MARINE AND FISHERY SCIENCES 35 (1): 115-122 (2022)

https://doi.org/10.47193/mafis.3512022010104

ABSTRACT. Digitally accessible Primary Biodiversity Data (PBD) are currently available

through a number of web-based platforms. This information is allowing for a growing number of

ecological, biodiversity informatics or conservation projects. Most of this information comes from

Natural History Collections (NHC) worldwide. Despite well-known limitations, NHC data are par-

ticularly useful as a source of data on invertebrates, which comprise about 99% of animal life.

However, a presumably very high amount of PBD is still not digitally accessible. Even the most

important scientific collections in developed countries are not fully inventoried or digitally accessi-

ble. Furthermore, species determination rates remain alarmingly low for some collections, and most

existing determinations probably should be retested. This is particularly true of expensive, difficult-

to-obtain deep-sea benthic samples. This paper reviews the database on material collected by the

German RV ‘Walther Herwig I and II’ during 1966-1978 research cruises to the Southwestern

Atlantic Ocean (SAO), stored in the invertebrate collection (invertebrates except Mollusca,

Arthropoda and Annelida) of the Zoological Museum of Hamburg. We found that out of 3,305

records, at least 204 species were identified, 72% are still undetermined at species level and 65% at

genus level. While sampling of more remote geographic areas and the use of a wider variety of sam-

pling methods are still necessary, supporting the training of an army of taxonomists will be of para-

mount importance to overcome the daunting task of analyzing the huge number of samples. stored

in museum collections. A community-wide effort is urgently needed to address this issue, and inter-

national cooperation must play a key role in this endeavor.

Key words: Oceanographic research, Southwestern Atlantic, benthic biodiversity, natural history,

collections.

Invertebrados bentónicos recolectados por el BI “Walther Herwig I y II” en el Océano

Atlántico Sudoccidental (1966-1978): una revisión de la colección de invertebrados del Museo

Zoológico de Hamburgo

RESUMEN. Actualmente, los datos primarios de biodiversidad (DPB) accesibles digitalmente

están disponibles a través de una serie de plataformas basadas en la web. Esta información ha posi-

bilitado la generación de un número creciente de proyectos ecológicos, de informática de la biodiver-

sidad o de conservación. La mayor parte de esta información proviene de las Colecciones de Historia

Natural (CHN) de todo el mundo. A pesar de sus conocidas limitaciones, los datos de las CHN son

particularmente útiles como fuente de datos sobre invertebrados, que comprenden alrededor del 99%

de la vida animal. Sin embargo una cantidad presuntamente muy alta de DPB todavía no es accesible

digitalmente. Incluso las colecciones científicas más importantes de los países desarrollados no se

115

*Correspondence:

alvar.carranza@gmail.com

Received: 16 August 2021

Accepted: 17 September 2021

ISSN 2683-7595 (print)

ISSN 2683-7951 (online)

https://ojs.inidep.edu.ar

Journal of the Instituto Nacional de

Investigación y Desarrollo Pesquero

(INIDEP)

This work is licensed under a Creative

Commons Attribution-

NonCommercial-ShareAlike 4.0

International License

Marine and

Fishery Sciences

MAFIS

NOTE

Benthic invertebrates collected by the RV ‘Walther Herwig I and II’ in the

Southwestern Atlantic Ocean (1966-1978): a review of the Zoological

Museum of Hamburg invertebrates collection

ALVAR CARRANZA1, 2, *, PABLO LIMONGI1and ANDREAS SCHMIDT-RHAESA3

1Departamento de Ecología y Gestión Ambiental, Centro Universitario Regional Este-Cure, Sede Maldonado, Maldonado, Uruguay. 2Área

Biodiversidad y Conservación, Museo Nacional de Historia Natural, C.C. 399, CP 11000, Montevideo, Uruguay. 3Zoologisches Museum,

Leibniz Institut für die Analyse des Biodiversitätswandels (LIB), Hamburg site Martin-Luther-King-Platz 3, 20146 - Hamburg, Germany.

ORCID Alvar Carranza https://orcid.org/0000-0003-3016-7955

Natural history collections (NHC) worldwide

harbour more than 3 billion of specimen records

(Pyke and Ehrlich 2010). Multiple collectors over

long periods of time have developed these ad hoc

data sets which contain invaluable information on

past and present biodiversity (Krishtalka and

Humphrey 2000). Over the last years, this huge

reservoir of information has been available since

computing and the exchange of data through net-

works allowed solving the problems of digitizing,

georeferencing and data distribution. Therefore, a

large amount of museum data is, and many more

will be, available to worldwide users.

