5 ISSN 0016-531X

" DES ANA

ZOOLOGIA , 1983 N* 46

ha yO

SOUTHERN PACIFIC

BRYOZOA: A GENERAL

VIEW WITH EMPHASIS

ON CHILEAN SPECIES

UENSENACRE SIDAD DE CONCEPCION

ES E

MIREN ALBERDI

Universidad Austral de Chile

SERGIO ÁVARIA

Universidad de Valparaíso

Danko Brncic

Universidad de Chile, Santiago

Ebuarbo Busros

Universidad de Chile, Santiago.

Huco Campos

Universidad Austral de Chile

ACULTAD DE CIENCIAS BIOLOGICAS

Y DE RECURSOS NATURALES

UNIVERSIDAD DE CONCEPCION

C'H TL'E

DIRECTOR:

Parricio Rivera R.

REEMPLAZANTE DEL DIRECTOR:

Oscar Mar rmHen J.

REPRESENTANTE LEGAL:

GUILLERMO CLERICUS E.

PROPIETARIO:

Universidad de Concepción

DOMICILIO LEGAL:

Barros Arana 631, 3% piso, Concepción, Chile.

COMITE DE PUBLICACION

KkrisLER Alvear V. CLODOMIRO MARTICORENA P.

Oscar Marrnenr J.

CLEMENTE GONZÁLEZ S.

JorGE ARTIGAS C.

COMITE ASESOR TECNICO

Jurke Gral

Universidad de Múnchen, Alemania Federal

Mary KaLin ARROYO

Universidad de Chile, Santiago.

Luis RAMORINO

Universidad de Valparaíso

BERNABÉ SAN TELICES

Universidad Católica, Santiago

FEDERICO SCHLEGEL

Universidad Austral de Chile

Juan CarLOS CASTILLA

Universidad Católica, Santiago

FERNANDO CERVIGÓN

Fundación Científica Los Roques,

Venezuela

MónNIkaA DURRSCHMIDA

Jusrus-LiEBIG

Universitát Giessen, Alemania Federal

RaúL FERNÁNDEZ

Universidad de Chile, Santiago

JosÉ Sruarbo

Universidad de Concepción

Tob SrueEssy

Ohio State University, U.S.A.

GUILLERMO TELL

Universidad de Buenos Aires, Argentina

HaroLbo Toro

Universidad Católica, Valparaíso

ISSN 0016-531X

GAYANA

ZOOLOGIA 1983

SOUTHERN PACIFIC BRYOZOA:

A GENERAL VIEW WITH EMPHASIS

ON CHILEAN SPECIES

Hugo I. Moyano

UNIVERSIDAD DE CONCEPCION

CELLDOE

> 46

“Los infinitos seres naturales no podrán perfectamente conocerse

sino luego que los sabios del país hagan un especial estudio de

ellos”.

CLAUDIO GAY, Hast. de Chale, 1:14 (1848)

Impreso en

Ebrrortal UNIVERSITARIA

San Francisco 454-Casilla 10220

Santiago-Chile

SOUTHERN PACIFIC BRYOZOA: A GENERAL VIEW

WITH EMPHASIS ON CHILEAN SPECIES*

Hugo 1. Moyano!

RESUMEN.

Este es un estudio sobre la taxonomía, zoogeografía, poli-

morfismo y diversidad zoarial de las Faunas Pacíficas de

Bryozoa con énfasis en las formas australes orientales. El

número total de Bryozoa marinos chilenos alcanza a 267

especies, que se distribuyen en 19 del orden Ctenostoma-

ta, 46 del orden Cyclostomata y 202 del orden Cheilosto-

mata. Entre éstas se hallan las 24 especies y un nombre

nuevos que se describen aquí. Zoogeográficamente la fau-

na briozoológica chilena puede distribuirse en las cuatro

provincias siguientes: Magallánica (42% — 56"30'5), Chi-

lena (18%S — 42%S), Juan Fernández (33%38'5; 7552)

Isla de Pascua (27"07'S; 109%22'W).

Taxonómicamente, en el Pacífico suroriental se en-

cuentran proporcionalmente poco representadas las fa-

milias Reteporidae, Vittaticellidae, Adeonidae y Serupo-

cellariidae, y completamente ausentes Eurystomellidae,

Euthyrisellidae, Conescharellinidae, Euthyroididae, y

Parmulariidae.

En todo el océano Pacífico más la Antártica y la cuenca

del Artico, el polimorfismo y la diversidad zoarial de los

Cheilostomata alcanzan los mayores valores en el lado

occidental desde la Antártica a las Filipinas. Y los valores

más bajos aparecen en el lado oriental desde la región

magallánica al Artico. Lo mismo aparece cuando se com-

paran estos parámetros calculados para las faunas tercia-

rias de Bryozoa de Argentina, Nueva Zelandia y Sudaus-

tralia.

Sobre la base de que la alta diversidad zoartal de toda la

briozoofauna y el alto polimorfismo de los Cheilostomata

indican especies de selección K, los valores proporcional-

mente mayores de éstos parámetros de las faunas de Bryo-

zoa de las plataformas continentales del Pacífico occiden-

tal desde la Antártica a las Filipinas, estarían reflejando

una historia de ambientes más predecibles. El Pacífico

oriental, por el contrario, con valores más bajos reflejaría,

al menos para la costa chilena sudamericana, una gran

variabilidad de las condiciones ambientales en el largo

plazo, las que realmente se hicieron presente durante una

gran parte del Terciario.

"Departamento de Zoología, Universidad de Concep-

ción Casilla 2407, Concepción, Chile.

*Contribution presented to the “Second International

ABSTRACT

This is a study on the taxonomy, zoogeography,

polymorphism and zoarial diversity of the Pacific

Bryozoan Faunas with emphasis on the austral eastern

forms. The total number of the Chilean marine Bryozoa

reaches to 267 comprehending 19

Ctenostomata, 46 Cyclostomata and 202 Cheilostomata.

These include twenty four new species and a new name

being described here. Zoogeographically the Chilean

Bryozoan Fauna can be devided into four provinces,

namely: Magellanic (42"S-56"30'S); Chilean (18%5-429S);

Juan Fernández (33"38'S; 75%52'W) and Easter Island

(27%07'S; 109%22'W).

Taxonomically, in the Eastern South Pacific the

families Reteporidae, Vittaticellidae, Adeonidae and

Scrupocellariidae are proportionally poorly represented,

whereas the Eurystomellidae, Euthyrisellidae,

Conescharellinidae, Euthyroididae and Parmulariidae

are altogether absent.

In the whole Pacific Ocean plus the Antarctic and the

Artic basin, the Cheilostomata polymorphism and the

zoarial diversity attain the highest values in the western

side from the Antarctc to the Philippines and the lowest

ones in the eastern side from the Magellanic region to the

Artic, the same reveals when comparing these parameters

«calculated for the Tertiary Bryozoan Faunas from

Argentine, New Zealand and South Australia.

On the basis that high zoarial diversity and high

Cheilostomata polymorphism values indicate more

K-selected bryozoan species, then the proportionally

higher values of these parameters in the Western Pacific

shelves bryozoan faunas from the Antarctic to the

Philippines, would reflect a history of more predictable

environments. The Eastern Pacific, on the contrary, with

lower values would reflect at least for the South-american

Chilean coast, a great variability of the environmental

conditions, which were really present during a large part

of the Tertiary.

now species

Key Words: Bryozoa, Taxonomy, Zoogeography, Zoarial

Diversity, Pacific Basin.

Symposium on Marine Biogeography and Evolution in

the Pacific”. Sydney, Australia, 5-17 July, 1982.

INTRODUCTION

Obtaining general views in a global scale, both

in time and space, has been the aim of several

contributions during the last decades in the

evolutive, paleontological, zoogeographical,

botanical and ecological fields (Durham,

1979; Fleming, 1979; GrantMackie, 1979;

Heck and McCoi, 1979, Knox, 1979 and

Kubanin, 1980). Whithin the bryozoan field,

Schopf (1973) considers the bryozoan colony

as the unit in which the natural selection

operates producing the polymorphs such as

the avicularia and vibracularia.

applied this view to compare polar and

tropical faunas, showing that the

polymorphism of the Cheilostomata

significatively varies between them. In his

discussion, he puts together the evolutive

environmental forces governing the evolution

of bryozoan species and the possibility to

characterize bryozoan assemblages from

diferent areas of the world. Extending this

view, Moyano (1975) compares Antarctic,

Arctic and Indo-Pacific bryozoan faunas and

characterizes them according to their global

polymorphism values. More recently, Moyano

(1978, 1982) further extendes this perspective

to the bryozoan zoarial types, measuring

zoarial diversity of polar, temperate and

tropical bryozoan faunas. In addition, the

combined analysis of the polymorphism plus

the zoarial diversity of both recent and fossil

bryozoan faunas would reflect present and

past environmental conditions modeling

them. This latter approach was pioneerly

developed and applied to fossil bryozoan

faunas of the Australia region (Stach, 1936,

1938).

Global fossil (Ross, 1978) or recent (Schopt,

Fisher 8 Smith, 1978; Schopf, 1979)

zoogeographical schemes, using the Bryozoa

as the unique or principal group have been

recently developed. A more local scheme has

defined the Magellanic Bryozoan Region

(Moyano, 1982). Schopf et al. predicted 32

faunal provinces of the world's continental

shelves using Bryozoa, Pelecypoda,

Foraminiferida and Coral faunas. In this

scheme, the South Pacific —from the Equator

to the Ross Sea— embraces part of the

Malayan, Antarctic, Falkland and Mexican

provinces, and the whole extension of the

Schopf

East-Australian, Tasmanian, New Zealand,

and Chilean provinces.

The bryozoan faunistic knowledge of the

Southern Ocean started at the very

beginning of the Nineteenth Century. Species

collected in the Magellanic region were first

published by Quoy £ Gaimard (1824) and

d'Orbigny (1841-1847); many others

gathered in the Australian seas, South

Atlantic, South Pacific and Indian Oceans

appeared in Busk's Catalogues of the marine

Polyzoa in the collections of the British

Museum (1852, 1854). The collections made

by the Challenger (Busk, 1881, 1884, 1886;

Waters, 1888) and by the vessels of several

other European Expeditions to the Southern

Seas (Ridley, 1881; Calvet, 1904, 1909;

Jullien, 1888; Waters, 1904, 1905; Kluge,

1914; Thornely, 1924; Livingstone, 1928;

Borg, 1926; Hastings, 1943; Borg, 1944)

greatly enriched the knowledge of the

Magellanic, Subantarctic and Antarctic

Bryozoa.

