145 1tJ.-23O 35-110 0-60 50-70 150-180
CHLOROPHYTA (14 species)
Enteromorpha bulbosa
(SUHR.) KiiTz. 590-610 310-690 270-570 410-640 415-680 195-465 1-380 50-150 1tJ.-175 120-240 155-255
Enteromorpha gunniana J .AG. 1tJ.-175
Blidingia minima (NAEG.) KYLIN 295-570
Percursaria percursa
(C.AG.) ROSENY. (145-195)
Viva lactuca L. 1»-440 70-3SO 140-465 100-540 125-415 1-100 20-150 40-190 2tJ.-14O 2tJ.-245
Viva nemaloidea (BolY) 125-130 2tJ.-170
Prasiola slipilala SUHR. 1100-380 225-275
Cladophoropsis brachyarlhra
(SYED.) BORG. 1100-380
Chaelomorpha aerea (DILLW.) KiiTz. 190-210 280-360 195-245
Rhizoclonium lorlUOsum
DILLW. KiiTz. 200 570-630 40-210
Rhizoclonium riparium
(Rom.) HARY. 550-6S0 280-360 545-640 145-195 160-300
Cladophora falklandica
HOOK. f. & HAllY. x
Cladophora subsimplex Kfrrz. 480-515 190-275
Cladophora incompla
HOOK. f. & HAIY. SO

PHAEOPHYTA (17 species)

Pylaiella liloralis (L.) KJELLM. 195-245
EClocarpus confervoides
(ROTH.) KJELLM. 105-130
.... Feldmannia chilinicola
VI (SAUND.) LEYI. 7G-150 20
VI
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......
VI Se.no Reloneav! area, tidal amplitude 720 em Aneud area, tidal amplitude 226 em
a..
M22 M91 M90 M59 M37 M139 M82 M31 M10 M55 M56 M57
Sphaeelaria furcigera KiiTZ. 70-150 145-325 35-70
Glossophora kunthii (C.AG.) J.AG. 20-20
Myrionema strangulans GREV. 10-170
Ralfsia australis SKOITSB. 50-70 60-170
Ralfsia californiea SETCH. & GARDN. 60-80
Leathesis difformis (L.) ARESCH. 130 10-90 60-110
Myriogloia ehilensis LEVR. 90
Desmarestia herbaeea (L.) LAMOUR. ?20-20
Adenoeyestis utrieularis
(BoRv) SKOITSB. 125-?
Seytosiphon lomentaria
(LVNGB.) ENDL. 310-350
Colpomenia sinuosa
(ROTH.) DERB. & SOL. 60-110
Seytohamnus fascieulatus
(HOOK. f. & HARV.) CorrON 360-425 295-410 360
Lessonia nigreseens BoRY ?50-30
Macroeystis pyrifera (L.) C.AG. ?0-40 ?0-40 75-215 ?-35
CYANOPHYTA (4 species)
Calothrix scopulorum
(WEBER & MOHR) C.AG. 380-630
Anabaena variabilis KiiTz 170-200
Lyngbya aestuarii (MERT.) LIEBM. 550-650 ?570-61O? 265-435
Mieroeoleus ehtonoplastes
(MERT.) THUR 615-630

RHODOPHYTA (42 species)

