Breviora

US ISSN 0006-9698

Cambridge, Mass. 14 December 2016 Number 550

NINETEENTH AND EARLY TWENTIETH CENTURY EXPEDITIONS: THEIR
EXPLORATIONS IN BRAZIL, WITH THE FIRST CONTRIBUTIONS TO
KNOWLEDGE OF THE HYDROZOA

P. A. Grohmann,1 A. L. Quijada,1 and D. R. Calder2

Abstract. Prior to 1800. exploring expeditions were usually of a commercial and political nature, undertaken
to colonize or expand relations with new countries and territories. Among those to be explored as the 19th century
progressed was Brazil, the largest nation in South America. This account is a brief historical overview of some
19th and early 20th century expeditions to Brazil, focusing on those that included investigations of hydrozoans
(phylum Cnidaria). Among the more important of these expeditions were /’ Expedition de La Coquille (France), the
Voyage of the Beagle (Great Britain), the United States Exploring Expedition, I’Expedition HSWMS Eugenie
(Sweden), the Thayer Expedition (United States), the Challenger Expedition (Great Britain), the Albatross
Expedition 1887-1888 (United States), the Scottish National ( Scotia ) Antarctic Expedition, the British Antarctic
Expedition, and the Hartt Expedition (United States). Maps of routes taken and names of expedition commanders
and scientists are provided in the supplemental materia! online. Also included is a list of hydrozoan species
collected in Brazilian waters by the expeditions (with original and current names of taxa), together with dates and
exact locations of collection, station numbers with coordinates and depths (when available), and the museums
where material is presently housed, with respective catalog numbers and sources of information. Citations of
sources of updated species names are also provided. Figures (diagrams and old plates prepared by expedition
illustrators) are added.

Key words: 19th century expeditions; 20th century expeditions; Brazilian coast; Cnidaria; Hydrozoa; exploration;
natural history

1 Departamento de Zoologia, Instituto de Biologia da
Universidade Federal do Rio de Janeiro, CCS, Bloco
A, Ilha do Fundao, Rio de Janeiro, Brazil, CEP
21941-590; e-mail: grohmann@biologia.ufrj.br.

2 Department of Natural History, Royal Ontario
Museum, 100 Queen’s Park, Toronto, Ontario,
Canada M5S 2C6; e-mail: dalec@rom.on.ca.

INTRODUCTION

Geographically, Brazil is the largest coun-
try in South America and fifth largest in the
world. Its discovery and development
opened rich opportunities for studies in
natural history. Fundamental advances in
knowledge of the biota of the country were

The President and Fellows of Harvard College 2016.

?

BREVIORA

No. 550

made by pioneering scholars such as the
naturalists Jose Mariano da Concei^ao
Veloso (Frei Veloso) (1742-181 1), Georg
Heinrich von Langsdorff (1774-1852), Carl
Friedrich Philipp von Martius (1794-1868),
Charles Wilkes (1798-1877), Alcide Charles
Victor Marie Dessalines d’Orbigny ( 1 802—
1857), Jean Louis Rodolphe Agassiz (1807-
1873), Charles Robert Darwin (1809-1882),
Pierre-Joseph van Beneden (1809-1894), and
Johann Friedrich Theodor Muller (Fritz
Muller) (1822-1897). Moreover, the coast-
line of Brazil, ninth longest in the world
(World Resources Institute, 2012), was
traversed by some of the great oceanograph-
ic expeditions of the 19th and early 20th
centuries.

This paper provides a brief historical
overview of several of these expeditions,
focusing on those that included collections
of cnidarians and especially hydroids. Speci-
mens are deposited in several museums
around the world, principally in Europe and
the United States. Among these are the
Naturhistoriska Riksmuseet in Stockholm,
Sweden (Schwedischen Reichsmuseum, after
Jaderholm, 1903); the Rijksmuseum van
Natuurlijke Historic in Leiden (now the
Naturalis Biodiversity Center) and the Zoo-
logical Museum at Amsterdam, both in The
Netherlands (Vervoort, 1946); and the Royal
Scottish Museum (now the National Museum
of Scotland) in Edinburgh, Scotland (Ritchie,
1909; Rees and Thursfield, 1965). In the
United States, the most important institutions
holding material from Brazil are the National
Museum of Natural History (Smithsonian
Institution) in Washington D.C.; the Yale
Peabody Museum of Natural History in New
Haven, Connecticut; and the Museum of
Comparative Zoology at Harvard University
in Cambridge, Massachusetts.

Knowledge of the hydrozoan fauna of
Brazil was advanced only modestly by those
expeditions, yet accounts of expedition

materials by foreign workers (e.g., Allman,
1883, 1888; Nutting, 1900, 1904; Jaderholm,
1903; Vervoort, 1946) constituted much of
what was known about the fauna of the
country until the mid- 1940s (Grohmann et
a/., 2011). Significant progress has been
achieved since then through the work of
many Brazilian hydroid specialists, begin-
ning with the publications of Marta Van-
nucci (1921-) of the University of Sao Paulo.
Vannucci served at the Instituto Oceano-
grafico from 1946 to 1969 and left the
Institute to join UNESCO in 1969 (Schu-
maher and Vital Brasil, 2000).

MATERIALS AND METHODS

Criteria for choosing which expeditions to
highlight in this work were:

1. presence of hydroids collected on the
coast of Brazil based on data from
available reports;

2. records of hydroid material from expedi-
tions in online catalogs of several muse-
ums in Europe and in the United States,
such as those of the Swedish Museum of
Natural History (Stockholm, Sweden);
the Natural History Museum (London,
U.K.); the University Museum of Zoolo-
gy (Cambridge, U.K.); the National
Museum of Natural History-Smithsonian
Institution (Washington D.C., U.S.A.);
the Museum of Comparative Zoology of
Harvard University (Cambridge, Massa-
chusetts, U.S.A.); and the Yale Peabody
Museum of Natural History (New Haven,
Connecticut, U.S.A.). Thus, data were
extracted from both expedition publica-
tions and from information available via
the Internet on material deposited in
museum collections.