In the Australian area, the contributions by

MacGillivray (1887, 1895); Waters (1881-

1906); Maplestone (1899-1909) Hamilton

(1898) and more recently Harmer

(1915-1957); Marcus (1922); Livingstone

(1927-1929); Stach (1936-1938); Hastings

(1932); Brown (1952-1956); Silén (1954) and

Powell (1967), give a rather sound knowledge

of this area.

The South Pacific Tertiary Bryozoa are

well known in the Australian area

(MacGillivray, 1895; Waters, 1881, 1882;

Brown, 1952), but comparatively poorly

known in the South-American area where the

Tertiary Patagonian Bryozoa studied by Canu

(1904, 1908, 1911) need to be reevaluated in

the light of modern taxonomy. On the other

hand, the Antarctic Tertiary Bryozoa are

practically unknown although being necessary

for correlations and comparisons of both

recent and fossil South-Pacific Bryozoan

Faunas.

Genera! compiling and comparative work

have been done in the last decades (Hastings,

1943; Brown, 1952; Rogick, 1965; Powell,

1967; Androsova, 1968, 1972; Viviani, 1969;

Moyano, 1975, 1978, 1982) generating a

partial perspective of the South-Pacific

bryozoans, principally focused not only on the

Antartic and Subantarctic regions, but also on

the New Zealand area. Thus, the complete

perspective of the whole South-Pacific region

including the many archipelagos in its

vastness is far from being reached.

A contribution to that perspective

—making emphasis on the Chilean Bryozoan

Fauna— using the polymorphism and the

zoarial and zoogeographical parameters

outlined above, is the aim of this work.

MATERIALS AND METHODS

The information. used in this work has been

obtained from: a. old and recent works of the

Australian area sent to the author by Dr.

Robin Wass of the Universty of Sydney; b.

data derived from Viviant's 1969 and 1977

publications, and from Moyano 1982; c.

additional material collected along the

Chilean coasts.

A. THE SAMPLES

The studied samples have the features

indicated hereinafter and were collected at:

1. Easter Island (27%07'S; 109922'W): 1934,

coral blades bearing zoaria in their dorsal side;

collected by Dr. Ottmar Wilhelm G. at 76 m

depth; 1972, intertidal material on algae and

small stones collected by the author.

2. Juan Fernández Archipelago (33%38'S;

78%58'W): 1964, top of a guyot off Más a

Tierra island at 220-280 m depth; collectors

H.I. Moyano and E. Alarcón; 1966, intertidal

collection made at Más a Tierra island by A.

Angulo; 1967, a diving collection at 2-5 m

depth in Cumberland Bay, Más a Tierra

island, made by G. Sanhueza.

3. Caldera (27%03'S; 70%30'W): 1982, scallop

shells bearing species unrecorded at that area;

presented to the author by Mariana

Rodríguez.

4. Coquimbo (29%53'S; 71%19'W): 1964,

shells and stones collected in the intertidal

zone by the author.

5. Los Molles (32%12'"S; 71%27"W): 1980,

stones with several cheilostomatous Bryozoa,

obtained by diving at 8-10 m depth under a

Lessomia sp. “forest”; collector Eduardo

Villouta.

6. Algarrobo (33%21'S; 71%41'W): 1982,

stones collected at 8-12 m depth by diving;

presented to the author by C. Villalba.

7. Off Punta Nugurúe (35%40'S; 72%50'W):

1980, large stones collected by trawl at 350 m

depth by A. Rivera and A. Wendt; presented

to the author by the collectors.

8. Colvumo (36"32'S; 72%56'W): 1981; a large

stone collected at 8 m depth by diving;

presented to the author by Jacqueline

Fernández.

9. Reque Cove (36"45'S; '73“11"W): 1982,

rock pieces collected between 10-25 m depth

by diving; presented to the author by C.

Villalba and J. Fernández.

10. Off Lebu(37%37'S; 73%259'W): 1979, seve-

ral zoaria collected at 600 m depth by mem-

bers of the Department of Oceanology, Uni-

versity of Concepción, Chile.

11. Melinka (43%54'S; 73%45'W): 1980,

subtidal stones covered by crustose stony algae

and Bryozoa, collected and presented by E.

Bay-Schmith and C. Werlinger.

12. IFOP Ol Expedition: 1964,

oceanographical and biological expedition

undertaken from Coquimbo (29%35'S;

71219 W) to Chiloé island and Juan

Fernández Archipelago. The studied samples

came from Valparaíso to Concepción, and

from 37 m to more than 200 m depth. The

samples collected by trawl and Petersen 0.1 m?

grab sampler consist of small stones covered

by Bryozoa, solitary corals, ascidiae and

foraminifera. This expedition was sponsored

by the Instituto de Fomento Pesquero and the

Chilean Navy; and the samples collected by

the author, T. Antezana and E. Alarcón.

B. SYSTEMATICS

The systematic account includes all the

presently known species from Arica (18%29'S;

70%19'W) to Diego Ramírez islands (56%30'S;

68%45"W) the ones from Juan Fernández

islands and those from Easter island. For each

new species only a short diagnosis will be given

which includes the type numbers. The type

material will be deposited in the Museo

Zoológico of the Universidad de Concepción,

Chile (MZUC).

In Table 1, species indicated by a generic

name followed by “sp. n. Viviani” correspond

to those appeared in a Doctor Rerum Naturae

As they are recognized as good

taxonomical entities, and useful for

zoogeographical purposes, they will be cited

as such to avoid nomina nuda problems.

Thesis.

C. POLYMORPHISM AND ZOARIAL DIVERSII Y

The polymorphism of Cheilostomata was

evaluated following Schopf (1973) and

Moyano (1975, 1978). This procedure has

been also utilized regarding the

polymorphism of the Tertiary Bryozoan

Faunas from Argentina, South Australia and

New Zealand. When speaking of

polymorphism in general percentages for a

given area, no distinctions have been made

among vibracularia and avicularia.

The calculations of the zoarial diversity

were made following Moyano, 1978. When a

species shows more than one zoarial form, the

most frequent one is computed. The same

criterion has been used in evaluating the

Tertiary Bryozoan Faunas from Australia,

New Zealand and South America. The zoarial

diversity values of the Australian recent

Victorian Bryozoa are only approximated

owing to the dubious or incomplete

descriptions of the zoarial form in many

species of that area.

D. ZOOGEOGRAPHY

The zoogeographical affinity of the Bryozoa

along the Chilean coast was evaluated using

the Kulczinsky-2 index, following Sibouet

(1979), Monniot (1979) and Moyano (1982).

The Chilean South-American shelf was

arbitrarily divided into segments of four

latitudinal degrees each, except CH10 from

46%S to 52%S. Juan Fernández archipelago and

Easter island were considered as one unit

each. Two sources of species were evaluated

by the Kulczinsky-2 index: 1. Viviant's Chilean

littoral Ectoprocta which includes 52 species,

mostly collected in the intertidal zone from

Arica (18%59'S) to Quellón in the

southernmost part of Chiloé island (43%08'S),

and 2. Table I which contains 267 species

inhabiting the continental shelf from Arica to

Diego Ramírez islands plus those of Juan

Fernández and Easter islands.

In Table I, species marked* come from

beyond 200 m depth. The number of

“Magellanic” species appearing in Table Lis

smaller than that included in a previous work,

(Moyano, 1982) because were only considered

those collected inside Chilean waters and not

those present in the Patagonian shelf between

Tierra del Fuego and the Falkland islands.

RESULTS

A. THE NEW TAxaA

Antropora paucicryptocysta Sp. n.

Fig. 18

Diagnosis: Zoarium encrusting, unilaminar

to plurilaminar. Zooids thin-walled, with

gymnocyst and cryptocyst slightly developed.

Opesia covering almost the whole frontal

surface. No spines. Cryptocyst fimely

granulated. A pair of small triangular

avicularia at the proximal side of each

autozooid, but each zooid being encircled by

[e]

six avicularia, two of its own and four of the

neighbouring autozoords. Avicularian

mandibule very small and semicircular.

Ovicell not observed.

Species being different from the others

described by Osburn (1950) for the Pacific

coast of America due to the slight

development of the cryptocyst.

Holotype: MZUC 10953. One colony

encrusting a stone, off Punta Curaumilla

(33%06'S; 71%49'W); 137 m depth; October

1964. Collectors: H. Moyano and E. Alarcón.

Paratypes: MZUC 10954,

holotype collecting data.

one colony;

Arachnopusia areolata sp. n.

Fig. 4

Diagnosis: Zoarium encrusting, unilaminar,

white. Zooids with a complete frontal pericyst

having round, oval or reniform holes; one or

two weak small oral spines rarely present;

lateral and proximal borders of the pericyst

with a row of small areoles. One to three small

pointed or rounded avicularia on the distal

apertural border of the pericyst. Ovicell

hyperstomial with a more calcified distal belt

which has small pores on its proximal rim.

Species differing from A. monoceros (Busk)

and A. admiranda Moyano, for its smaller

avicularia, weak —1f present— oral spines and

for the row of marginal areoles.

Holotype: MZUC 10956, one colony

encrusting a large stone collected off Punta

Nugurúe (35%40'S; 72%50'W); 350 m, 1980;

Collectors: A. Rivera € A. Wendt.

Paratypes: MZUC. 10957, several colonies

encrusting small stones; off Punta Curaumilla

(33%06'S; 71%49'W); 137 m depth; October

1964; collectors: H.1. Moyano e E. Alarcón.

Aplousina gymnocystica sp. n.

Fig. 22

Diagnosis: Zoarium encrusting, unilaminar.

Zooids somewhat irregular, widely or

narrowly ovate or pyriform, separated by

deep furrows. Distal and lateral gymnocystal

borders elevated. Proximal gymnocyst

reaching one fifth of the zoecial length;

descending cryptocyst laterally narrow, but

proximally enlarged reaching to one fourth of

the zoecial length. With marginal, minute,

decidueous spines encircling the opesia.

Endozoecial ovicell widely open, formed by

two calcareous superposed sheets. No

avicularia; no dietellae. With a large

uniporous distal septula and two or three

more in each lateral wall.

Species clearly different from other

members of the genus Aplousma by the

development of the cryptocyst and

gymnocyst.