Goniotriehum elegans
(CHAUV.) LE JOLIS 195-295
Erythrocladia irregularis ROSENV. 40-210
Erythrocladia subintegra ROSENV. 40-210
Erythrotriehia earnea (DILLW.) J.AG. 200 70-360
Erythrotriehia polymorpha HOWE 35-70
Porphyra eolumbina MONT. 220-440 250-620 300-400? 375-545 380-440 245-415 295-570 200-265 185-275 75-200 200-280
Acroehaetium eatenulatum HOWE 200 40-210 195-245
Aeroehaetium leptonemoides LEVR. 200
Aerochaetium porphyrae
(DREW) SMITH 390-440
Chaetangium faitigiatum
(BORY) J.AG. 190-350 130-280 170-375 100-400
Gelidium pusillum
(STACKH.) LE JOLIS 35-75 10-190
Gelidium pseudointrieatum
SKOTTSB. & LEVR. 380-540 400-600 230-520 175-545 260-470 145-305 80-200 60-280
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Seno Reloneavi area, tidal amplitude 720 em Aneud area, tidal amplitude 226 em
M22 M91 M90 M59 M37 M139 M82 M31 M10 M55 M56 M57
Gelidium lingulatum J.AG. 60-110 2~120
Hi/denbrandtia lecannellieri HARlOT 20-110 105-160
Litothamnion pauciporosum LEM. 210-240 180-210 390 1~145 20-140 20-100 2~1O 0-150
Lithothamnion caroli LEM. 70-90 390 (}...6()
Litothamnion validum FosL. 2~1O ?0-30?
Crodelia accedens (FosL.) LEVR. 130-210 x 1~15 20-50 35-120 0-160 ?2~100 0-120
Corallina officina/is (L.) LAMOUR. 70-140 2~70
Corallina chi/ensis DCNE. 2~1O 0-120
Pugetia chi/ensis (J. AG.) K YLiN 50-140
Trematocarpus dichotomus KiiTZ 3~9O 4IJ...50 0-100
Agardhiella tenera (J.AG.) SCHMITZ. 0
Catenellafusiformis (J.AG.) SKOTTSB. 490-540 410-480 410-590
Chondrus canaliculatus
(C.AG.) GREV. 270-300 200-250 160-390 145-410 155-180
Gigartina chamissoi (C.AG.) J.AG. 190-210 360-425
Iridaea boryana
(SETCH. & GARON.) SKOTTSB. 110-190 10-180
Iridaea membranacea J. AG. 40-70
Iridaea dichotoma
HOOK. f. & HARv. 90-415
Iridaea undulosa BORY 120-130
Iridaea ciliata KiiTZ. 180-250 60-425 105-335 40-180 3~9O ?3(}...6()
Iridaea crispata BoRY 150 295-335 350-450
Dendrymenia skottsbergi
(DAWS.) LEVR. 0 2~30?
Ceramium rubrum (Huos.) C.AG. 15-60
Ceramium stichidiosum J.AG. 230 ?-100 20-90
Ceramium dozei HARlOT 0-150
Centroceras clavulatum
(C.AG.) MONT. 80-140 1~90
Heterosiphonia punicea
(MONT.) KYLIN 2(}...6() 50-110
Polysiphonia abscissa
HOOK. f. & HARv. 80-90 70-150 145-245 5-150
Bostrychia hookeri HARv. 490-540 550-600 360-465 375-630 410-520 x6 60-100 ?-200 195-265
Bostrychia harveyi MONT. 595-635 520-590 1160-630 315-570
Herposiphonia su/ivanae
(HOOK. f. & HARV.) FALKENB. 2~50 50-150?

1. Var. hanseni STEPHEN.

2. Among Elminius, level not noted.
3. All varieties.
4. One shell only, with Pagurus sp.
5. Shells with Pagurus villosus also between 90 and 140.
6. Upmost in the Enteromorpha zone .
......
Vl
-...j
 Appendix 2A. Species only found in rock pools allhe Ancud 51alions. Appendix 2B. Species found bolh in rock pools and alone or more 51alions
in Ancud or jusl oUlside Ihese.
Polycysti.r yagana MAIcus MS6
Porrocysti.r aasimilis (LEVINSEN) MlO AlIIhopleurtl. hermafrodilica MSS,MS6
Vortictros dahl; MARCUS MS6 ParlUllMopsis C""'_tII MSS
Promonotus pardus MAIcus MI0 AlIIhotJwe chik",is MSS
Mia.a t.e/itaat MARCUS MI0, MSS, MS6 AiptllSwmorpha elongata MSS
Proceroda trIIlCroslOmo (DARWIN) MS6 Pe"o/is/hts laevigatw MSS,M56
NOlOplana puma MAIcus MS6 Ha/icarcinw planatw M10,M55
Be"'w emargillatw H. MILNE-EDWARDS MSS ACJJnlhocyclus albatrossis M10
Neomysis sopayi HOLMQUIST MI0, MSS, MS6 Cancer pa/yodon MlO
Neomysis iIytlpfli HOLMQUIST M10, MSS, MS6 HomalllSpis plana M10, M55
PateUoitJIJ orbigllyi (DALL) MS6 Hemigrapsus crenu/atus MlO
Anisodoris punctuolatll (ORBIGNV) Exosphaeroma lanceola/ll M10
var. cymilla MARCUS MS6 AmphoroUka typa MS7
'f1tecDctra dJuwini PaUVor-FoL MS7 Chaetopleura peru.iana MS5
Meye_"'r gelatinosw (MEYEN) MS6. MS7 Tonicia elegam f. grayi MlO
DamartSliIJ llgultml. (LIGHTF.) LuJoUR. MIO Concholep.. concholepas MS6
CalothrU: confervicola (RotH.) C.AG. MS6 PhytitJsp. MS6
Lyngbya confervoidD C.AG. MIO, MSS, MS6, MS7 Hormomya granu/a/Il MS5
O.trea chiIe",is MS6
Amphipholis .q/lllllJa/a M10,MS5
Corella eumyotil MS5
Chaetomorpha aerea M10,M5S
Macrocy.ti.r pyrifera M10
CoraUina offit:iMlis MIO,MS7
ChondruS CtUJaliculalru MS6
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Giganina chonWsoi MI0

