Proceedings 9th International Coral Reef Symposium, Bali, Indonesia 23-27 October 2000, Vol. 1
Coral barnacles: Cenozoic decline and extinction in the Atlantic/East Pacific
versus diversification in the Indo-West Pacific
A. Ross1 and W.A. Newman1
The pyrgomatid coral barnacles, first appearing in the late Oligocene of the western Atlantic, underwent a
Miocene diversification unparalleled by any other group of sessile barnacles. Diversification in the Indo-
Pacific (eastern Tethys) coincided with retreat of the tropics from higher latitudes, especially in the
Atlantic. Fragmentation of the tropics, due to the breakup of the Tethys seaway, and wholesale
extinctions of their host corals beginning in the Oligocene of Europe, Mediterranean and eastern Pacific
resulted in relictual distributions and regional endemism. This was followed by Neogene extinctions of
many host coral genera in the western Atlantic which were not replaced by originations. The exceptional
diversity of pyrgomatids now evident in the Indo-Pacific was tied to the survival and radiation of the
corals found there. Curiously, our knowledge of pyrgomatid numbers and diversity has shifted from the
Indonesian to peripheral centers of distribution.
Keywords Cirripedia, Pyrgomatidae, Biogeography.
1 Scripps Institution of Oceanography, La Jolla, California 92093-0202, USA
The pyrgomatids are obligatory symbionts or parasites
that have undergone numerous adaptations for living on
some 200 different coral, hydrozoan and sponge hosts
(Ross and Newman 1973, Ogawa and Matsuzaki 1992).
Aside from an extensive fossil record they are well
represented today by 67 living taxa (Figs. 1-2).
The pyrgomatids, beginning with a six-plated wall and
separable opercular plates (Newman and Ladd 1974a),
and progressing to a single-plated wall with two inse-
parable opercular plates (Table 1) had an archaeobalanid
ancestry. This is based on shell morphology, growth
patterns (Ross and Newman 1973, 2000), cirral and other
behaviors (Anderson 1992), and sperm ultrastructure
(Healy and Anderson1990).
Fig. 1 Extinction and reliction versus radiation in the Pyrgomatidae. Within the Mediterranean, Atlantic and eastern
Pacific, six species are extant and more than 30 extinct whereas none is extinct in the Indo-Pacific. Comparing these
faunas there are 10 times the number of Indo-Pacific species; our studies suggest well over 100 species. Numbers of
species given here reflects work in progress; Table 1 lists only nominal taxa.
Table 1 List of nominal pyrgomatids; genera and higher taxa in approximate phylogenetic order, species alphabetically;
asterisk (*) denotes type species, dagger (†) extinct taxa. Geologic time range follows name of each genus. Wall and
opercular plate patterns (6 x 4; 4 x 4; 1 x 4 etc.) are indicated diagramatically and each applies to the taxa immediately
following. Ceratoconcha voksae (early Miocene, Chipola Fm.), C. aderca (early Pliocene, Tamiami Fm.) and
Moroniella cystosa (early Pliocene, Tamiami Fm.) from Florida, mentioned by Zullo and Portell (1992b), not listed
below, are nomina nuda.
Pyrgomatid paleobiogeography reflects adaptations
to the diversity and distribution of their hosts. The history
of the corals and their exceptional diversity in some areas,
and decline or extinction in others, is tied to tectonic and
climatic changes bordering the Tethys seaway (Rosen
1984, Wilson and Rosen 1998), and the narrow regime
within which they thrive. The global maximum species
diversity of corals today is in the Indo-Pacific (Veron
1995). Although Indo-Pacific pyrgomatids underwent a
concomitant rapid diversification during the Neogene the
wide spectrum of fossils suggests a pre-Miocene radiation
no later than the early late Oligocene in the Caribbean
(Zullo in litt., cf. Wilson and Rosen 1998).
Tropical provincialism began with the collision of
Africa and Eurasia (20-17 Ma), severing the Indian Ocean
from the Atlantic (Rosen 1984). This was followed by the
Messinian "salinity crisis" (6 Ma) resulting in the
extinction and later replacement of the tropical Medi-
terranean fauna by a warm temperate one. The partial
closure of the Indonesian (ca. 7 Ma; Wilson and Rosen
1998) and complete closure of the Panamic seaways (3.5
Ma) completed development of tropical provincialism for
the pyrgomatids and their hosts. The western Atlantic
coral fauna suffered a further decline, the "Plio-
Pleistocene faunal turnover," in which about 90% of the
Mio-Pliocene species and 37% of the genera went extinct
with no new generic originations since (Budd et al. 1993).