=J

Holotype: MZUC 9790, a colony encrusting

a valve of Chama sp. Cumberland Bay, Más a

Tierra island; April 1967. Collector: G.

Sanhueza.

Paratypes: MZUC 9796, two zoaria

encrusting an Arca sp. valve; Cumberland

Bay, Más a Tierra island; August 1965;

Collector: A. Angulo.

Cellaria humilis sp. n.

Figs. 15,16, 17

Diagnosis: Zoarium irregulary branching,

articulated; — joimts tubular; branches

appearing everywhere as lateral offshoots

through the autozooidal oral apertures; this

way of branching causes the zoarium to be

irregularly cereiform. Zooids long, roughly

hexagonal; four making a whorl around the

internodal axis. Oral aperture at the

beginning of the distal zovidal third; two

lateral parenthesis-like cryptocystal thin

elevations encircling it, but without fusing

themselves proximally or distally. Oral

aperture semicircular, the proximal border

gently arcuated, without proximal or distal

internal denticles. Operculum reinforced by a

complete marginal sclerite. Avicularia scarce,

small, triangular, one fourth of the frontal

zoecial length; with a semicircular mandibule

wider than long. Ovicell apertures not seen.

Species differing from the other austral

forms by its zoarial structure and small

avicularia.

Holoptype: MZUC 9863; a ramified zoarial

fragment; off Punta Nugurúe (35%40'S;

72%50'W);, 350 m depth; 1980; collectors: A.

Wendt £ A. Rivera.

Paratypes: MZUC 9864. Several internodal

fragments from the same locality of the

holotype.

Cribralaria labiodentata sp. n.

Fig. 5

Diagnosis: Zoarium encrusting. Zooids lar-

ge, white, glistening, separated by well mar-

ked furrows zoecial pericyst made of 12 to 14

frontal ribs; two rows of small pericystal holes

separating every two ribs. Oral frame with two

distal and two proximal spines; the two proxi-

mal with a serrated free border and the two

distal with a pair of proximally directed lateral

horns; the latter may be inconspicuos. Inter-

zoecial avicularia large, asymmetrical, roughly

squared latero-diagonally produced into the

asymmetrical avicularian rostrum; avicularian

mandibule large, falciform, reaching one half

of the zoecial length. Ovicell endozoecial, in-

mersed in the avicularian chamber or not.

This species diffterentiates from those of

the North Tropical Pacific and from those of

the Australiam and New Zealand areas by 1ts

larger avicularium and the two rows of pores

between each pair of frontal ribs.

Holotype: MZUC 9862; a large colony on a

coral blade; Easter Island; 76 m depth; 1934;

collector: Dr. Ottmar Wilhelm.

Escharoides molinal sp. nm.

Fig. 23.

Diagnosis: Zoariuium encrusting and

unilaminar, or eschariform being composed

of one or two lamellae. Zooids strongly

calcified becoming very thick, with imprecise

contours. Frontal wall a pleurocyst: the large

marginal areolae being separated by radiating

ribs. Secondary aperture proximally elevated,

with a V-shaped sinus. Primary aperture not

seen in frontal view and lacking any type of

proximal or lateral denticles. Some strong,

decidueous spines present in the distal part of

the secondary aperture. Two types of

latero-oral avicularia; a patr of small, pomted,

triangular distally directed no longer than the

apertural diameter; and one or two larger,

pointed or spathulated, pointing latero-dis-

tally, lying obliquely at one or both sides of the

secondary aperture; these larger ones with a

huge perforated avicularian chamber. Ovicell

imperforated, hyperstomial, having marginal

areolae separated by ribs; becoming inmersed

in strongly calcified zooids.

This species is dedicated to the abbott Juan

Ignacio Molina who wrote the first Natural

History of the Chilean Fauna at the end of the

Eighteenth Century.

It seems to be very akin to E. angela

(Hutton) and E. excavata (MacGillivray) from

the Australian area. It differentiates,

however, from the first one by the less

developed peristome and for the two distally

directed tiny avicularia and the two

latero-distal giant ones; from the second

because it lacks the central and lateral

denticles inside the primary aperture and for

frequently having four instead of two

avicularia.

Holotype: MZUC 9960, a very calcified

zoarial piece; Cumberland Bay, Más a Tierra

island, April 1967; collector: G. Sanhueza.

Paratypes: MZUC 10968, several small

pieces; off Cumberland Bay, Más a Tierra

island; 60 m; October, 1964; collector: H. IL.

Moyano.

Fenestrulina microstoma sp. n.

Figs. 3,9

Diagnosis: Zoarium encrusting, unilaminar,

white-yellow. Zooids large, hexagonal, very

inflated, separated by deep furrows. Frontal

wall a densely perforated tremocyst, leaving a

central imperforated area proximal to the

ascopore. —Centrally situated ascopore

separated from the oral aperture by two or

“three rows of pores; ascopore outline oval or

reniform without spines or internal denticles.

Small oral aperture separated from the distal

zoecial rim by two rows of pores; oral spines

rare. Ovicell hyperstomial, imperforated with

short marginal costae.

Species differing from £. malus: by its larger

size and the frontal wall almost completely

perforated; it differentiates too, from the two

new taxa of this genus proposed by Viviani

(1969), by its more centrally placed aperture

leaving more than one row of pores between it

and the distal zoecial rim.

Holotype:: MZUC 10979, one ovicelled

zoarium; off Punta Carranza, IFOP

Expedition (35%35'S; 72%42'W); 37 m depth;

November, 1964; collector: H.I. Moyano.

Paratypes: MZUC 10980, several colonies;

holotype collecting data.

Macroporella areolata sp. n

Fig. 8

Diagnosis: Zoarium encrusting, unilaminar,

light brown. Zooids large, almost hexagonal;

frontal wall a convex tremocyst pierced by

excessively small pores and, with apparent

marginal areoles. Frontal ascopore in front of

a low umbo, transversely oval. Oral aperture

relatively small. A latero-distally directed

avicularium with setiform mandibule. A

densely perforated hyperstomial ovicell with

marginal areoles.

Species characterized by its areolar pores

and the almost inexistent tremocystal ones.

Holotype: MZUC 10959, one encrusting

zoarium; Off Punta Curaumilla (33%06'S;

71%49'W); 137 m depth; October, 1964;

collectors: H.I. Moyano e E. Alarcón.

Paratypes: MZUC 10960, several colonies

encrusting small stones; holotype collecting

data.

Opaeophora brown sp. n.

Higsiz3/4Zo!

Diagnosis: Zoarium encrusting, small,

patchy, unilaminar. Zooids yellow-white

separated by deep furrows. Primary frontal

wall membranous; a complete cryptocyst with

a pair of lateral large round opesiules. Zoecial

aperture semicircular, the proximal border

straight, encircled by four long spines. Ovicell

hyperstomial, large and smooth. Large

interzoecial asymmetrical avicularia, slightly

shorter than autozoecia; avicularian

mandibule linguiform, laterally curved.

Circular ancestrula with a complete cryptocyst

pierced by two large round opesiules;

provided with marginal spines.

Species dedicated to Dr. A. Brown, eminent

Australian Bryozoologist. This species looks

very similar to O. lefada monopia Brown from

the upper Pliocene of New Zealand.

Holotype: MZUC 9698, Cumberland Bay,

Más a Tierra island; 5 m depth; April 1967;

collector: G. Sanhueza.

Paratypes: MZUC 9770, five small zoaria;

holotype collecting data.

do)

Parasmittina proximoproducta sp.n.

Figs. 6, 7.

Diagnosis: Zoarium encrusting. Zooids

rhomboidal to irregularly squared. Frontal

wall granulated, with small scattered pores

being a pleurocyst. Oral area elevated and

strongly produced proximally; primar;

aperture with moderate wide lyrule and a pair

of lateral cardelles; secondary aperture

bordered proximally by a tall vertical

elevation and by four distal spines; with a long

vertical furrow along the internal side of the

oral vertical elevation. Two lateral oral

narrowly triangular avicularia, one bemg

always larger than the other; one or more

additional and similar ones may be present

around the apertural area. Hyperstomial

ovicell globular, not inmersed.

Species different from other similar from

the Indopacific area (Soule an Soule, 1973) by

its long proximal elevation on the proximal

apertural side.

Holotype: MZUC 9860; Easter Island, a

zoarium on back side of a coral blade; 67 m

depth; 1934; collector: Dr. Ottmar Wilhelm.

Paratypes: MZUC 9861. Holotype collecting

data.

Phylactella problematica sp. n.

Fig. 19:

Diagnosis: Zoarium encrusting, unilaminar,

white. Zooids large, strongly convex. Frontal a

tremocyst with many small pores; areolae not

clearly set apart. Oral area produced into a tall

peristome. Primary aperture without lyrula or

cardelles; secondary aperture proximally

notched. A globular, densely perforated,

hyperstomial ovicell looking as dependence of

the distal part of peristome. No avicularia was

observed.

Species notoriously different from those

present along the Chilean coast, with dubious

generic relations, therefore it has been

tentatively adscribed to the genus Phylactella.

Holotype: MZUC 10961, a colony encrusting

a stone; off Punta Curaumilla (33%06'S;

71%49'W); 137 m depth; October, 1964;

collectors: H.I. Moyano é E. Alarcón.

Paratypes: MZUC 10962; several colonies

encrusting stones; holotype collecting data.

Reteurgula zoecrulifera sp.n.

Fig. 10.

Diagnosis: Zoarium encrusting, unilaminar

loosely pluriserial. Zooids elliptical, separated

by interzoecial

membranous,

spaces. Frontal wall

lateral

cryptocyst; gymnocyst proximally developed.

covering a Narrow

Uwo small pointed spines on both sides of the

oral valve. Many small zoeciules scattered

between zooids; these have a round or

elliptical opesia «covered by a frontal

membrane; zoeciule size variable measuring

one fourth to one half of the zoecial length.

No avicularia or avicularian mandibules on

zoeciules. Hyperstomial ovicells round, with a

very large dorsal non-calcified area.

Species close to a Tertiary New Zealand

torm of R. acuta (Hincks) referred to by

Brown (1962), but lacking the avicularia

present in the fossil form. Species easily

identifiable by the large amount of zoeciules

separating the autozoids.

Holotype: MZUC 9865; a colony encrusting

a stone; off Punta Curaumilla (33%06'S;

71%49'W); 137 m depth; October, 1964;

collectors: H.I. Moyano € E. Alarcón.