Ceramium rubrum MSS
QlIlTOCtrao cla.u/aIwn MlO

Appendix 2C. PoIychacta and Bivalvia laken in sand and Ibe slalions and levels (in em) al which Ihcy were found. Deplhs in italics arc below zero.
Species M22 M91 M90 MS9 M37 MIO MSS MS6

POLYCHAETA
H"'ydna brevisetosa KtNBERG 156-180
H"'ydna patllgonico KlNBERG llG-l30
EuhIlia strlga/ll EHLERS G-SO
Dalhousiella arocuda WESENBERG-LuND 19G-2SQ
SylHs brachychoettl ScHM.u.DA 180-210
SyIHsIlllOp$ EHLERS 180-210
PeriMreis vallatII (GaUBE) 43(}-S10 46S-S20 105-S4S 9G-S4Q + + +
Perinertis falJcJluulit:o RAMsAv 21G-230 + + +
PmuIonereis galhlpagemis KIN.EBG 21G-230 + +
Platynereis maga/haeluis KlNBEBG 130-310 llG-l30 105-190 9G-340 + + +
GlyartlltmgissiIruJ ABWlDSSON 256-310
Gylunl papiJlo6a GaUBE 9G-26S +
Hemipodus .implu (GaURE) 256-390 9G-26S +
Marphy.a _ (BLANCHARD) 38S + +
BIVALVIA
DiplodonIIJ ineonspicIIIJ I'IIIUPft 19G-310 7G-9O S6-26S
Ameghinomya IlllliqIUl KING 156-180 S6-70 10-90 105-140
Clausintlla gayi Huot 105-125
Gari .olitJIJ GRAV S6-70 105-140
Tagelus dmnbeii L.ul.u.a 9G-110 S6-70
Semele corruga/ll SowI!uY OUlside IG-30 105-140
Ensis macho MOUNA OUlside
Hialella .a/itJIJ SoWERBV G-SO 310 7G-14S

Since sand-Hving species were nul scarched for Ihc levels don't show Ibeir toIal vertical distribulion, only al which levels Ihey were occasionally found.