Eoceratoconcha first occurs in the early Miocene of
Jamaica, and then middle Miocene of Trinidad (Newman
and Ladd 1974a), and Pliocene of Florida (Zullo and
Portell 1990, 1991). None is known from the Paratethyan
region, and not one is living today. Thus, the center of
origin for the Ceratoconchinae could well be the
The derived Ceratoconcha appear in the Oligocene of
Puerto Rico (Zullo in litt.), early Miocene of Jamaica
(Newman and Ladd 1974a), Florida (Zullo and Portell
1991, 1992), middle Miocene of Trinidad (Newman and
Ladd 1974a), Mio-Pliocene of Cuba (Withers 1953), late
Pliocene of Florida (Brooks and Ross 1960, Weisbord
1972, Zullo and Portell 1991), and Pleistocene of
Barbados (Withers 1926). In the Mio-Pliocene there were
several species of Ceratoconcha in southeastern Cali-
fornia at the northern end of the Imperial Seaway
(33°57'N, unpubl.). There are at least four living species,
known from the western Atlantic: Brazil (Young 1988),
Belize (Highsmith et al. 1983), Trinidad (Bacon 1976),
Dominican Republic-Haiti (DesMoulins 1867), Jamaica
(Scott 1987), Barbados (Scott 1987), Texas (Pequenot and
Ray 1974), Florida (Pilsbry 1931, Wells 1966) and
Bermuda (Zullo et al. 1972, Bromley 1978, Southward
Based on fossil evidence, Ceratoconcha reached its
greatest diversity (15 spp.) in the middle to early late
Miocene in Paratethys (Baluk and Radwanski 1967a,
1967b), other records are for Spain and Algeria ( Moisette
and Saint Martin 1990). Extinction in Europe and north-
ern Africa, stemming from the interaction of tectonics,
glacio-eustatic sea level changes, and local climate
followed during the Messinian "salinity crisis".
Fig. 2 Known Indo-Pacific localities having two or more taxa. Japan and the Ryukyu Is. Have more species, but less
generic diversity compared to Australia. No taxa occur in Hawaii. Eleven fossil taxa in five extant genera (late Miocene
to Holocene) occur in Japan, Ryukyu Is, Fiji and Marshall Is., but likely represent living forms. Fractions indicate
actual number of genera/species known at a given locality. Dot on the right, beyond 140°W and below the 20°S tick
mark represents records from the Tuamotu Is. The limited number of genera in the Indo-Malayan region likely reflects a
collecting bias in our data.
Central American plate tectonics late in the Miocene
led to closure of the trans-isthmian seaway between the
Caribbean and eastern Pacific (Pindell and Barrett 1990).
The recent, poorly developed, scattered and relatively
young reefs in the eastern Pacific contain almost wholly
Indo-Pacific taxa (Veron 1995). A combination of paleo-
tectonic activity, followed by rapid sedimentation,
increased upwelling and shoaling of the thermocline
(Kennett et al. 1985, Newman 1992) likely accounts for
their demise there. This niche was soon occupied by a few
eurytopic Indo-Pacific coral taxa and by the late Pliocene
to Recent archaeobalanid, Hexacreusia (Ross 1962, Zullo
1967, Zullo et al. 1972, Johnson and Ledesma-Vázquez
Existing evidence supports a ceratoconchine dispersal
from the Caribbean to the Mediterranean. Based on
negative evidence, further support comes from the
absence of Ceratoconcha in the Red Sea or bordering
Megatrematines, lacking any Miocene records, occur
in the early Pliocene of Sicily (Moroni 1967), late Plio-
cene of Crete (Baluk and Radwanski 1967c), and the Plio-
Pleistocene of Italy (Withers 1953). They also occur in
the Coralline Crag of England (Tilbrook 1997) and
Pleistocene to Holocene deposits in Japan (Sakakura
1934, Asami and Yamaguchi 1997).
Closure of the Iberian portal likely signaled the
demise of Megatrema in the Mediterranean (6 Ma). With
refilling of the Mediterranean, M. anglicum, now ranging
from the British Isles (Rees 1962, 1966) to Nigeria
(Stubbings 1967) northern Angola (Zibrowius in litt.), and
eastward to Meteor Bank (Young 1998), Madeira and
Canary Is. (Zibrowius in litt.), re-invaded and is the only
nominal megatrematine known to occur there (Relini
1980, Moisette and Saint Martin 1990, Roca 1992),
although there appears to be another species at Oran,
Algeria (cf. Relini 1980). In northern Africa, M. anglicum
extends from Oran, (Moisette and Saint Martin 1990) east
to Cap Bon, northeastern Tunisia (Zibrowius in litt.).