Schizoporella maulina sp. n.

Fig. 12.

Diagnosis: Zoarium encrusting, thick,

unilaminar, white. Zooids hexagonal, slightly

convex; frontal wall a tremocyst with evenly

distributed small pores. Oral aperture

schizoporelloid, with a wide U-shaped

proximal sinus. Hyperstomial ovicell closed by

the operculum having a great less calcified

transversely oval frontal area; periovicellar

marginal pores present. Avicularia lacking.

Species adscribed to the genus Schizoporella,

considered in a wide sense.

Holotype: MZUC 10963, a large zoarium

encrusting a stone; off Punta Carranza

(35%35'S; 72%42'W); 35 m depth; November

1964, collectors: H.I. Moyano £ T. Antezana.

Paratypes: MZUC 10964; several colonies.

Holotype collecting data.

10

Smittina fragaria sp. n.

Figs. 20, 21.

Diagnosis: Zoarium encrusting, unilaminar,

white, glistening. Zooids rectangular to

hexagonal with straight rims; frontal wall a

tremocyst, without distinct marginal pores.

Oral area moderately elevated; primary

aperture lacking a lyrule but with two lateral

denticles; secondary aperture at the end of a

short peristome bearing a small proximally

directed avicularium with a short semicircular

mandibule. Hyperstomial ovicell with a large

frontal densely perforated exposed area,

these pores are relatively large.

This species has been adscribed to the

genus Smitina, considered in a very wide

sense.

Holotype:: MZUC 10958, one zoarium

encrusting a stone; off Punta Curaumilla

(33%06'5; 7149 W); 137 m depth; October

1964; collectors: H.I. Moyano «e E. Alarcón.

Smiíttina jacquelinae sp. n.

Figs. 11, 14.

Diagnosis: Zoarium encrusting, unilaminar,

light yellow. Zooids hexagonal, gently convex;

frontal wall an evenly pierced tremocyst. Oral

area elevated in a proximally notched

peristome; the two lateral sides of the

peristome may coalesce leaving a suboral

orifice; primary aperture with a slightly

denticular small lyrule. A proximal oral

avicularium completely or partially inside the

peristome, settled just on the proximal notch;

avicularian mandibule distally rounded,

slightly linguiform. Hyperstomial ovicell

becaming inmersed, with one or more small

frontal pores. In very young zoecia there are

four oral spines.

This species is a S. purpurea (Hincks)-like

form, but being different in its colour and the

poorly developed lyrule. On the other hand,

this species has a large distribution from

Melinka (43%54'S; 73%45'W) to Los Molles

(329125; 71%40'W) and it may grow side by

side the “typical” S. purpurea (Hincks).

Holotype: MZUC 10975, a large zoarium on

a stone; Reque Cove (36%45'S: 73%11 "W);

10-20 m depth, 1982; collectors: César

Villalba € Jacqueline Fernández (to whom the

species is dedicated).

Paratypes: MZUC 10976, one zoarium on a

stone, Melinka, 1980; collectors C. Werlinger

and E. Bay-schmith. MZUC 10977, an

encrusting zoarium; Los Molles; 8-10 m

depth; 1980; collector: E. Villouta.

Smittina undulimargo sp. n.

¡NS llo

Diagnosis: Zoarium encrusting, unilaminar,

thick, brown. Zooids strongly calcified,

shortly after their arising at the marginal

growing rim. Mature zooids with an

undulated border, clearly set off by a very

narrow brown ribbon following the border

and frequently crossing the inmersed ovicells

in a diagonal way. Frontal wall nearly flat,

evenly perforated. Apertural area not

produced or elevated. Primary aperture with

a moderate to wide lyrule flanked by two

poined lateral denticles. Secondary aperture

proximally notched, the peristome being as

thick as the frontal thickness; with a minute,

proximal-obliquely directed oral avicularium

which is not seen from above owing to its place

inside the peristome. Hyperstomial ovicell

deeply inmersed, with an irregular row of

radial large pores.

A typical smittinid distinctly different from

all the others along the Chilean coast for its

wavy outline.

Holotype: MZUC 10965, a colony encrusting

a stone; off Punta Carranza (35%35'S;

72%42'W); 35 m depth; November, 1964;

collectors: H.I. Moyano and T. Antezana.

Paratypes: MZUC 19066, some colonies with

the holotype colecting data.

Smittina volcanica sp. n.

Fig. 13

Diagnosis: Zoarium encrusting, unilaminar,

white and shining. Zooids large, irregularly

quadrangular, lateral rims wavy; a thin

epitheca covers the whole frontal wall.

Frontall wall a densely perforated tremocyst;

oral apertural area elevated in a tall peristome

with four marginal spines, proximally

notched. Primary aperture with a narrow

lyrule but lacking cardelles. A tiny median

elongated, oral avicularium inside the

peristome, proximally and upwardly

directed. Ovicell appearing as a huge inflation

of the distal part of the peristome; its exposed

walls having the same constitution of the

frontal zooidal wall, so it looks as 1f it were

inmersed in the distal zooid as an endozoecial

ovicell.

This species seems to be different from all

the other known smittinids im the South

Eastern part of the Pacific Ocean. It is

considered as part of the genus Smttina, by the

presence of a lyrule and the oral avicularium

placed over it and inside the peristome.

Holotype: MZUC 10978, one small zoarium

on a stone; off Punta Curaumilla (33%06'S;

71%49'W); 137 m depth; October 1964;

collectors: H.1. Moyano é E. Alarcón.

Smittina jullieni n. nom.

Smttia purpurea Jullien, 1888:54; pl. 2, Fig.

4.

Not Smittia landsborovi (Johnston)

purpurea Flincks 1881:123.

var.

Diagnosis: As described and illustrated by its

author.

This species has a light purpurine colour

making it different from the other smittinids

present along the Chilean coast. It

differentiates from the typical Smittina

purpurea (Hincks) apart its lighter colour for

having a very small oral avicularium

completely inside the peristome and for its

tendence to become very calcified.

Spiroporina reteporelliformis sp. n.

Figs. 24, 27.

Diagnosis: Zoarium vinculariform, white,

calcareous, irregularly branching. Branches

with two faces, the frontal one having

autozooids and the dorsal one showing

kenozooids and avicularia. Zooids irregularly

hexagonal; frontal wall a pleurocyst with

scattered marginal pores. Primary aperture

with a U-shaped proximal sinus; secondary

aperture with a pair of small triangular,

distal-laterally directed avicularia.

Hyperstomial ovicell, somewhat inmersed, its

frontal wall lacking calcification. Dorsal

(basal) zoarial wall covered by one or more

kenozooidal superposed sheets. Ovate

avicularia on dorsal kenozooids.

Species having Reteporellina-like

Reteporella-like form; clearly set off from other

Spiroporima species for developing a dorsal

zoarial face provided with kenozooids and

avicularia as in the species of Reteporidae.

Holotype: MZUC 9666, a broken colony; on

a guyot off Más a Tierra island; 220-280 m

depth; October 1964; collectors: H.I. Moyano

8c E. Alarcón.

Paratypes: MZUC 9667, more than eighty

complete colonies.

or

fragments and small

Holotype collecting data.

Cnisia parunternodata Sp. n.

Figs. 38, 39.

Diagnosis: Zoarium small, white, articulated

and branching. Free zooidal ends frontally

and laterally curvated; zoecial aperture

circular to ovate. Joints with short, disciform

yellow nodes. Sterile internodes having 1-5

zooids, those with one or two in the basal part

of the colony being more common. The new

branches alternate to right and left, budding

from the first autozooid of internodes with 3-5

zooids; fertil internodes with 4-7 zooids, the

first producing most of times, a lateral branch,

and the second being always the gonozooid.

Pyriform gonozoovid with a terminal

oveciostome adnate to the third autozoecial

tube in the internode; oecial aperture

transversely oval facing frontally. Some

autozooids with spiniform processes.

A small species, densely branched looking

like C. eburnea (L.), but differing from it by the

small number of autozooids per internode

and by its proportionally smaller gonozooids.

Fertile zoaria measure 2,0 to 3,4 mm high;

gonozooidal length 0,7 to 0,95 mm;

gonozooidal width 0,22 to 0,3 mm.

Holotype: MZUC 9792; one zoarium with

three gonozooids; Punta Loberías,

Cumberland Bay, Más a Tierra Island; April

1967; 2-5 m depth; collector: G. Sanhueza.

Paratypes: MZUC 9798; seven small fertile

zoaria. Holotype collecting data.

112

Frondipora masalierrensis Sp. N.

Fig. 36.

Diagnosis: Zoarium vinculariform,

branched in one plane, white, becoming

retiform by fusion of its branches. Stems

giving short and alternate branches, but

frequently irregular. Zooids in fascicular

clusters facing frontally and laterally,

projecting beyond the lateral border; among

fascicules there are nude areas without zoecial

tubes. Autozooidal tubes long, prismatic in the

fascicles and circular when traversing the

gonozooid. Dorsal zoarial side without zoecial

apertures but with clear cut zoecial

boundaries and pseudopores. Gonozooids as

large as wide, pyriform or irregularly circular

structures, developed between fascicles and

traversed by many autozooids. Oeciostome in

the centre or in the distal third of the

gonozooid, slightly oval, associated with an

autozooidal tube, from which it differentiates

by its larger diameter and facing

proximally.

Species differing from other Frondipora

species by its large gonozooid evenly traversed

by zoecial tubes.

Holotype: MZUC 9662, a reticular zoartal

piece; off Más a Tierra island; 220-280 m;

October 1964; collector: H.I. Moyano.

Paratypes: MZUC 10967, zoarial fragments;

Más a Tierra island; 60 m depth; October

1964; collectors: H.I. Moyano « E. Alarcón.

for

Desmeplagioecia irregularis sp. N.

Figs. 30, 31.

Diagnosis: Zoarium encrusting, fan shaped,

very flat, widening distally, producing here

and there lateral offshoots. Zooids emerging

from a flat zoarial surface, distally and

upwardly directed; emerging isolated in the

younger part of the zoarium but becoming

connate throughout forming clusters or

fascicles towards the distal and mature zoarial

part; clusters composed from two to six zoo1ds

each; zooids stout, very calcified. Gonozooid

nearly flat, wider than long, small or very

large; two or more gonozooids seemingly

coalesce forming a larger one; this gonozooid

is usually traversed by isolated zooidal tubes;

oeciostome short and distally outflared,

issueing from the central part of the

gonozooid, always proximally connated to an

isolated autozooidal tube; its distal transverse

oval aperture facing proximally and

upwardly.