Information presented here is arranged in
the same order for each expedition: route of
the voyages (with map, when available) and a

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HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

3

brief history, names of the commanders of
the vessels, names of the naturalists, localities
traversed in Brazil, hydrozoan taxa collected
(as contained in contemporary reports or in
label data surveyed in museums), and collec-
tion localities. To better present the infor-
mation and enable comparison, Appendix
Table I provides (where possible) data for the
date and station of collection, coordinates,
depth, museum where material is deposited,
number of the sample in the catalog of the
institution, and source or sources where this
information was obtained. When more than
one source is cited, they are complementary
(one contains data for species/station, an-
other cites data for coordinates, and so
forth). Appendix Table II contains updated
names of species and some of the sources
where synonymies were obtained.

The Hartt Expedition has been treated
separately here because it comprised a series
of explorations that extended inland as well
as along the coast of Brazil.

Particularly important were reports of the
various voyages made by the key researcher
or researchers themselves, such as the narra-
tive of the Thayer Expedition by Agassiz and
Agassiz (1868). Another was the account of
Agassiz and Hartt (1870), containing one of
the most complete studies on geology and
physical geography of Brazil. The latter
publication, also containing illustrations
and maps, was largely the work of the
second author. When no information was
available in the published literature, addi-
tional data were obtained via the Internet.

At the end of each expedition report,
references are cited where additional infor-
mation about each of them can be obtained.

Based on the existing published plates,
comparisons were also made among species
collected in the 19th and early 20th centu-
ries with lists of species recorded from
Brazil in current literature. Sources of
information are listed in the right column

of Appendix Table II. All species names
were checked on the World Register of
Marine Species (WoRMS), and a justifica-
tion is given in the respective discussion
about the presence or absence of this
acronym on the list.

ACRONYMS AND CODES USED IN
THIS PAPER

BMNH

British Museum (Natural
History), now Natural History
Museum, London

GBIF

Global Biodiversity Information
Facility, Copenhagen, Denmark

KU

Kansas University

MCZ

Museum of Comparative
Zoology, Harvard University

NMNH

National Museum of Natural
History (Smithsonian

Institution)

NZ

Nomenclator Zoologicus

OEB

Organismic and Evolutionary
Biology (Harvard University)

RSGS

Royal Scottish Geographical
Society

SNAE

Scottish National Antarctic
Expedition

UFRJ

Universidade Federal do Rio de
Janeiro

worms

World Register of Marine
Species

YPM

Yale Peabody Museum of
Natural History, Yale

University

Exp.

Expedition

#

collecting station number

Nr

number

RESULTS AND DISCUSSION

/' Expedition de La Coquille (1822-1825)
Figure 1

History. Ordered by King Louis XVIII of
France (1755-1824) and carried out aboard

4

BREVIORA

No. 550

Figure 1. Route of VExpedition de La Coquille. Alienor.org (1998), redrawn with permission.

the corvette of the same name. La Coquille ,
the objective of this expedition was to
explore the “southern seas” and what was
then called Australasia. Under the command
of Lieutenant Louis Isidore Duperrey ( 1 786—
1865), with Lieutenant Jules Sebastien Cesar
Dumont d’Urville (1790-1842) as second in
command. La Coquille left Toulon on 1 1
August 1822 and entered port in Brazil on 16
October of the same year at Santa Catarina.
Its naturalist was Rene Primevere Lesson
(1794 1849), and the locality sampled was
Ilha de Santa Catarina (16 Oct 1822). The
hydrozoan collected was the medusa Eudora
rosacea Lesson, 1830.

Discussion. There is no figure of E. rosacea
in the plate published by Lesson (1830),
although illustrations are provided of two
other species (E. discoides and E. hydropotes )
that the author assigned to the same genus
(Fig. 2). Both closely resemble E. rosacea ,
which he described as . . completement
hyaline, assez epaisse au milieu, bordee sur sa
cir conference de une membrane mince ...”
(Lesson, 1830, chap. XIV, pi. 9, figs 2, 3).
Lesson was not a specialist in Cnidaria: he

was a medical doctor. Director of the
Botanical Garden of Rochefort, and a
representative of the Museum of Paris. He
may well have confused a medusa of the
family Aequoreidae, considered a group of
“Scyphozoa Discophorae” at the time (Long,
1841:121), for Eudora. In likely being unfa-
miliar with the species Aequorea forskalea
Peron and Lesueur, 1810, the name E. rosacea
was given to the specimen he encountered.
The aequoreid species A. forskalea (Hydro-
zoa, Leptothecata) is relatively common off
the Brazilian coast. These medusae readily
lose their tentacles during collection or when
reaching the beach, thereby resembling Les-
son's account of the species. Moreover, when
examining the bell or plate of the medusa in
lateral view (Fig. 2A), specimens of Eudora
(Scyphozoa, Discophorae) are much more
flattened than those of Aequorea (Fig. 2C).
The author may have been confounded by
these confusing details. From the diagrams by
Lesson (1830) of E. discoides and E. hydro-
potes (Fig. 2B), one can compare his concept
of Eudora (Fig. 2A) with Aequorea (Fig. 2C)
(both by Long, 1841).

2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

5

Figure 2. A. Specimen of the genus Eudora seen from the aboral side, in profile, and from the oral side; B. E.
hvdropotes (the two smaller specimens at top) and E. discoides (the large one at bottom); C. specimens of Aec/uorea
(one whole specimen and one in oral-aboral section through the mouth). Note the mesogleal thickening in the apical
part, resembling a delicate bell jar. Panels A and C after Long (1841:121, vol. XIX); panel B after Lesson (1830, chap.
XIV, pi. 9, figs 2 and 3).