158
 Appendix 3
BIOLOGY AND COLOURS OF SOME ANIMALS
Our field notes contain a lot of information on the biology of the animals
studied, on the months during whicb eggs and young stages were observed,
and on the colours of the living animals. Some of this information has been
published in the LUCE reports on the diverse taxa, but as this information
is disseminated in many publications it is not easily accessible. The many
unpublished notes, hidden in the field diaries, are still less easy to trace.
Since some of this information may be of interest for certaim purposes, it
will be recorded here. Some of the information refers to species found at the
stations but outside the sections studied.
ANTHOZOA
Phymactis clematis. Green, brown·green or brown.
Anthopleura hermafroditica. Mostly on the underside of stones at the border
to the sand. - Greenish brown, tentacles blue-green.
Bunodactis hermafroditica. On stones in sand. One specimen observed eating
a nereid polychaete. - Greyish brown or greyish, tentacles blue-green.
Parantheopsis cruentatD. Occurrence as Authopleu'a. - Grey-green.
Paranthus niveus. Pink.
Antholoba achates. Brown.
Cereus? htrpetodes. Encrusted with shell fragments and sand grains. Trans-
verse fission observed in January and February.
Anthothoe chilensis. Grey with brown lines and yellow-white tentacles.
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Aiptasiomorpha e/ongala. Salmon-rcd, white lines.
TuRBELLARIA
Aprostarum stiliferum. Grey-brown.
Notoplana australis f. huina. Grey.
Notoplana chierchiai. Grey-brown.
Notoplana palta. Grey.
NEMERTIN[
Euborlasia nigrocincta. Chocolate-brown, square-striped.
Euborlasia corrugatus. Brown or dark black-brown.
Nemertopsis gracilis. Dark grey-green with light dorsal stripe.
Atyponemertes chi/ensu. Grey-green.
POLYCHAETA
Eulalia strigata. Dark black-green with light-green parapodia.
Perinereis val/ata. Spawning 31 January.
Platynereis magalhaensis. Tough tubes between Macrocystis pyrifera holdfasts.
Chaetopterw ?variopedatus. Under stones. Pinnixa bahamondei in the tubes.
CiRR[PEDlA
Chthama/w cirratw. On stones and molluscs but missing at the base of the
stones. - Grey but often green from small algae.
Chthamalus scabrosus. On·stones and molluscs, often covered by green algae.
Balanus laevis. Only found on Balanus psittacus.
Balanus psittacus. Often with Balanus laevis and B. flosculus on the shells.
Balanus flosculus. Very common on big boulders and molluscs. Sometimes
tube-shaped, especially on seaward side of boulders.
Elminius kingi. On stones and Mytilus edulis chilensis. Also found on rocks
in almost fresh water at Stn M31 and brackish water at M82. No other
barnacles found at these two stations. - White, but growth-stages of
Enteromorpha sp. and diatoms give the shells a brownish colour.
DECAPODA
Alpheus chilensis. Grey-brawn-green with reddish legs.
Pagurus edwardsi. Most common in shells of Tegula#atra but also in shells
of Nucella calcar. - Red with red claws.
Pagurus villosus. Very common on sand, especially among Pyura chilensis,
in shells of Nassarius gayii, sometimes also in shells of Nassariius denti-
ferus.
Petrolisthes laevigatus. Very lively. Hides under stones. keeping tight to
these. - Violet-blue.
Petrolisthes angulosus. Grey-mottled or red.
Taliepus dentatus. Brown.
Pisoides edwardsi. Red, grey or grey-brown. Claws red.
Halicarcinus planatus. Sluggish. - Greyish or brown with spots of varying
colours.
Acanthocyclus albatrossis. Under stones. Very sluggish. Eggs seen in April
and May, young specimens in January and February. - Blueish-black or
grey-brawn. Leg ends white with brown tips. Tip of left claw white with
violet underside. Tip of right claw white, at the base and along the inner
side violet. Eggs vermilion-red.
Cancer edwardsi. Young specimens observed in January and February. -
Violet, dark-violet, red-violet or dark blood-red.
Cancer plebtjus. Young specimens taken in January. - Violet, light violet,
wine-red or flesh-coloured with light spots.
Cancer polyodon. Red with black claws.
GaudichaudUJ gaudichaudi. Red or violet with black claws.
Homalaspis plana. Red, red-violet, violet or blue-red. Claws black.
Pinnotheres politus. Commensalistic under Crepipatella diiataJa. Big speci-
mens often found in small Crepipatella. In one sample from low down