This species seems to be more related to the

genus Desmeplagivcecia than to Tubulipora. It

has the zooidal clusters and the large

transverse gonozooid of the former and the

fascicles of the latter. Considering the

gonozooid structure as the basis of the

cyclostome taxonomy this species comes

nearer to Desmeplagioecia than to Tubulipora. 1t

differentiates from D. lineata (MacGillivray)

inhabiting the Magellanic and Australian

regions by its fan-shaped instead of circular

zOarium.

Halotype: MZUC. 10973, a large zoarium

measuring 18 mm in its larger diameter;

Reque Cove (36%45'S; 7321 1''W); 20-25 m

depth; April 1982; collectors: J. Fernández

and C. Villalba.

Paratypes: MZUC 10974, several small

ovicelled zoaria; Algarrobo (33%21'S;

71%40'W); 8-12 m depth; May, 1982;

collectors: J. Fernández and C. Villalba.

Tubulipora protewa sp. n.

Figs. 32, 33

Diagnosis: Zoarium encrusting,

white-bluish, composed or irregularly

branched lobules, widening where a

gonozooid develops and becoming narrow

distally. A completely developed zoarium

becoming irregularly star-like, with lobules

touching each other or fusing. Zooids

0,100-0,125 mm diameter, their free end

rather produced, disposed in alternating

series 2-6 tubes each, connate throughout or

becoming free and separated at their distal

ends; the zooidal tube pattern alterates where

a gonozooid exists forming irregular series or

tascicles. Gonozooid large, lobulated; the

ramifyimg lobules occasionally fuse encircling

series or fascicles; veciostome cylindric, 0,075

mm diameter, not terminal; shorter, equal or

longer than an autozoectal tube, associated

with a tube or fascicle.

Species differing from the others described

for the Magellanic region and the Pacific coast

of America due to its zoarial pattern and large

gonozooids.

13

Holotype: MZUC 10969, one large and

irregularly stellate zoarium, 8.4 mm Xx 7.2 mm

having nine gonozooids; Los Molles (32%12'S;

71%27'W); 8-10 m depth, 1981; collector: E.

Villouta.

Paratypes: MZUC 10970, three zoaria

encrusting a stone. Holotype collecting data.

Tubulipora tuboangusta sp. n.

Fig. 37.

Diagnosis: Zoarium encrusting, long and

not bifurcated, or bifurcating into two

opposed adhering elongated lobules; a basal

lamina provided with several rows of

apertures. Zooids disposed in transverse,

oblique alternating series composed each of

two to seven connate zoecial tubes, the more

common having'5-6 per series. The series may

fuse at the end of zoarial lobules originating

transverse fascicles. Gonozooid inflated and

dilatated between the series and fascicles;

oecial tube long, narrow, one half to one third

of the autozoecial diameter; adnate only by its

base to the more internal tube of a series.

The name of the species indicates its

narrow and long oecial tube, making it

different from 7. stellata Busk, T. anderssoni

Borg and 7. spatiosa Borg from the Magellanic

region.

Holotypes: MZUC 9788, one zoarium

measuring 6 mm diameter; Punta Loberías,

Cumberland Bay, Más a Tierra island; April

1967, 2-5 m depth; collector: G. Sanhueza.

Paratypes: MZUC 9778; same collecting

data as the holotype.

Tubulipora tubolata sp. n.

Figs. 34, 35.

Diagnosis:

bluish,

Zoarium encrusting, white-

regularly stellated from its very

beginning; wide zoarial lobes composed of

regularly ordered and alternating zoarial

series, 3-5 zooids each. Zooids stout

0,125-0,200 mm diameter, very calcified;

their free ends moderately long. Gonozooids

fairly abundant, inflated pyriform to

irregular producing lobes beyond the

oeciostome, occasionally encircling one or

more zooids; oeciostome large, clearly set off,

strongly dilatated and distally outflared,

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23

facing proximally, associated to an

autozooidal tube.

Species related to 7. stellata Busk and T.

anderssoni Borg but being different in the

form of the oecial tube.

Holotypes: MZUC 10971, a large stellate

zoarium, 16 mm diameter, having 18 lobules;

Los Molles (32%12'S; 71%27'W), 1980, 8-10 m

depth; collector: E. Villouta.

Paratypes: MZUC 10972,

zoaria. Holotype collecting data.

two stellated

B. THE CHILEAN BRYOZOAN FAUNA

This fauna, excluding the antarctic species, IS

composed of 19 of the order

Ctenostomata (7.11%), 46 of the Cyclostomata

(17.22%) and 202 of the Cheilostomata

(75.65%). These 267 species (Table 1) inhabit

the continental shelf from Arica (18%29'5S;

70%19'W) to Diego Ramirez islands (56%30'S;

6845'W) and the shelves of the Juan

Fernández and Easter islands. As the

bryozoan assemblages in each of these areas

are different, they will be analized separately

in the folowing paragraphs.

species

l. The Magellanic Bryozoan Fauna: The

general features of this fauna have been

recently revised by Moyano (1982). It may be

summarily characterized as having 7.69% ot

ctenostome species, 23.59% of cyclostome

species and 68.75% of cheilostome species.

These figures indicate a proportionally larger

amount of cyclostome forms when compared

with the whole Chilean Bryozoa. The

Cheilostomata show 74.62% of species having

one or more polymorphs, and the total fauna

a zoarial diversity (H'H'max.100) reaching

62.95%. Endemic elements are the families

Calvetiidae and Pseudidmoneidae, and the

genera Andreella Jullien, Flustrapora Moyano,

Orthoporidroides Moyano and Johetina Jullien.

Zoogeographically it extends to the Southern

Atlantic Ocean reaching the archipelagos

Tristan da Cunha and Falkland,

Kerguelen in the Indian Ocean.

and

2. The Northern and Central Chale Bryozoan

Fauna: A general account of it was first given

by Viviani (1969). This author studied the

litoral entoprocts and ectoprocts from Arica

(18%29'S) to Quellón (43%08'5) describing 52

ectoproct species. This number Is Now

24

elevated to 107 species. This assemblage, as

now understood is characterized by the

excessive predominance of encrusting species,

which make up the main bulk of any sample

collected from the intertidal zone to the

border of the continental shelf (ca. 200 m

depth). Taxonomically, the ctenostome

ectoprocts comprise 7 species, the cyclostomes

12 and the cheilostome 88. The intertidal

forms are almost the same from Arica to

Quellón, because many have very large

latitudinal ranges, for instance Alcyoniduum

mamillatum, A. polyoum, Bowerbanka gracias,

Buskia setigera, Bicrisa edwardsiana, Aetea

anguina, A. ligulata, A. recta, Beania magellanica,

Bugula neritina, Cauloramphus — spuaferun,

Celleporella — hyalna, Chapenella acanthina,

Electra hastingsae, Fenestrulima malusa, Happothoa

flagellum, Hippothoa divaricata, Membrampora

hyadest, M. tuberculata, Macroporella cihata,

Scruparia ambigua, Smttina maplestoner, Smttina

purpurea and Umbonula alvarexana.

Several new species sensu Viviani pertaining to

the genera Aetea, Aplousma, Fenestrulina,

Hippaliosina, Lagenicella and Membranaipora,

also have a great latitudinal distribution (See

Table D.

The study of several samples from 35 m

depth to more than 350 m depth has revealed

the existence of a relevant bryozoan faunule

altogether different from the intertidal and

the upper subtidal ones. In it, many of the new

species appearing in Table 1 were found.

Some of the new and the known species

belong to genera hitherto unknown or nearly

«so for the Chilean fauna, namely: Antropora,

Bellulopora, and

“Phylactella”.

Several species uknown north to 425 were

now discovered as normal components of the

encrusting communities at 10-30 m depth in

central Chile. Among these, two are

particularly important: Inverstula — nutrix

Jullien, a typical antarctic and subantarctic

species now extending to 30% along the

Chilean coast, and Romancheina martala,

hitherto known only from the southernmost

Chilean archipelagos, proves to be a very

common encrusting element at 10-20 m depth

near Concepción (368). This means that it 1s

necessary to explore and systematically study

Retevirgula, Arthropoma

the narrow Chilean continental shelf from

1898 to Chiloé island.

3. The Juan Fernández Archapelago Bryozoan

Fauna: In 1921 Marcus published the first

account of the Bryozoa collected by several

expeditions to these islands. He reported 20

species including 5 Cyclostomata and 15

Cheilostomata. Now that number elevate to 41

as indicated in Table 1, and comprises 1

ctenostome (2.43%), 13 cyclostomes (31,7%)

and 27 cheilostomes (65.85%). The

cyclostome component is marked by two

diastoporids known only from the Magellanic

area and by Frondipora masatierrensis sp. N.,

which is the first record of the genus for the

eastern south Pacific. The Cheilostomata

characteristically show two species of Caberea,

the subantarctic Ogivala elegans, the

australasian Microporella lumifera and

Arthropoma — biseriale, the north Pacific

Crassimarginatella kumatae, and new species of

the genera Escharoides Opaeophora, Aplousima

and Spiroporima. Among these Spiroporina

reteporelliformis sp. n. shows a reteporidan

condition in having vibice-like kenozooids

covering the rear part of the zoarial branches.

The endemic elements include at least 10

species making 24.39%.

4. The Easter Island Bryozoan Fauna: In a

previous paper (Moyano, 1973) ten species

comprising one cyclosttome and nine

cheilostomes were reported. Now that

number is elevated to 16. It is most likely that

many species remain to be discovered because

only two sets of samples have been examined.

One littoral survey yielded the first ten known

species, and a second one on coral blades

obtained beyond 50 m depth provided the rest

described here. This whole and incomplete

assemblage is principally characterized by 3

smittinids, 3 cosmopolite forms and 3

cribrimorphs, with an endemism reaching

25%. The analysis in Table 1 proves that this

fauna is composed of 8 (50%) of species

having a pantropical or cosmopolite

character.

C. ZOOGEOGRAPHY OF THE CHILEAN BRYOZOAN

FAUNA.