As for the name Eudora , it has been applied
to genera in several taxonomic groups other
than Cnidaria (GBIF, 2004; NZ?2004; C. D.
C. Oliveira, personal communication), in-
cluding polychaetes ( Eudora Rafinesque,
1815); mollusks {Eudora Gray, 1852); crusta-
ceans {Eudora de Haan in Siebold, 1833, for
Decapoda; Eudora Bate, 1856, for Cumacea),
and insects {Eudora de Laporte in Brulle,
1840, for Coleoptera; Eudora Robineau-
Desvoidy, 1863, for Diptera). Although
Eudora Peron and Lesueur, 1810, is the senior
homonym, the generic name is no longer used
in Cnidaria (WoRMS).

For more detail, see Duperrey (1826),
Lesson (1830), Souza Sobrinho (1972), and
Morgat (2005).

Beagle Expedition (1831-1836)

Figure 3

History. The Voyage of the Beagle (1831-
1836) under the British flag is one of the
most widely known expeditions to pass
through Brazil. Under the command of
Captain Robert FitzRoy (1805-1865), the

Beagle departed Plymouth, southwest Eng-
land, on 27 December 1831. Charles Robert
Darwin (1809-1882), the naturalist on
board, was the first to write about the
sandstone and coral reefs along the coast of
Pernambuco state in the northeast. Lasting
nearly 5 years, this expedition passed
through Brazilian waters on two occasions.
On the outward leg of the voyage, the ship
sailed southwestwards along the coast of
Brazil towards Rio de Janeiro. En route, it
passed the Arquipelago de Sao Pedro e Sao
Paulo (St. Peter and St. Paul Rocks) (16 Feb
1832), Fernando de Noronha (20 Feb 1832),
Pernambuco, Bahia (29 Feb 1832), and
Abrolhos (18 Mar 1832), arriving in Rio de
Janeiro on 4 April 1832. Members of the
crew collected colonies of the hydroid family
Tubulariidae while in port. On the return leg,
Beagle traversed waters off Bahia (1 Aug
1836) and Pernambuco (12 Aug 1836) in the
northeast.

Discussion. Harmer (1901) compiled a
photographic report of manuscripts and data
taken from the catalog of specimens orga-

6

BREVIORA

No. 550

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Figure 3. Route of the Beagle Expedition. Mayr (1993, map between pages 80 and 81), reproduced with
permission.

nized by Darwin during the expedition.
Darwin’s catalog documented materials col-
lected at various localities and deposited in
collections. It is known, for example, that a
sample collected in Rio de Janeiro during
June 1832 (referred to as Nr 282), contains
colonies of the hydroid families Sertulariidae
and Tubulariidae. Another (from sample Nr
297) contains a fertile specimen of Sertular-
iidae, showing gonangia. In zoological notes
and listings of specimens, Keynes (2000)
alluded to a third sample containing Sertu-
lariidae (from sample Nr 265) that was
discarded by Harmer because of its extensive
state of decomposition. As for material
observed in Rio de Janeiro (4 Jul 1832),
Darwin (1842) reported on two species of
hydroids (family Tubulariidae) observed in
abundance on the ship's anchor after a month
in port and was struck by the rapidity of their
growth. Unfortunately, the specimens were

not preserved in alcohol, as was the custom of
the period (recorded in the annotations of
Keynes, 2000, as “. . . not in spirits”). Today,
hydroids are known to be among the first
invertebrates to establish themselves on
bacterial films in the process known as
bioencrustation (Sentz-Braconnot, 1966).

For more detail, see Darwin (1871),
Keynes (2000), and Rookmaaker (2009).

United States Exploring Expedition
(1838-1842)

Figure 4

History. On 18 August 1838, the United
States Exploring Expedition, composed of
six vessels (the sailing ships U.S.S. Vincennes ,
U.S.S. Peacock , U.S.S. Porpoise , U.S.S.
Relief \ U.S.S. Flying Fish , and U.S.S. Sea
Gull) set sail from Norfolk, Virginia, U.S.A.
The objectives of this expedition were as
much commercial as scientific, with a view

2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

7

Figure 4. Route of the United States Exploring
Expedition. Philbrick (2003:xx-xxi, Preface), partially
reproduced with permission.

toward expanding American influence and
interests in the eastern North Pacific. Com-
manded by Lieutenant Charles Wilkes
(1798-1877), expedition naturalists included
James Dwight Dana (1813-1895), Charles
Pickering (1805-1878) and Joseph Pitty
Couthouy (1808-1864). While in Brazilian
waters, the expedition collected at Cabo
Frio, northeastern Rio de Janeiro, on 22
November 1838 and in the city of Rio de
Janeiro on the following day.

Specimens identified by Dana from the
United States Exploring Expedition were
deposited in collections within the U.S.A.,
including at Peale’s Museum, the Cabinet of
the Academy of Natural Sciences (now the
Academy of Natural Sciences of Drexel

University in Philadelphia), and the Boston
Society of Natural History (now the Muse-
um of Science, Boston) (Dana, 1846). Only
one species of hydroid was collected, de-
scribed as Tubularia ornata Couthouy, 1846
(Fig. 5).

Discussion. The physiography of many
localities mentioned in the bibliography as
points of collection during some expeditions
has changed over time. This is the case, for
example, for collections made by the United
States Exploring Expedition in Guanabara
Bay, Rio de Janeiro, in November 1838
(Wilkes, 1852). The cove of Praia Grande,
where some of the material was collected, is
today the section of shore from Armagao
Hill to Gragoata Hill in Niteroi and is now a
level land-filled area. Only one species of
hydroid, described as Tubularia ornata Cou-
thouy, 1846, was collected (in Dana 1846:19,
vol. 7, chap. 3, fig. 1). That name of the
species (Fig. 5) is no longer considered valid
(WoRMS). However, Ectopleura crocea (L.
Agassiz, 1862) is relatively common in
Guanabara Bay (Absalao, 1993; Omena et
al., 1995) where vessels of the United States
Exploring Expedition anchored on 23 No-
vember 1838. Possibly, one of the tubulariids
collected some years before by the Beagle
Expedition (1832) (see discussion of the
Beagle Expedition above) and reported by
Darwin (1842) was E. crocea.