on the shore 10 of 25 Crepipatella had Pinnotheres. Females were domi-
nant. Berried females observed January to April. On 31 January 12 of
16 females were berried at Stn M37. - White to grey-brown.
Pinnixa bahamondei. Common in tubes of Chtutoplerus ?variopedatus in
Seno Reloncavi.
Pin1lOJtodes chilensis. [n the intestine of Loxechinus albus.
Htmigrapsus crenulatus. Fairly lively. Under stones and common among EI-
minius kingi. One large specimen with undetermined alga on the legs. -
Mostly grey but also black··grey, bl~ck, black-brown, brown, brown-green
or blue-grey with some lighter pans. Tubercles on carapax black-grey.
Underside and claws white.
Cyc/ograpsus cinereus. Fairly lively. High up on the shore, even among ter-
restrial plants. Hides under stones. - Brown.
ISOPODA
Jaeropsis bidens. Among holdfasts of Lessonia nigrescens and Macrocystis
pyrifera.
[socladus calcarea. Curling up like Sphaeroma. - Brownish or often yellow.
Exosphaeroma lanceolata. Curling up like Sphaeroma. - Brown.
Dynamenella eatoni. Curling up like Sphaeroma. At one station found among
holdfasts of Lessonia nigrescens.
Amphoroidea typa. Eggs observed in February. - Grey, grey-green, green,
brown or red-brown with rust-brown segment borders. Eggs yellow.
Paradynamenopsis lundae. Curling up like Sphaeroma.
PvCNOGONIDA
Tanystylum cavidorsum. Among holdfasts of Marcrocystis pyrifera.
POLYPLACOPHORA
Plaxiphora aurala. Heavy cover of aJgae on the plates where also Balanus
floscuJus may be found.
Chaetopleura peruviana. White.
lschnociton imitator. At M56 on holdfasts of MacTocystis pyrifera. - White.
Ischnochiton pusio. Mostly under stones where it is moist when the shore is
emerged. - Black.
Chiton granosus. Black or grey-green.
Chiton latus. Under stones. - Forma subfuscus has plates with different
shades of red-brown. Margin black.
Tonicia atrata. Under stones. - Margin black. Some specimens became yel-
low-grey, others orange in alcohol.
Tonicia e/egans. Moves around quite a lot when the shore is submerged. -
Plates greenish or black, margin light-coloured, much mottled, yellow,
orange or black. Forma chilensis, margin black.
PROSOBRANCH[A
Fissurella costata. Calcareous algae and spirorbids on the shell. - Mottled.
Fis~'urella maxima. Red-striped.
Fissurella nigra. Sometimes with Balanus flosculus on the shell. - Black,
black-blue or black-violet, white around top hole.
Fissurella oriens. Light or black-violet, radially striped.
Fissurella picta. Radially striped.
Nacella aenea. Mottled or reddish with dark spots or light with red margin.
Nacella magellanica. Sexually mature in April and May. - Mottled, inside
of shell uncoloured, foot grey.
Patel/oida ced/;ana. Some specimens very high. - Dark, radially striped with
angle-shaped dots. dark ring around top. Inside blue or blue-green.
Patelloida orbignyi. Black with dark margin.
Patelloida zebrina. Mottled with angle-shaped dots.
Monodonta nigerrima. Grey-blue. blue or black.
Tegula atra. Strolls widely when the water floods. seeks back to base of
stones when the shore is emerged (Fig. 3). Specimens often overgrown
with calcareous algae. Naked scars where specimens of Patel/aida cecil·
iana have been fixed. One big specimen with Balanus psittacus on the
shell. Some shells with Pagurus edwardsi. On an overhanging boulder 5-8
specimens hung together. forming a chain. Some specimens wilh egg cap·
159
 sules of Crepipatella dilatata on the shell in March. - Dark-blue, blue
violet, black or black-violet.
Prisogaster niger. Violet or black-violet, operculum brown.
Littorina aroucana. Small specimens often among Chthamalus cirratus and
Elminius kingi, together with Lasaea petitiana. Large specimens on the
rocks. mostly in depressions and crevices. - Grey.
Crepipate[{a dilatata. Mostly on stones, in some places covering them, especi-
ally the sides and free undersides, but also on Balanus psittac... and Tegu-
la atra, young specimens also on Fissurella costata. Shells with Balanus
have impressions from the riffles of the barnacle. Many specimens with
Pinnotheres polit... below. Chain-fonning (Fig. 11), specimens decreasing
in size towards the top. The two lowest specimens in a chain had eggs
in January and February. Egg capsules observed January to May. -
White or pink. eggs yellow-grey.