Having a core of 80 “bryozoan

(ectoprocts plus endoprocts) and

species”

a large

bibliographic information, Viviani (1969)

proposed a zoogeographical scheme for the

Chilean coast. This scheme included four

provinces: a. Peruvian (2*S-20%S), b. Chilean

(205-4095), c. North-Patagonian (405-4698),

d. South-Patagonian (4695-5695).

If we take such a fact into account: a.

Vivian's scheme did not considered the

subtidal bryozoans until 200 m depth, which is

the average extension of the submarine

environment normally used in zoogegraphical

analises (Ekman, 1953; Briggs, 1974; Schopf,

1979), b. the number of ectoprocts has

duplicated since the Viviani's study; c. the

Peruvian bryozoan fauna was in 1969 and is

until now almost unknown and, therefore, not

allowing a clear and sound zoogeographical

relation between it and the Chilean bryozoan

fauna, and d. the convenience of comparing

the Bryozoa from Juan Fernández and the

continental ones, makes it necessary to deeply

study the Chilean Bryozoogeography.

In Table II Viviani's Chilean ectoprocts are

zoogeographically analized through the

values of the Kulczinsky-2 index applied to

each pair of compared localities from A to R.

On the left side of the diagonal we find the

numbers of common species for each pair of

areas, and on the right, the corresponding

Kulczinsky-2 index values. If the locality A is

examined, it is possible to see that all the

localities have more than 50% of affinity;

locality E with 31 species shows the same

feature as well as localities C, D and F. If now a

median locality is analized, for instance K, the

same feature is present; the same is true of the

species richest one, i. e. locality L. These

results confirm the general impression that an

observer of the intertidal fauna of the Chilean

coast obtains, that is, the existence of a large

amount of common species extending from

the far north to a point between Concepción

and Chiloé island. From Table IH it is possible

to see that the zoogeographical affinity of

locality A steadily decreases to the south, and

the reverse occurs when looking at the values

of locality R which decrease towards the

north.

Table III analyzes by means of the

Kulczinsky-2 index values all the known

Chilean ectoprocts listed in Table I, that have

been recorded during more than a century

Table Il

ZOOGEOGRAPHY OF THE CHILEAN ECTOPROCTA AFTER THE DATA BY

VIVIANI (1969) USING THE

KULCZINSKY-2 INDEX.

Localities ABCDEFGH Nik TE MINO POR

(Number of species per locality) 90 20 23 29 31 29 20 14 22 26 25 33 22 1 18 14 19 16

A. Arica (18*28'S). 65 79 71 65 67 60 66 57 62 67 56 57 0 57 61 61 51

B. Iquique (2012'S) 13 79 88 82 80 85 66 72 79 72 71 62 0 68 60 62 51

C. Tocopilla (22%02"S). E a 90 83 86 79 57 80 74 79 66 67 0 69 63 67 58

D. Antofagasta (23%35'S). 17 21 23 87 86 76 69 76 80 78 71 64 0 68 64 65 58

E. Coquimbo (30*03'S). 16 20 22 26 87 78 67 78 74 76 69 62 0 66 62 64 57

F. Los Molles (32”18'5). 16 1922 :25::26 42 69 80 77 82 75 68 0 72 64 74 63

G. Zapallar (32%50'5). 12 7 17 18: 19 10 61 76 75 72 64 67 0 69 67 62 56

H. Isla Negra (33%28'5). LALO ES 18 138 10 64 66 67 61 64 0 70 79 74 7

I. Llico (34%45'S). 12 15 18 19 19 20 16 11 71 81 76 68 0 71 70 69 59

J. Hoca (34%55'5). 14-18-18;22121 21 47 12 17 75 86 63 52 75 71 64 71

K. Tomé (36%40'5). 15: 16: 19:21 21 22 16-12 19 19 77 68 0 76 72 74 62

L. Mehuín (39*20'5). 14 18: 18: 22 22 23: 1612 20. "25-22 76-52 17 IN TIRA

M. Pelluco(39%35'5). 12 19; 15.16 15 17 14 11 15 16: 16:20 0 80 82 74 81

N. Carboneros(39%50'S). 070.00. .070 0-0 0-41 0 150 53 54 0 53

O. Maullín (41%40'S). 11 138 14 15 15 16 13: 11 14 16 16 18. 16 1 89 76 83

P. Pargua (41%47'S). 10) MASAS 18 11 12 18138 14 187 ASA 81 87

Q. Lacúi (41%50'S; 74%03'W). 12 14 14 15 16 17 12 12 14 14 16 18 15 0 14 13 75

R. Quellón (43%08'S; 73%35"'W). 9 10 11 12 12 13 10 11 11 14 12 16 15 1 14 13 13

and a half of bryozoan research. Table HI also greater — affinity with the Panamic

includes several localities from Perú to Alaska

in the eastern Pacific and from the south

Atlantic to Australia in the south Pacific.

Localities l and L show very small affinity

values with all other regions becoming

separate entities. Locality R compared from A

to L shows still lower values; but these increase

rapidly to locality Q making evident the

obvious relations between the subantarctic

and antarctic areas when compared with the

Chilean bryozoan fauna. Locality F has great

affinity with localities G, H, J, Kand M, that is,

from Arica (18%S) to a point near Chiloé

island; the same is true of each of these six

geographical points. Locality N is closer to

locality M, which is closer in turn to the six

indicated above. On the other hand, locality O

is closer to P, which in turn is closer to Q.

s The above analysis indicates that the

Chilean bryozoan fauna may be divided in

four zoogeographical groups, namely: a.

Easter Island Bryozoa (Locality 1); b. Juan

Fernández islands Bryozoa (Locality L); c.

North and Central Chile Bryozoa (Localities

F.G, H,J,K, Mand N), and d. Southern Chile

Bryozoa (Localities O and P). The first has

zoogeographical area owing to its cosmopolite

and pantropical elements; the second is

simultaneously closer to the Galapagos

islands, New Zealand and Australia because of

its large amount of cosmopolite or

“panpacific” components. The third, North

and Central Chile, shows a northernmost

subgroup composed of the localities F, G, H,

being closer each other probably because their

littoral elements are the only ones known; a

second subgroup of the following three J.K.

and M is the richest in species number and the

core of the non-Magellanic Chilean bryozoan

fauna; locality N being the link with the

Magellanic region exhibits, however, closer

affinity to localities M an K owing to the fact

that its known elements are principally

littoral, nevertheless, species obtained in

deeper waters —A. giganteum for instance—

favours its inclusion with localities O and P.

The fourth, including localities O and P, has

the richest and most diversified bryozoan

fauna; this is probably related to a larger

disponibility of coast and continental shelf. It

near 200 species extend to the South Atlantic

and Indian Oceans (Moyano, 1982), which is

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27

reflected by the Kulezinsky-2 index values of

locality Q in Table HI.

According to the evidences acumulated so

far, the Chilean Bryozoan Fauna can be

divided into four provinces:

a. Chilean: from Arica to Chacao Channel

(1895-4295).

b. Magellanic: from Chacao Channel to

Diego Ramírez islands (42%5-56"30'5).

c. Juan Fernández Islands.

d. Easter Island.

The limit between the Chilean and Magellanic

provinces is only approximated, because most

of their known Bryozoa are shallow water

forms including many littoral ones. Species

obtained at deeper waters have proved to be

typically Magellanic. This is the reason for

considering the Chacao Channel, where the

austral archipelagos begin, as the

southernmost limit of the Chilean bryozoan

province. Probably the actual limit be at a

point near 40%S, as suggested by several

zoogeographers when discussing the general

zoogeography of the Chilean coast (Ekman,

1953; Briggs, 1974).

D. THE SOUTHERN PACIFIC BRYOZOAN FAUNAS.

In the following account these faunas will be

featured and compared in a threefold

scheme: the taxa, the polymorphism and the

zoarial diversity.

The taxa: From a taxonomic point of view

the Southern Pacific Bryozoan Fauna can be

divided into three large groups

corresponding to the Australian, Antarctic

and South-American elements.

a. The Australian area has the largest

number of species reaching near 400 in the

southern part of the Australian continent

(MacGillivray, 1887, 1895); ca. 350 in the

northern coasts between the continent and

New Guinea, and amounting to 640 between

the Philippines and the former (Schopf et al.

1978). This vast area has a number

endemic, or nearly so, families such as

Eurystomellidae, Euthyrisellidae,

Conescharellinidae, Euthyroididae and

Parmulariidae. On the other hand the

Reteporidae, Adeonidae, Onchoporidae,

of

Orbituliporidae and Catenicellidae are

principally represented there.

Families giving a special distinction to this

area are the Catenicellidae,

Conescharellinidae and Reteporidae. Their

species are characterized by very specialized

zoaria and high polymorphism values and

contribute to a large extent, to the great

zoarial diversity of the Western Pacific

Bryozoa.

The cyclostome Bryozoa are also well

represented both fossil and recent

(MacGillivray, 1887, 1895). Some genera are

seemingly endemic to this area, viz. Favosipora

and Densipora. Other specialized forms, e.g.

Crisma radians and Desmeplagioecia lneata,

make connections with the Antarctic and

Subantarctic regions respectively.

The Australian Bryozoan Fauna as a whole

features corresponding more

K-selected species owing to high

polymorphism, what means a keen zoecial

division of the work at colonial level (Table

IV), a higher zoarial diversity (Table V) and

an extreme taxonomic diversity of the species

compared in number and specializations with

other regions of the world. (MacGillivray

1887, 1895; Harmer, 1926, 1933, 1957; Bock,

1982).

to

Its

shows

b. The antarctic and subantarctic zones

have been carefully characterized by Hastings

(1943), Borg (1944), Rogick (1965),

Androsova (1968) and Moyano (1975, 1978).

Rogick indicated, with some hesitation, the

existence of 321 species inside the Antarctic

Convergence. Among these, 179

considered endemic (55.94%), being

composed of 1 entoproct, 2 ctenostomes, 18

cyclostomes, 89 anascan cheilostomes and 69

ascophoran cheilostomes.

To these figures many new species have

been added in the last years (Moyano, 1978)

causing the endemism to be higher. In fact,

along the Antarctic peninsula the endemism

varies from 70% to 90% on a basis of 128

species (Moyano, 1978).