See details in Dana (1846, 1849), Jenkins
(1850), Wilkes (1852), Tyler (1968), and
Philbrick (2003).

Eugenie Expedition (1851-1853)

Figure 6

History. At the direction of the Swedish
government, the Eugenie Expedition left
Stockholm on 24 September 1851 and came
close to circumnavigating the globe. Under
the command of Captain Christian Adolf
Virgin (1797-1870), and with naturalist

8

BREVIORA

No. 550

Figure 5. Tubularia ornata Couthouy, 1846. A.
Whole polyp; B. detail of a gonophore. Based on
Dana (1846:19, vol. VII, chap. Ill, fig. 1).

Johan Gustav Hjalmar Kinberg (1820-1908)
on board, the frigate Eugenie sailed from
Stockholm in 1851 visiting Madeira, Rio de
Janeiro, Buenos Aires, the Magellan Straits,
Chile, Peru, Panama, the Galapagos Islands,
the Sandwich Islands, San Francisco, Tahiti,
Sydney, Canton, Manila, Batavia, and the
Cape of Good Hope. The frigate was at

Cabo Frio, Rio de Janeiro, in December
1851 en route to Rio Grande do Sul (off the
Plate River estuary) and the Strait of
Magellan. At Cabo Frio, the expedition
collected the hydroids Sertularella tenella
(Alder, 1856); Obelia bidentata Clark, 1875;
and Lafoea cylindrica von Lendenfeld, 1885.

Discussion. The three hydroid species
mentioned above were cited by Jaderholm
(1903) as existing in the Rijksmuseum van
Natuurlijke Historie in Leiden. However, the
online database of the Smithsonian National
Museum of Natural History lists a sample of
Sertularella tenella Alder, 1856 (accession
USNM 70696), with the same data as
material at the museum in Leiden. It is
possible that part of the collected sample was
sent to the Smithsonian, or this may be a
case of exchange of specimens between
institutions. In the appendix of a paper on
hydroids of the coast of Sao Sebastiao,
Brazil, Migotto (1996) questionably referred
the hydroid identified as S. tenella by
Jaderholm (1903) to Sertularella conica All-
man, 1877. Later, Migotto et al. (2002)
recorded S. conica from Rio Grande do Sul
and from Sao Paulo to Espirito Santo states.

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Figure 6. Route of the Eugenie Expedition. Skogman (1854-55).

2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

9

while making no mention of the name S.
tenella. Meanwhile, S. tenella has been
reported from Bahia (Grohmann et al.,
2003) and from Rio de Janeiro (Grohmann
et al., 2011). Both S. tenella and S. conica are
treated as valid species in WoRMS. Galea
(2013:22) maintained that the hydroid iden-
tified as S. cornea from South Carolina,
U.S.A. (Calder, 1983), and Bermuda (Cald-
er, 1991, 1993, 1998, 2000, 2013) was a new
species that he renamed Sertularella calderi.
However he, himself, regarded records of S.
conica from Brazil as likely referable to
Sertularella peculiaris Leloup, 1974. As for
the hydroid identified as Lafoea cylindrica
von Lendenfeld, 1885, Migotto (1996) con-
sidered it likely identical to Hebella scandens
(Bale, 1888) and different from the true L.
cylindrica ( =Hebellopsis cylindrica).

For more detail, see Skogman (1854-55),
and Jaderholm (1903).

Thayer Expedition (1865-1866)

Figure 7

History. The Thayer Expedition was spon-
sored by American financier and philanthro-
pist Nathaniel Thayer, Jr. (1808-1883), and
organized under the leadership of Louis
Agassiz of Harvard College. The expedition
to South America departed Boston, Massa-
chusetts, aboard the steamship Colorado on
2 April 1865 and arrived in Rio de Janeiro
on the 23rd of that month. Agassiz had
proposed the expedition in part to recuperate
from health problems and in part because of
his fascination with the diversity of the fauna
of Brazil after studying a collection of fish at
the request of K. F. P. von Martius.
Moreover, Agassiz wished to prove a theory
of continental glaciation — a project later
dropped — within Brazilian territory.

Under the command of Captain George
Bradbury (?-?) and with the participation of
other naturalists, including Joel Asaph Allen

(1838-1921) and Charles Frederick Hartt
(1840 1878), the expedition collected in
several states during 1865, including Rio de
Janeiro (23 Apr), Espirito Santo (no exact
date), Bahia (28 Jul), Alagoas (30 Jul),
Pernambuco (31 Jul), Paraiba (2 Aug), Ceara
(5 Aug), Maranhao (6 Aug), and Para (10
Aug), and also passed through other inland
states. This expedition collected nearly all of
the species of the hydrozoan genus Mdlepora
Linnaeus, 1758, that are known from Brazil:
Millepora sp. (Maceio, Alagoas); Mdlepora
insignis Verrill, 1864 (Guarapari, Espirito
Santo, and Porto Seguro, Bahia); Millepora
alcicornis var. fenestra ta Linnaeus, 1758
(Porto Seguro, Bahia); Millepora nitida
Verrill, 1868 (Porto Seguro, Bahia); and
Millepora boletus Pourtales, with no year or
locality specified on the labels in the MCZ.
Some minor inconsistencies in localities
where the material was collected exist in the
data given by Verrill (1902:197). In that
work, he recognized three varieties of M.
alcicornis'. var. cellulosa Verrill (Rio Formo-
so, Pernambuco; Paraiba), var. digitata
?Esper (Rio de Janeiro, Cabo Frio; Porto
Seguro and Abrolhos, Bahia; Maceio, Ala-
goas), and var. fenestrata Duchassaing and
Michelotti (Abrolhos, Bahia). Verrill never-
theless suggested that when carefully studied,
changes or additions might be necessary to
the varieties he had recognized.