Parenthetically, the notion that M. anglicum in Japan is
the same as that now found in the eastern Atlantic has not
been convincingly demonstrated (cf. Ogawa and Mat-
suzaki 1994, 1995, Asami and Yamaguchi 1997).
There is one record for a Pleistocene megatrematine
in the Falmouth Formation on the north coast of Jamaica
(Portell in litt.) but no records for the eastern Pacific. Why
they seemingly never ranged this far to the west, despite
favorable westward currents (Iturralde-Vinent and
MacPhee 1999), is unknown. The sole species in the
Caribbean today, M. madreporarum, occurs in Brazil
(Young 1988), Tobago (Bacon et al. 1984) Bonaire
(Southward 1975) Jamaica (Scott 1987), Barbados (Scott
1987) and Florida (Ross and Newman 1973).
In the Indo-Pacific, living taxa are known from the
Great Barrier Reef (unpubl.), Rottnest I., Western
Australia (Jones 1993), Japan and Ryukyu Is. (Utinomi
1967, Ogawa and Matsuzaki 1995, Asami and Yamaguchi
1997) South China Sea (Ren 1986) and Gambier Is
(unpubl.). There are records (Broch 1931, Hiro 1935,
Nilsson-Cantell 1938) for what purports to be M.
anglicum, but the descriptions or discussions of these are
inadequate and likely apply to taxa now found in Japan
(Utinomi 1967, Ogawa and Matsuzaki 1995). Despite a
presence in the Indo-Pacific, megatrematines never
attained a diversity comparable to the pyrgomatines.
By no later than the middle Miocene the seaway
between the Mediterranean and Indian Ocean was severed
(Rögl 1998), thereafter the pyrgomatid faunas in the two
provinces became sharply differentiated. Although
recognizably different, migration from one to the other
region was possible up until this time, but Ceratoconcha,
from which the pyrgomatines are likely derived, failed to
survive in the Indian Ocean. The wholly Indo-Pacific
Pyrgomatinae includes three morphologically and
ecologically distinct tribes (Table 1; Ross and Newman
Pyrgopsellini. - Sponge-inhabiting pyrgomatines are
not likely to fossilize because they have a chitinous basis
and a partly chitinous wall. Among the known species,
one is from the Andaman Is., another from Hong Kong
and the Philippines (Rosell 1975).
Hoekiini. - This tribe includes wholly parasitic
species which occur only on a few species of Hydnophora
(Ross and Newman 1995, 1999, Ross 2000). If there is a
fossil record, if not by their characteristic and fragile
amoeboid wall, it will likely be for the calcareous portion
of the basis, which has an amoeboid outline.
Pyrgomatini. - This tribe encompasses some 60 or so
species, and has a diversity of shell characters far greater
than any other pyrgomatids. Fossil records include Nobia
and Savignium, from the late Miocene and Pleistocene of
Fiji and the Marshall Is. (Pilsbry 1945, Newman and Ladd
1974b, Newman et al. 1976), and Trevathana in the
Pleistocene of Japan (Mimoto 1991). All others are from
Holocene terraces in Japan, including Cantellius, Dar-
winiella and Galkinia (Asami and Yamaguchi 1997). The
paucity of records provides no clue as to the time of
origin and subsequent diversification of the Indo-Pacific
fauna, but noteworthy, genera representing modern forms
appear in the late Miocene.
Pyrgomatids probably originated in the Paleogene of
western Tethys and hence extant populations in the
Atlantic and Indo-Pacific represent Tethyan relicts.
Despite broad scale extinctions in the west, the greatest
diversity of forms occur in the Indo-Pacific where
pyrgomatids underwent an extensive radiation. Western
Atlantic pyrgomatids, like their coral hosts, are far less
diverse than they were during the Neogene (Budd et al.
1993). By comparison, there are some 100 coral genera in
the tropical Atlantic, and about 1000 in the Indo-Pacific,
and instructively the proportion of pyrgomatids in the two
regions, 7 to 69, is comparable.
Contrary to previous studies (Ross and Newman
1973, Newman et al. 1976) the distribution patterns of
either Indo-Pacific genera or species does little to amplify
their history. The center of distribution was once thought
to be the Malaysian Triangle. However, this "center of
distribution" has disappeared for pyrgomatids, partly
because our knowledge has shifted largely from
expedition-based collections to research centers that are
generally peripheral to the deep tropics.
Acknowledgments This paper is dedicated to H S Ladd
(1899-1982), V.A. Zullo (1936-1993) and H. Utinomi
(1910-1979) all of whom contributed in many ways. We
thank colleagues, friends and institutions for providing
specimens over the past 40 years. Financial support was
provided, in part (to AR), by the American Museum of
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