Taxonomically the Antarctic Bryozoan

Fauna is marked by a large number of species

belonging to the genera: Amastigia, Beanta,

Camptoplites, Notoplites, Cellaria, Chapenella,

Cornucopina, Cellarinella, Escharoides, and

Smittina. At family level, this fauna is sharply

were

Table IV

POLYMORPHISM OF SOME RECENT PACIFIC CHEILOSTOMATOUS BRYOZOAN FAUNAS

Species

Faunas

number

No poly-

morphs

l or more 2 or more 3 or more

polymorphs polymorphs polymorphs

Philippine region

(Canu € Bassler, 1929). 296 (100%)

Indo-pacific area

(Harmer, 1926, 1934, 1957). 437 (100%)

Antarctic region

(Moyano, 1975). 194 (100%)

Magellanic region

(Moyano, 1982). 134 (100%)

Chile, 42%S-18%S

(Viviani, 1969; Moyano) 91 (100%)

Galapagos islands

(Osburn, 1950, 1952, and 1953). 126 (100%)

Arctic region

(Kluge, 1962). 200 (100%)

defined by the Bugulidae, Scrupocellariidae,

Flustridae, Cellarinellidae, and Smittinidae.

Among these, Cellariidae has the endemic

genera Cellanaeforma, Mawsonia and

Paracellaria. Cellarinellidae shows only one

species —C. dubha Waters— beyond the

Antarctic Convergence to the north. The

family Reteporidae is fairly well represented

with 8 species; in this feature the Antarctic

Bryozoa come nearer the Australian fauna

where the largest number of reteporids exists.

c. The South-American coast from the

Galapagos islands to the south shows at least

two different bryozoan faunistic components:

the more austral is also the more studied one

having abouth 200 species (Moyano, 1982),

and the northernmost one from ca. 40%S to the

Galapagos is beginning to be known. In this

study more than 100 species are adscribed to

it. The Galapagos Bryozoan Fauna itself

comprises 150 species and is closer to the

Panamic zoogeographical region, so it shares

most of the species with the Gulf of California

(Osburn 1950, 1952, 1953, Soule, 1963:

Moyano, 1982). The continental shelf from

Guayaquil (02%00'5S) to Arica (18%69'5) IS

68 (22.97%)

66 (15.10%)

37 (19.07%)

34 (25.37%)

41 (45.05%)

30 (23.80%)

57 (28.50%)

2

1

228 (77.03%) 52 (17.56%) 3 (2.36%)

y

r

371 (84.90%) 109 (24.94%) 37 (8.46%)

157 (80.92%) 46 (23.71%) 15 (7.73%)

100 (74.62%) 34 (25.37%) 2 (1.49%)

50 (54.94%) 7 (7.69%) 3 (3.29%)

96 (76.20%) 31 (24.60%) 6 (4.76%)

143 (71.50%) 31 (15.50%) 4 (2.00%)

poorly known but itis possible to presume that

it has, at least in part, a mixed bryozoan fauna

composed of panamic, peruvian and chilean

elements. (See Table 1).

The polymorphism: "The measure of

polymorphism can feature a given Bryozoan

Fauna reflecting its evolutive history (Schopf,

1973). This criterium has been used by Schopf

to characterize tropical and polar cheilostome

faunas. According to this author the defensive

polymorphs are increasingly K-selected from

the arising of the Cheilostomata up to the

present time (Schopf, 1977). On the basis of

these ideas, a measure of the polymorphism

for Recent and Tertiary Pacific Bryozoan

Faunas is presented.

Table IV shows that the Indo-Pacific and

the Antarctic have the highest values, and

Chile has the lowest ones. Highest values seem

to be associated with the tropics, being the

Antarctic an exception. When considering the

polymorphism of the Tertiary Bryozoan

Faunas (Table V) it is possible to see that it

increases from South-America to Australia,

but New Zealand has a larger proportion of

species having two or more polymorphs. The

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possible significance of these figures will be

discussed later.

The zoarial diversity: Stach (1936, 1938)

developed a new approach for considering

the bryozoan species as a whole within a given

faunule. He classified the extant bryozoan

colonies in nine zoarial forms, namely:

Membraniporiform, Petraliform,

Catenicelliform, Eschariform, Reteporiform,

Vinculariform, Cellariform, Flustriform, and

Lunulitiform, correlating them with

particular conditions existing where they

grow. He applied this view to estimate the

batimetry of Lower Cenozoic formation

fossils. From the beginning he stressed the

idea of the variability of the zoarial form

possessed by certain species, for instance,

Caleschara denticulata (MacGillivray) which

becomes eschariform in shallow waters of the

Southern Australian coast, but it becomes

vinculariform from 30-200 fathoms in Bass

Strait. The number of zoarial forms has

grown from the times of Stach to the present

day, but the basic types he described are the

more common and easier to identify.

In previous works (Moyano, 1978, 1982)

the zoarial diversity of both tropical and polar

Recent Bryozoan Faunas was calculated.

These calculations showed that different

values were present, being higher in the

Indo-pacific and Antarctic regions. This

finding was associated to environmental

stability. It was also demonstrated that the

highest values of Zoarial diversity correspond

to the highest figures of polymorphism.

Now, in this work the same ideas have been

applied to several bryozoan faunas of the

Pacific basin. In Table VI, the main Pacific

Bryozoan Faunas from the Arctic to the

Antarctic and from the western to the eastern

continental shelves are compared in this

sense. The zoarial diversity expresed as H/H'

max attains the highest values from the

Antarctic to the Philippine islands

(71%-82,6%), median values from the

Magellanic area to the Arctic, and the lowest

one for 107 species of the Chilean coast

between 18%S to 42%S. This means that there

are two different large assemblages around

the whole Pacific basin when considering the

zoarial types and the probability of finding

one of them in studying a given fauna; one

Sl

includes the tropical Western Pacific plus the

Antarctic, and the other the Eastern cold

temperate, temperate and tropical Pacific plus

the Arctic. If the idea that environmental

stability favours higher zoarial diversity values

were true, then the Eastern Pacific Bryozoa

Faunas would have born a certain degree of

environmental uncertainty having been

higher than the one affecting the Western

Pacific Faunas. This idea can hold true when

considering that one of the world areas

supposed to have been very unstable during

the last million years is the Arctic, where the

zoarial diversity attains low values. Then, why

the Chilean coast Bryozoan Fauna has the

lowest value? One response could be: the

incomplete knowledge of its fauna, and the

other one supposes this area to have been

affected by great environmental unstability

during the Tertiary.

On the other hand, the Argentinean

Tertiary Patagonian Bryozoa exhibit ca. 78%

eveness whereas New Zealand has 72.9% and

South Australia 83.23% when analyzing the

Cheilostomata plus the Cyclostomata. If only

the zoarial diversity of the Cheilostomata is

analyzed, the Argentinean figure diminishes

to 74.58% and the Australian one increases to

84.39%. If the Tertiary Argentinean

Bryozoan extended to the present Magellanic

region, then, during the Patagonian in the

Tertiary period the environmental conditions

were different, since the present zoarial

diversity values are lower. In the Australian

area, the Victorian Tertiary Bryozoa had

higher diversity values than those from today,

indicating conditions not very different from

the Recent ones. Nevertheless, it 1s always

necessary to remember that the fossil record

has always the danger of being incomplete

and then conducing to wrong deductions, if

not properly evaluated.

From the threefold analysis just finished

one aspect seems to emerge: the Western

Pacific Bryozoa are marked by high values of

polymorphism, zoarial diversity, and

taxonomic diversity. This characteristic can be

also extended to the Antarctic. Thus,_the

Antarctic and the Australian area converge in

this sense although being far apart today. This

convergence also extends to the family

Reteporidae. which has its greater

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diversification in both areas; in comparison,

the Magellanic area has at most two

reteporidan species which reach to 8 in the

Antarctic and more than 30 in the Australian

area.

Although the differences stressed above

between the western and easter sides of the

Pacific basin, many connections among them

can be established when studying their taxa.

The physical connecting element providing

relatively similar conditions in the Suban-

tarctic Pacific area is the West-Wind-Drift.

Probably, to its intermission 33 species have a

circumaustral or nearly so distribution

(Moyano, 1982).

Each of the four Chilean bryozoan

provinces defined above has some common

elements with the Australian area. The

Magellanic province has the 33 species already

indicated. The Chilean province shares the

genera Retevirgula, Heteropora, Opaeophora,

Arthropoma, Aspidostoma, Plesiothoa,

Crepidacantha, Cryptosula, Ellisna, Escharordes,

Hippomenella, Hippopodmella, Inversiula,

Osthimosia, Spiroporina, Vittaticella and Bicrisia.

The Juan Fernández the

following species the

Australian area: Arthropoma biseriale (Hincks),

Caberea zelandica (Gray), Celleporima costata

(MacGillivray), Crepidacantha crimspina

has

in

province

also present

Levinsen, Chapenella cervicornas (Busk),

Hippomenella vellicata (Hutton), Maicroporella

lunifera (Haswell), Spiropormma pentagona

(d'Orbigny), Smittina maplestone:

(MacGillivray) and Smittina purpurea (Hincks).

The Easter island province shares with the

Australian the genera Escharnma,

Cribralaria and Canda.

area

DISCUSSION

The Chilean Bryozoan Fauna is defined by a

large amount of encrusting species and low

values of zoarial diversity and polymorphism.

A close examination of its components and its

distribution shows that there is a Magellanic

zone, richer in species and having higher

polymorphism and zoarial diversity values,

and a Northern zone with less species and very

small values of the other parameters. The

Magellanic area is directly influenced by the

West-Wind-Drift waters, and the shelf north

to 40% under the regime of the cold

Humboldt current which flows north

affecting the western South-american coast

reaching Guayaquil. This cold flow deflects

the superficial isotherms to the north; being

that corresponding to 20"C. near the

Equatorial Line. Thus the whole Chilean coast

is affected by cold waters. Nevertheless, the

presence of warmer, more saline and less

oxigenated waters running to the south under

the Humboldt current is not negligible for

their effects acting on pelagic and benthic

species under 100 m depth along Central and

Northern Chile. This has been tested with

mesopelagic fishes (Craddock and Mead,

1970) and itis probalby the explanation of the

33

presence to the latitude of Valparaiso of the

tropical bryozoan Bellulopora bellula.

If the present oceanographic conditions

had been present during the whole Tertiary

period or during large part of it, making this

area more stable, then the Bryozoan Fauna

perhaps would have had polymorphism and

zoarial diversity figures similar or near to the

present antarctic ones. Fleming (1979) states

that during the Neogene, the Chilean Fauna

was hit more than other cool temperate

regions by thermic and faunistic oscillations

causing many Panamic and Indo-pacific

elements to arrive from the north during

warmer periods, and which were extinct

during the cooler ones along the Ice Ages.