Mention should also be made that one of
the participants in the expedition was Wil-
liam James (1842 1910). He tired of natural
history collecting during the trip, eventually
turned to philosophy and psychology in-
stead, and became widely known later as
“the father of American psychology.”

Discussion. One of Louis Agassiz’s objec-
tives of the Thayer Expedition was to seek
evidence that would refute the transmutation
of species theory promulgated by Charles
Darwin. A creationist, Agassiz’s intent was
to prove that all land life would have been

10

BREVIORA

No. 550

Figure 7. Stations covered by the Thayer Expedition in the Amazon and in the southeast (according to Higuchi,
1996). Source: MCZ. Museum of Comparative Zoology, Harvard University (http://mcz.harvard.edu/Departments/
Ichthyology/expeditions_thayer_hassler.html), redrawn with permission.

destroyed by Pleistocene glaciation at sea
level in the tropics. He therefore believed
that the actual existence of such life could
have been possible only through Divine
creation. Charles Frederick Hartt, one of
the expedition geologists, was skeptical
about what was held to be glacial drift, a
key to Agassiz’s hypothesis. Hartt eventually
concluded, publicly, that the supposed drift
was simply material from the process of
intense bedrock weathering, thereby refuting
the ideas of the famous expedition leader.

This expedition without doubt collected
the most hydrocorals on the coast of Brazil.
The MCZ holds specimens collected by the
expedition that have been identified as
Millepora alcicornis, M. insignis , M. nitida ,
and M. boletus. In his report on milleporids
from Brazil, Boschma (1962) reported Mil-
lepora braziliensis Verrill, 1868, Millepora

squarrosa Lamarck, 1816, M. nitida , and M.
alcicornis. According to WoRMS, M. insig-
nis has been relegated to the synonymy of
Millepora platyphylla Hemprich and Ehren-
berg, 1834. Millepora boletus, a species
attributed to Pourtales, is not cited in recent
lists of cnidarians from Brazil and was not
found in any of the synonymy lists of other
species. It is not listed in WoRMS and
appears never to have been formally de-
scribed and named. Support for this hypoth-
esis is based on the fact that no description
of it could be found in any bibliography.
According to Hartt (in Agassiz and Hartt,
1870), Verrill distinguished three forms of
the genus Millepora Linnaeus, 1758, along
the Brazilian coast: M. alcicornis Linnaeus,
1758; M. braziliensis Verrill, 1868; and M.
nitida Verrill, 1868. More recently, Amaral et
al. (2002) recognized four distinct species in

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HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

Brazil: M. alcicornis, M. braziliensis, M.
nitida , and an undescribed species referred
to as Millepora sp. a Amaral et al., 2002. In
that article, the authors suggest that the
Caribbean species M. squarrosa is distinct
from M. braziliensis. Amaral et al. (2008)
subsequently published an overview of Bra-
zilian Milleporidae, finally describing and
naming the new species as Millepora labored
Amaral, 2008. It is possible that this species
corresponds to M. boletus , the undescribed
species attributed to Pourtales (see above).

See details in Agassiz and Agassiz (1868),
Agassiz and Hartt (1870), Dick (1977), and
Higuchi (1996).

Challenger Expedition (1873-1876)
Figure 8

History. The renowned Challenger Expe-
dition of 1873-1876, conceived by two great
British biologists, William Benjamin Car-
penter (1813-1885) and Sir Charles Wyville
Thomson (1830-1882), was financed by the
British Treasury. The prime objective was to
prove that life existed in the deep sea,
notwithstanding the great pressures, the
cold, and the lack of light, although inves-
tigations were also undertaken on the
chemistry, geology, and physics of the open
ocean. To carry out the mission, the corvette
H.M.S. Challenger was selected. Modifica-
tions were made in its structure to facilitate
the work of the scientists, including con-
struction of a chemistry laboratory on
board. The expedition was commanded by
Captains Sir George Strong Nares (1831 —
1915) and Frank Tourle Thomson ( 1 829—
1884). The team of scientists included Sir
Charles Wyville Thomson himself as well as
John Murray (1841-1914), John Young
Buchanan (1844-1925), Henry Nottidge
Moseley (1844-1891), and Rudolf von Wil-
lemoes-Suhm (1847-1875). The expedition
passed through Brazil in 1873, its route

1 1

passing the Arquipelago de Sao Pedro e
Sao Paulo (28 Aug), Fernando de Noronha
(2 Sep), Pernambuco (9 Sep), Alagoas (11
Sep), Sergipe (12 Sep), and Bahia (14 Sep).
Hydroids (other than hydrocorals) of the
expedition were examined by George James
Allman (1812-1898), a leading 19th century
specialist of the group (Calder, 2015). Of all
expeditions reviewed here, the largest collec-
tion of hydroids from Brazil was obtained by
naturalists of the Challenger.

Specimens were deposited in the British
Museum (Natural History) (now the Natural
History Museum, London) (Allman, 1883,
1888). Among the species identified were
Ag/aophenia calamus Allman, 1883; Campa-
nularia ptychocyathus Allman, 1888; Desmo-
scyphus acanthocarpus Allman, 1888;
Desmoscyphus obliquus Allman, 1888; Des-
moeyphus pectinatus Allman, 1888; Idia
pristis Lamouroux, 1816; Lytocarpus race-
miferus Allman, 1883; Sertularia cylindrithe-
ca Allman, 1888; Sertularia integritheca
Allman, 1888; and Thyroscyphus ramosus
Allman, 1877, all in Bahia; Halicornaria
plumosa Allman, 1883, and Stylaster duch-
assaingi Pourtales, 1867, at Barra Grande,
Alagoas; and Thuiaria hyalina Allman, 1888,
in Sergipe.