Argobuccinum arg.... Brownish. striped.
Trophon xanthostoma. Light violet.
Concholepas concholepas juv. Grey-green.
Nucella calcar. Some shells with Pagur... edwardsi. Egg capsules seen in
December. April. May. and July. - Colour very varied. dark grey-violet,
yellow or flame-coloured, small specimens white. Blue or white around
the aperture.
Euthria magellanica. Violet-blue. blue. blue-green or green.
Nassari ... dentiferus. Shells sometimes with Pagurus vi/losus. - Banded.
Nassarius gayii. On stones and sand. Hides in holes in the sand when the
shore is submerged. Many shells with Pagurus villos.... - Brown or violet-
brown. with brown spiral.. bands. also violet-brown or dark violet. striped.
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Inside of shell white.
OPISTHOBRANCHIA
Onchidella marginata. Often on Chthamalus cirralUS. Eggs observed in early
March. - Black or black-green.
Berthella platei. White.
Rostanga pu/Chra. Bright orange-red.
Neodoris erinacea. Yellow or red-brown. underside orange.
Anisodoris punctuolata var. cymina. Slimy, dissolves fast after capture. -
White-yellow or orange-yellow.
Anisodoris tesselata. Yellow or yellow-grey. dorsal side with fine black net-
pattern.
Thecacera darwini. Yellow with black dots.
Phidiana inca. Papillae rust-brown with white tips.
Aeolidia papil/osa var. serotina. Grey-brown or red-brown. foot salmon-red,
papillae blue-violet or grey-brown.
PULMOl'olATA
Phytia sp. Among Littorina araucana.
Siphonarians. possibly more than one species. Mainly in holes. - Brown-
green with radial stripes from top to margin. Inside dark violet with
white margin. Foot yellow-white with grey-blue sides with white spots.
Chi/ina bulloides. In brackish and almost fresh water. Eggs observed in Janua-
ry. - Shell with dark spots.
BIVALVIA
My til... edulis chilensis. Often together with Lasaea petitiana and with Elmini-
... kingi and a variety of algae on the shell. Newly settled specimens in
April. - Blue, blue-brown or brown.
160
Aulacomya ater. Common on lower margin of stones. With spirorbids.
Chthamalus <iTTatus, and bryozoans on the shell. Newly settled specimens
in April. - Young specimens yellow.
Hormomya granulata. Yellow. striped.
Brachidontes purpuratus. On stones but also burried in sand at the base of
the stones. Often together with Chthamal... ciTTatus. Observed as high
up as +540 cm. in Seno Reloncavi, with different algae on the shells. -
Blue, violet or black- violet.
OS/Tea chilensis. Locally common on exposed side of boulders. together with
Crepipate/la dilatata.
Lasaea petitiana. Common among Chthamalus cirralUS, Balanus flosculus, and
mussels. - Pink.
Tagelus dombeii. Digging at least 50 em deep in the sand.
Diplodonta inconspicua, Ameghinomya antiqua, Clausinella gay;, Gari solida,
and Hiatella solida. Sand-living. - White.
ASTEROIDEA
Cycethra verrucosa. Orange-yellow, vermillon or orange-red.
Patiriella fimbriata. Red, eosin-red or dark-red.
Anasterias varium. Green, moss-green or black-green. Fast being red in alco-
hol.
Cosmasterias lurida. Colour varying, most often violet but also red-brown
or grey-brawn-green.
OPHIUROIDEA
Ophiomitrella chilensis. Yellow·orange or yellow-red.
Ophiactis asperula. Mottled. like Ophiopholis aculeata (L).
Amphiura magellanica. Grey.
ECHINOIDEA
Loxechinus albus. Some specimens with Pinnaxodes chilensis in the intestine.
- Green.
HOLOTHURIOIDEA
Cucumaria godeffroyi. Yellow.
Pseudocnus dubiosus leoninus. Greyish or yellow-red.
ASCIDIACEA
Amaroucium fuegierue. Under stones and on Pyura chi/ensis, together with
Didemnum chi/ense. - Grey, red around siphons.
Didemnum chi/erue. Under stones and on Pyura chilensis and Hormomya
granulata.
Lissoclinum caulleryi. On Pyura chilensis. - Yellow-brown.
Distaplia occidentalis. Red, wine-red or red-brown.
Corella eumyota. Some with copepods in branchial sac. - White. milk-white.
vitreous, branchial sac pink or red.
Pyura chilensis. Very common under and between stones to which specimens
are fixed. Cements together stones and shells. With Amaroucium fuegien-
se, Didemnum chilense, and Lissoclinum caulleryi on mantle. - Red or
pink but can be green from algae.
PISCES
(Sicyaces? sp.) 'Peie sapo'. On boulders. one specimen covering fry in Janua-
ry. at +385 cm, in Seno Reloncavi. about MSL (Fig. 12).