These facts mean instability and, probably,

these are the cause that have featured the

extant Chilean Bryozoa, easily detected in its

low polymorphism and zoarial diversity.

Taxonomically the Chilean Bryozoan

Fauna shows traces of the facts depicted

above, being composed of Indo-Pacific,

Panamic Subantarctic, Antarctic and endemic

elements. Some species apparently show an

antitropical distribution, viz. Membramipora

isabelleana (2 M. uillosa) associated to

Macrocystis kelps, as do too M. hyadesi. 1

Nicholson's (1979) view is correct, this species

came from the Northern Hemisphere when

the genus Macrocysti invaded the Southern

Hemisphere.

The division of the Chilean Bryozoan

Fauna into four provinces, two continental

and two insular, results from the data in

Tables, 1, Hand HI. This agree with most of

the zoogeographical models already proposed

for the Chilean Fauna (Ekman, 1953;

commented and summarized in Briggs,

1974). The present knowledge does not

support Viviani's proposal of dividing

continental Chile into four provinces. In the

littoral zone (intertidal and upper subtidal)

many species —those wich Viviani dealt

with— go far south, whereas in the sublittoral

below ten meters depth others go far north.

The littoral Retevirgula areolata reaches as far

as the Guaitecas archipelago (445), whereas

the antarctic and subantarctic Inversuula nutrix

reaches 3098, and the magellanic Romanchema

martiali 368. This fact would support the

existence of an intermediate “mixed”

province between the Magellanic and the

Northern one, composed of a mixing of

southern and northern elements.

Nevertheless, the study of samples collected

below 35 m depth to 200 or more m depth

shows that in Central Chile there exists a

particular Bryozoan fauna different from the

Magellanic one at the same depth. The

probability exists that north Valparaíso the

same species which have been found between

Lebu and Punta Curaumilla may appear at

100-150 m depth.

The present-day bryozoan diversity

around Australia is substantially higher than

in the South-american coast. Polymorphism

and zoarial diversity have the highest values in

Australia and in the Indo-Pacific region. This

feature also characterizes the Antarctic. An

interpretation of the latter could be to

consider the long and stable isolation of the

Antartic being separated from Australia by

the circumantarctic current (30-25 Ma; Grant

Mackie, 1979), Although the extreme

environmental conditions now present in the

Antarctic, its persistence for several million

years rendered it an stable environment; this

probably caused the bryozoan species to be

K-selected, what means slow growth, long life

as in Cellarinella species forming annual marks

that indicated that they have a life span of

several years, and the development of very

specialized avicularian and vibracularian

polymorphs. Australia was drifting to the

north while the Antarctic and the water mases

around it were progressively cooler, this fact

provided it with a more favourable

environmental condition than that occurring

at that time in the southermost part of South

America (Grant Mackie, 1979; Kennet in

Knox, 1979).

If environmental unstability is one of the

factors that induces species to be r-selected

and stability to become K-selected (Pianka

1970, in Schoptf, 1977), then the Bryozoan

Faunas from the Antarctic to the Philippines

would translate an evolutive history having as

frame a long environmental stability. On the

contrary, the Bryozoan Faunas from the

Arctic to Magellanic region are marked by low

zoarial diversity and lower values of

polymorphism showing a more r-selected set

of species indicating a less stable environment

in the long term.

The closer Tertiary connections between

Australia and Antarctica are perhaps still

observed in the existence of a relative large

amount of reteporidan species. Reteporine

species are highly specialized exhibiting an

important degree of polymorphism with up to

four different types of avicularia. On the

other hand, three of the Antarctic

Reteporidae belong to the genus Happellozoon,

wich is also represented in New Zealand,

meaning perhaps, some connections between

these islands and the Antartic, at least, in their

hydrological conditions during, the early

Pliocene to the late early Pleistocene (Knox,

1979).

CONCLUSION

The present study further shows that:

a. The Chilean Bryozoan Fauna comprises

267 species, including 19 Ctenostomata

(7.11%), 46 Cyclostomata (17.22%) and

202 Cheilostomata (75.65%).

b. Twenty four species and a name are

described as new:

ba. The Cheilostomata Aplousina gym-

nocystica sp. ne Antropora

paucicryptocysta sp. n., Arachnopusia

areolata sp. n., Cellaria humilis Sp. n.,

Cribralaria labiodentata sp. n., Escha-

rowdes molina: sp. n., Fenestrulina mi-

crostoma sp. n., Macroporella areolata

sp. n., Opaeophora brown Sp. N.,

Parasmittina proximoproducta sp. n.,

Phylactella problematica sp. n., Retevir-

gula zoeciulifera sp. n., Schizoporella

maulina sp. n., Smittina fragana sp.

n., Smittina jacquelinae sp. n., Smittina

undulimargo sp. n., Smittima volcanica

sp. n., Spiroporina reteporelliformas sp.

n.

bb. The Cyclostomata Crisia paruinter-

nodata sp. n., Desmeplagioecia irregu-

laris sp. n., Frondipora masatierrensis

sp. Nn., Tubulipora proteica sp. n.,

Tubulipora tuboangusta sp. n. and

Tubulipora tubolata sp. n.

bc. The new name Smiítina jullieni n.

nom. for the preoccupied Smittia

purpurea Jullien, 1888.

. From a bryozoogeographical point of

view the Chilean fauna is divided into

four units or provinces:

ca. Magellanac, from Chacao Channel to

Diego Ramírez islands (42%

-5630'S).

cb. Chilean, from Arica to Chacao

Channel (ca. 18%S-429S).

cc. Juan Fernández, (3338'S; 75%52'W).

cd. Easter island, (27%07'S; 109%22'W).

. From a taxonomic point of view, in the

Eastern South Pacific (Ocean, the

families Reteporidae, Vittaticellidae,

Adeonidae and Scrupocellariidae are

poorly represented in relation to the

Australian area, whereas, the Eurysto-

35

mellidae, Euthyrisellidae, Conescharel-

linidae, Euthyroididae and Parmula-

riidae are altogether absent.

e. In the whole Pacific basin plus the

adjacent polar areas the polymorphism

of the Cheilostomata and the zoarial

diversity of all marine living bryozoan

orders attain highest values in the

western side from the Antarctic to the

Philippines, and lowest ones from the

Magellanic area to the Arctic.

f. The calculated figures of polymorphism

and zoarial diversity for the Tertiary

Bryozoa from Argentine, New Zealand

and South Australia also increase to the

west. Nevertheless, the actual zoarial

diversity figures of New Zealand being

slightly lower than the Argentinean

ones.

g. On the basis of zoarial diversity and

Cheilostomata polymorphism, the

Western Pacific plus the Antarctic have a

more specialized -that 1s, a more

K-selected- Bryozoan Fauna reflecting

perhaps a history of greater stability in

the long term. The Eastern Pacific, on

the contrarv, shows substantially lower

values; this implies more generalized or

more r-selected bryozoan species

extending from the Magellanic region to

the Arctic.

h. The lowest polymorphism and zoarial

diversity figures of the whole Pacific

basin belong to the Chilean coast

between 18% and 42%S. This could be

explained, at least in part, by the great

variability of the environmental

conditions during a large part of the

Tertiary.

ACKNOWLEDGEMENTS

The author is deeply indebted to the

institutions and personnel of the Facultad de

Ciencias Biológicas y de Recursos Naturales,

Universidad de Concepción, Chile, whose

cooperation and assistance made this work

possible: the Chilean Navy; Departaments of

Zoology and Oceanology, Universidad de

Concepción; Instituto de Fomento Pesquero;

Dr. Jorge Artigas, Prof. Fidel Jeldes, Dr. Ariel

Gallardo, Prof. Enrique Bay-Schmith, Prof.

Andrés Angulo, and the collectors already

cited in Materials and Methods.

Dr. D. A. Brown of the Australian

University, Canberra and Dr. R.E. Wass of the

University of Sydney, are particularly

thanked for help to resolve systematic

problems (D.A.B.) and for sending valuable

papers on Australian Bryozoa (R.E.W.).

The author also thanks constributions of

Mónica Orellana and Miguel Angel Alarcón

(Departments of Zoology and English,

Universidad de Concepción) for reading and

revising the manuscript, José Bustos for

redrawing the plates and Mrs. Lily Yánez for

secretarial assistance.

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to)

Figs. 1, 2: Smttina undulimargo sp. n., Fig. 1: Mature ovicelled zoecia, Fig. 2: oral apertures at the growing end. Fig. 3:

Fenestrulina microstoma sp. n., ancestrula and first zooid. Fig. 4: Arachnopusia areolata sp. n. Fig. 5: Cribralaria labiodentata sp. n.

Figs. 6, 7: Parasmittina proximoproducta sp. n. Fig. 8: Microporella areolata sp. n.

40

Fig. 9: Fenestrulina microstoma sp. n. mature zooids. Fig. 10: Retewrgula zoecnulifera sp 0 Lugo 11.14: Smittina jacquelinae sp.

n. Fig. 11: New zooids at the growing end; Fig. 14: mature zooids. lg. 12: Sehrzoporella marndina sp. nm. Fig. 13: Sivttima

volcanica Sp. n.

41

e

Figs. 15,16, 17: Cellaria humilis sp. n. Fig. 18: Antropora paucionyptocysta sp. nm. Fig. 19: Phylactella problematica sp. an. Figs. 20,

21: $mittma fragaria sp. n. Fig. 22: Aplousina gymnocystica Sp. 0.

42

Fig. 23: Escharoides molinai sp. n. Figs. 24, 25,26, 27: Spiroporina reteporelliformas sp. n. Fig. 24: frontal zoartal view. Fig. 25

dorsal zoarial view. Fig. 26: frontal and dorsal views of growing up. Fig. 27: transverse section of a branch showing three

kenozooidal layers. Figs. 28, 29: Opaeophora brown sp. n.

43

. NL ] : E : . o

Figs. 30, 31: Desmeplagivecia irregularis sp. n. Fig. 30: large broken gonozooid. Fig. 31: Two small gonozooids. Figs. 32,33:

Pubulipora proterwca sp. n., Fig. 32: ovicelled zoarium; Fig. 33: a gonozooid on a widening lobe. Figs. 34, 35: Tubulipora tubolata

sp. n.

Tubulipora

45

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