Discussion. Names of most of the species
mentioned in all expeditions treated here
have changed since their reports were
published in the late 19th and early 20th
centuries. Ag/aophenia calamus Allman,
1883, was treated by Migotto (1996) as a
species of doubtful status. Allman (1883:39)
had remarked that it differed little from
typical aglaopheniids. Meanwhile, Vervoort
and Watson (2003:261) considered A. cala-
mus to be conspecific with Ag/aophenia
pluma (Linnaeus, 1758). For now, A. cala-
mus is listed in the WoRMS database as a
valid species. Not listed in WoRMS is
Campanularia ptychocyathus Allman, 1888,
considered synonymous with Clytia nolifor-

12

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No. 550

Figure 8. Route of the Challenger Expedition. Perry and Fautin (2003), redrawn with permission.

mis (McCrady, 1859) by Migotto (1996) but
not by Calder (1991:68, 2015:236). As for
the species that Allman (1888) assigned to
the genus Desmoscyphus, D. acanthocarpus
is referable to Diphasia digitalis (Busk,
1852), D. pectinatus is referable to Tridentci-
ta marginata (Kirchenpauer, 1864), al-
though WoRMS considers this species
Sertularia marginata (Kirchenpauer, 1864).
Desmoscyphus obliquus is likely referable to
the same species. Idia pristis Lamouroux,
1816; Lytocarpus racemiferus Allman, 1883;
and Sertularia cylindritheca Allman, 1888,
are now assigned to different genera, being
known respectively as Idiellana pristis (La-
mouroux, 1816), Macrorhynchia racemifera
(Allman, 1883), and Sertularelloides cylin-
dritheca (Allman, 1888) (Migotto, 1996;
Migotto et al., 2002; Bouillon et al., 2004;
Galea, 2013). Allman (1888:85-87) consid-
ered the diagnosis of Idiellana pristis by
Lamouroux to be “. . . short and inade-
quate,” so a new description was provided.
As for Sertularia integritheca Allman, 1888,
it was included in the synonymy of Sertular-
ella formosa Fewkes, 1881, by Migotto

(1996), then referred to the genus Synthe-
cium Allman, 1872, as Synthecium formosum
(Fewkes, 1881) by Migotto et al. (2002) and
is now assigned to the genus Hincksella
Billard, 1918, as Hincksella formosa (Few-
kes, 1881) (see WoRMS). The name Hal-
icornaria plumosa Allman, 1883, is an
invalid junior homonym of Halicornaria
plumosa Armstrong, 1879, and the name
Gymnangium allmanii (Marktanner-Turner-
etscher, 1890) has been applied to the species
(see WoRMS, although it is currently
misspelled therein as G. allmani ). Finally,
the name Thuiaria hyalina Allman, 1888, has
long been considered a synonym of Sertu-
larella diaphana Allman, 1885 (see Migotto,
1996).

According to Cairns (1986:2, 73), a record
of the stylasterid Stylaster duchassaingi
collected at station 122 at a depth of 732 m
is referred to as “. . . one dubious record,
strongly queried.”

See details in Thomson and Murray
(1885), Allman (1883, 1888), and Rehbock
(1992).

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HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

13

U.S. Fish Commission Steamer Albatross
Expedition (1887 1888)

History. Constructed in 1882, the Alba-
tross was the first ship designed and built
especially for marine research. Equipped
with on-board laboratories, deep-sea dredg-
ing gear, and hydrographic instruments, it
was the first ship belonging to the American
government to have electric lighting in-
stalled. During four decades of service, it
sailed across the North and South Atlantic,
North and South Pacific, and Indian oceans.
Its expeditions resulted in huge museum
collections and a massive number of scien-
tific publications and is the oceanographic
vessel to which is attributed the largest
number of newly described marine species.
During the 1877-1878 expedition, which
sailed around the southern tip of South
America to California, its commander was
Captain Zera Luther Tanner (1835-1906),
and the naturalist in charge was Leslie
Alexander Lee (1852-1908). Localities sam-
pled by the Albatross Expedition in Brazil
during 1887 included Ceara (14 Dec), Para-
iba (16 Dec), Bahia (18 Dec), Abrolhos (27
Dec), and Cabo Frio, Rio de Janeiro (30
Dec). Collecting efforts for cnidarians con-
centrated largely on Anthozoa (Octocorallia
and Hexacorallia), and only one hydrozoan,
identified as Thyroscyphus sp., taken at Joao
Pessoa, Paraiba, is recorded.

For more detail, see Allard (1999).

Scottish National ( Scotia ) Antarctic
Expedition— SNAE (1902-1904)
Figure 9

History. Organized by naturalist William
Speirs Bruce (1867-1921), this expedition left
Troon, Scotland, on 2 November 1902. On
board the ship S.Y. Scotia, commanded by
Captain Thomas Robertson (1855-1918),
the expedition objective was to increase
knowledge of the Antarctic region. The

SNAE succeeded in building the first mete-
orological station, named Omond House, on
the Antarctic continent. During its passage
off Pernambuco, it was the first to take
photographs of the Arquipelago de Sao
Pedro e Sao Paulo.

Brazilian hydroid material collected by
this expedition was limited to a single station
(#81), sampled at Abrolhos, Bahia (20 Dec).
The collection, deposited at the Royal
Scottish Museum (now the National Muse-
um of Scotland, Edinburgh), was examined
and identified by James Ritchie (1882-1958).
Taxa collected included Aglaophenia allmani
Nutting, 1900; Aglaophenia dubia Nutting,
1900; Aglaophenia minima Nutting, 1900;
Halicornaria longicauda Nutting, 1900;
Monostaechas quadridens (McCrady, 1859),
Sertularici cornicina (McCrady, 1859), Sertu-
laria heterodonta Ritchie, 1909; and Sertu-
laria rathbuni Nutting, 1904 (Ritchie, 1909).

Discussion. Again, changes have subse-
quently been made in the names of some of
these species. Aglaophenia allmani is now
referred to the genus Macrorhynchia Kirch-
enpauer, 1872, as Macrorhynchia allmani
(Nutting, 1900) (see Migotto, 1996; Migotto
et al., 2002; WoRMS). Aglaophenia minima
is a synonym of Aglaophenia latecarinata
Allman, 1877 (WoRMS). The name Hal-
icornaria longicauda is currently assigned to
the genus Gymnangium Hincks, 1874, and
taken to be a synonym of G. allmani
(Marktanner-Turneretscher, 1890). The spe-
cies has recently been redescribed by Galea
(2013). Sertularia cornicina is usually regard-
ed as identical with Dynamena disticha (Bose,
1802) (see Migotto, 1996). Sertularia hetero-
donta is conspecific with Tridentata distans
(Lamouroux, 1816); a syntype of the species
exists in the BMNH, with a photograph
from the catalog in the online database of
the collection. Sertularia rathbuni is a syno-
nym of Dynamena dalmasi (Versluys, 1899)

14

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No. 550

Figure i Traci chan of ih t Scotia 1902-1904

Figure 9. Route of the Scottish National Antarctic
Expedition. Speak (1992), reproduced with permission.

(see Migotto, 1996; Migotto et al. , 2002;
WoRMS).

See details in Ritchie (1907, 1909), and
Speak (1992).

British Antarctic ( Terra Nova) Expedition
(1910-1913)

History. On 1 June 1910, this expedition
left London for Wales, setting out from
Cardiff on 15 June aboard S.S. Terra Nova.
The prime goal of expedition leader Robert
Falcon Scott (1868-1912) was to be the first
to reach the South Pole. Associates Edward
Adrian Wilson (1872-1912), Denis Gas-
coigne Lillie (1888-1963), and Edward Wil-
liam Nelson (1883-1923) were given
responsibility for scientific aspects of the
expedition. Scott soon discovered that he
would have to contend with noted Norwe-
gian explorer Roald Engelbregt Gravning

Amundsen (1872-1928) to be first to reach
the pole. Off Brazil, en route to Antarctica,
Terra Nova cruised along the coast off Rio
de Janeiro and the Trindade and Martim
Vaz archipelago.

The misfortune of the expedition at its
final destination is well known. After suc-
cessfully reaching their intended destination
in Antarctica, Scott and four companions
discovered to their great dismay that
Amundsen had reached the South Pole a
month earlier. Tragically, all five died during
their attempt to return to the expedition’s
base.

An archive of photographs from this
expedition is deposited at the Scott Polar
Research Institute of Cambridge University
in England.

The localities sampled in Brazil were Ilha
da Trindade (26-30 Jul 1910), Espirito
Santo, and Rio das Ostras (2 May 1913),
Rio de Janeiro. However, the species Liriope
tetraphylla (Chamisso and Eysenhardt,
1821), collected by the Terra Nova and
deposited in the BMNH, is without exact
locality data.

Discussion. A majority of the cnidarian
specimens collected by the British Antarctic
Expedition were sea anemones (Anthozoa).
Data for the hydromedusa L. tetraphylla
collected during the expedition were ob-
tained through searches made in the BMNH
(under accession 1970.10.8.76-78 and
1970.10.8.89-100), although this information
is absent from the listing by Totton (1930).

See details in Jones (2005).

Hartt Expedition (1875-1877)

History. As a result of his participation in
the Thayer Expedition, geologist Charles
Frederick Hartt became engrossed with
Brazil and its geology. As a youth growing
up in Nova Scotia, Canada, he had learned
Portuguese from an immigrant shoemaker,

2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

15

facilitating his work in the South American
country. He returned in 1867 to undertake
investigations on geology of the Bahia
region and its coral reefs. Thereafter, Brazil
became the focus of his research, and he
returned to the country several times while
on the faculty of Cornell University, Ithaca,
New York, U.S.A. Hartt organized a series
of voyages and traversed the coast of several
Brazilian states in what became known as
the Hartt Expedition. Recognizing the great
need of a geological survey of the country,
Hartt was instrumental in the founding, in
April 1875, of the Comissdo Geoldgica do
Imperio do Brasil , with its headquarters in
the city of Rio de Janeiro, and he was named
its director. From the state of Rio de
Janeiro, where he temporarily settled with
his family (from 1875 to 1877, according to
Brice and Figueiroa, 2003:19), Hartt and his
team of researchers carried out geological
explorations across the rest of the country.
His contributions to geology in Brazil were
carried on by noteworthy students he had
influenced to undertake investigations there,
such as John Casper Branner (1850-1922),
Orville Adalbert Derby (1851-1915), and
Richard Rathbun (1852-1918) (Menezes,
1878; Brice and Figueiroa, 2003). His team
was also composed of a few Brazilian
collaborators, such as the engineers Elias
Fausto Pacheco Jordao (1849-1901) and
Francisco Jose de Freitas (?-?), and the
photographer Marc Ferrez (1843-1923).
With his colleagues, Hartt explored the
country from north to south (Amazonas to
Rio Grande do Sul), dedicating at least 12 of
his 38 years of life to studies of Brazilian
natural history. Given his professional back-
ground, his interests in Brazil mainly in-
volved the geology, paleontology, arche-
ology, and physical geography of the coun-
try. Nevertheless, in the research proj-
ect presented to the Brazilian government,
12 items were to be undertaken. One of them

(the ninth article) was directed to the study
of marine animals — the stony corals — stat-
ing (when translated into English): “The
investigation of useful and harmful animals
to man, and the study of stony coral reefs to
determine their structure, their mode and
speed of development, their position, their
extent, and their potential obstruction of
ports and navigable channels (Menezes
1878: 36).”

As Hartt became ill in early 1877, work
began to progress more slowly. Nevertheless,
his collection by that time comprised ap-
proximately 500,000 specimens housed in an
improvised house in Rio de Janeiro. By late
1877, however, finances of the Imperial
Government were so dire that the ministry
had to restrict public funding, and cuts were
made to all nonessential spending. Seeing
little immediate economic benefit and few
published results, funding of the commission
was reduced and then, by January 1878,
finally cancelled. In early 1878, Hartt con-
tracted yellow fever and died in a rooming
house in Rio de Janeiro on 18 March at age
38, surrounded by members of his commis-
sion. A few months earlier, his family had
returned to the U.S.A.

The available bibliography on the Hartt
Expedition cites only the localities sampled
on the coast, without mentioning dates or
coordinates of the collections. The same is
true for labels consulted from the museums
where the material is deposited. The Hartt
team visited Rio de Janeiro, where the coral-
like anthoathecates Stv/aster roseus (Pallas,
1766) and Millepora sp. were collected. Also
visited were Pernambuco, at Maria Farinha,
Santo Aleixo, Rio Formoso, and Recife de
Candeias. Elsewhere, at Paraiba, Alagoas
(Maceio), and Bahia (Mar Grande, Itacara-
nha, and Periperi), most material collected
was of true corals (Anthozoa, Scleractinia).
Additional sampling was undertaken on the
continental shelf and at Abrolhos. In Peri-

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BREVIORA

No. 550

peri the expedition collected the aglaophe-
niid hydroid Lytocarpus philippinus Kirch-
enpauer, 1872 [=Macrorhynchia philippina
(Kirchenpauer, 1872)]. As the data on
biological material cited here were obtained
by checking museum labels [online databases
of the NMNH and YPM (see Appendix
Table I)], no formal discussion of the data is
needed.

For more detail, see Menezes (1878),
Freitas (2002), and Kern (201 1).

Summary of the Expeditions

With the passage of time, ongoing “prog-
ress” can be noted relative to the types of
vessels used on expeditions. The earliest
vessels were sailing ships, such as La
Cocjuille, the Beagle , and ships of the United
States Exploring Expedition. In succeeding
decades, steamships such as the Colorado ,
used on the Thayer Expedition, appeared.
Later, H.M.S. Challenger was specially
modified and equipped for dredging in deep
water, with a laboratory for the convenience
of its scientists. The Albatross was the first
ship designed and constructed especially for
the purpose of marine research: it possessed
a laboratory, deep-sea dredging gear, and
modern hydrographic instruments, and was
the first American ship with electric lighting
in its facilities. Onboard reports of the
expeditions also continually improved, with
the inclusion of new data (e.g., coordinates
and depth), as well as greater precision and
more defined objectives.

Materials obtained by these expeditions
constituted the first contributions to knowl-
edge of the “Radiata” (a polyphyletic group
recognized at the time that included Cnidaria,
Ctenophora, and Echinodermata) from Bra-
zil. Samples obtained during /’ Expedition de
La Coquille included four cnidarians (two
scyphozoans; one hydrozoan, thought at the
time to be a scyphozoan; one anthozoan).

Collections by the United States Exploring
Expedition yielded one hydrozoan and six
anthozoans (two octocorals and four hexa-
corals) in samples from Rio de Janeiro. Three
species of hydrozoans were recorded from
Rio de Janeiro by the Eugenie Expedition.
Reports from the Thayer Expedition docu-
mented the most species of milleporids (five
species), along with eight anthozoans. The
largest collection of cnidarians was obtained
by the Challenger Expedition, with 13 species
of hydrozoans and 15 of anthozoans.

Increasing richness of resulting collections
as years advanced appears to correlate at least
in part with improvements in the vessels and
their sampling gear. However, most of the
expeditions simply passed along the Brazilian
coast en route to other areas, and their
exploration objectives, sampling efforts, and
interests of participating naturalists varied.
As for hydroids, many of which are fragile
and easily overlooked, effectiveness of sample
sorting becomes an issue in their discovery.

Critical Evaluation

Discoveries of hydroids and medusae
during expeditions of the 19th and early
20th centuries through Brazilian waters
provided initial contributions to knowledge
of Hydrozoa in the country. These obscure
invertebrates were totally unknown in the
region at the time. Reports on them,
especially Allman’s (1883, 1888) accounts
of species from the Challenger Expedition,
provided a foundation for later taxonomic
studies. Moreover, discoveries of a number
of new species in Brazil, a country almost
totally uncharted and unexplored at the time
and covering a large geographic area, were of
significance overall to science.

ACKNOWLEDGMENTS

We are indebted to Ricardo S. Cardoso,
Leonardo S. Avilla, and Tatiana F. Maria

2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

17

(Universidade Federal do Estado do Rio de
Janeiro) and Cleo D. C. Oliveira (UFRJ) for
helpful suggestions. Adam J. Baldinger,
Curator of the Collection of Marine Inver-
tebrates (OEB, Harvard University), and
Aude Andouche (Departement Milieux et
Peuplements Aquatiques du Museum Na-
tional d’Histoire Naturelle de Paris) provid-
ed information on Brazilian material
deposited at their institutions. Heartfelt
thanks are extended to my friend Vera Abud
for critically reading this paper and Janet W.
Reid (Virginia Museum of Natural History)
for help with the first English version and to
Axel M. Katz (UFRJ), Carlos A. M.
Barboza (Nucleo em Ecologia e Desenvolvi-
mento Socio-Ambiental de Macae, UFRJ),
and Rafael B. de Moura (Universidade
Federal de Pernambuco) for their contribu-
tions to figures. We are especially indebted to
those who gave us permission to reproduce
and redraw the maps, such as Vincent
Lagardere (Alienor.org, Conseil des Musees,
La Coquille Expedition ), Harvard University
Permissions Department (HUP reference
38291, Beagle Expedition), Jeffrey Ward
Inc. (cartography and graphic design. United
States Exploring Expedition), Project Rune-
berg ( Eugenie Expedition), Karsten Hartel
and Melissa Aja (Museum of Comparative
Zoology, Harvard University, Department
of Ichthyology, Thayer Expedition). Daphne
Fautin (Kansas University, Challenger Ex-
pedition), and Alexa Martin (Royal Scottish
Geographical Society, Scottish National
Antarctic Expedition). We are also grateful
to the reviewers and the editorial staff of
Breviora , Jonathan Losos and Deborah
Smiley, for their excellent and constructive
comments.

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2016

HYDROZOA FROM EARLY EXPEDITIONS TO BRAZIL

21

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