The Southern Ocean has been the focus of

oceanographic research since the late 17th centu-

ry, although it is only since the early 1960s that

international research in the area has begun to

grow in importance. During this time, the scien-

tific and technical systems of Argentina, Brazil

and Uruguay were boosted by increasing connec-

tivity due to the establishment of research net-

works and the development of bilateral pro-

grammes. Joint scientific efforts included cruises

carried out by research vessels from Germany

(‘Walther Herwig I and II’, ‘Meteor’), USA

(‘Vema I’ and ‘Atlantis II’), Japan (‘Kaiyo Maru’,

‘Orient Maru’, ‘Shinkai Maru’), Russia (‘Evrika’,

‘Dimitry Stefanov’) and Poland (‘Professor

Siedlecki’), triggering the development of region-

al ocean sciences (Angelescu and Sánchez 1995).

In particular, bilateral research programmes in the

second half of the 20th century (e.g. Argentina-

Germany exploration programmes) included four

visits of the German RV Walther Herwig in 1966,

1968, 1970-1971 (Schmidt 1971) and 1978. The

main objective of these research programmes,

within the FAO Project of Fishery Development

framework, was the survey and assessment of

fishery resources of the Patagonian shelf (Ange-

lescu and Sánchez 1995).

During the ‘Walther Herwig I and II’ (hereafter

WH) cruises in the 1960s and 1970s, a wealth on

material was collected in hundreds of stations in

the Southwestern Atlantic Ocean (SAO). Most of

these benthic invertebrate samples are currently

stored in the Zoological Museum of Hamburg

(ZMH), whose nine research collections house

more than ten million of scientific objects. Estab-

lished in 1843 as the Hamburg Museum of Natural

History, the Museum was later renamed as the

State Institute of Zoology and then the ZMH,

before it was finally incorporated into Universität

Hamburg in 1969 (Köstering 2018). Recently, it

fused with the Zoological Research Museum

Koenig in Bonn to form the Leibniz Institute for

the Analysis of Biodiversity Change (LIB). The

material is separated into four collections: Inverte-

brates, Malacology, Annelida and Crustacea. This

work analysed inventoried records housed in the

Invertebrates Collection database of ZMH collect-

ed by the WH between 1966-1978. In addition, it

analysed the taxonomic coverage, spatial informa-

116 MARINE AND FISHERY SCIENCES 35 (1): 115-122 (2022)

encuentran completamente inventariadas ni son accesibles digitalmente. Además, las tasas de determinación de especies siguen siendo

alarmantemente bajas para algunas colecciones, y probablemente la mayoría de las determinaciones existentes deberían volver a analizar-

se. Esto resulta particularmente cierto con respecto a las muestras bentónicas de aguas profundas, caras y difíciles de obtener. Aquí revi-

samos la base de datos sobre el material recolectado por el BI alemán “Walther Herwig I y II” durante los cruceros de investigación entre

1966-1978 al Océano Atlántico Sudoccidental (SAO), almacenado en la colección de invertebrados (invertebrados excepto Mollusca,

Arthropoda y Annelida) del Museo Zoológico de Hamburgo. Encontramos que de 3.305 registros, donde al menos 204 especies fueron

identificadas, el 72% todavía se encuentra sin determinar a nivel de especie y el 65% a nivel de género. Si bien el muestreo de áreas geo-

gráficas más remotas y el uso de una mayor variedad de métodos de muestreo aún son necesarios, el apoyo a la capacitación de un ejército

de taxónomos será de suma importancia para superar la abrumadora tarea de analizar la enorme cantidad de muestras almacenadas en las

colecciones de los museos. Se necesita con urgencia un esfuerzo de toda la comunidad para abordar este problema, y la cooperación inter-

nacional debe desempeñar un papel clave en esta empresa.

Palabras clave: Investigación oceanográfica, Atlántico Sudoccidental, biodiversidad bentónica, historia natural, colecciones.

tion, including depth of the oceanographic sta-

tions. This will provide a comprehensive synthesis

on the current status and potentialities of the Pri-

mary Biodiversity Data (PBD) associated with

this particular collection, in order to encourage

further similar research in NHC worldwide.

Research cruises herein analysed covered a

large portion of the SAO off Brazil, Uruguay and

Argentina. The Brazilian continental margin is

strongly influenced by the western contour cur-

rents, the Brazil Current (BC) flowing southward

and the Brazilian Northern Current (BNC) flow-

ing northward. The BC, which is shallowest

between 15° S and 20° S, transports saline and

oligotrophic tropical waters, receiving additional

contribution from the South Atlantic Central

Waters (SACW), reaching a vertical extension of

about 500 m, and continues to flow southwards

towards the Subtropical Convergence (33° S-38°

S) where it merges with the Malvinas Current and

then flows away from the coast to the east

(Burone et al. 2021).

In the confluence zone, the Río de la Plata

estuary represents the greatest freshwater inflow

to the region, being one of the few geographical

features (i.e., Valdés Peninsula, North Patagonic

gulfs, and the Magallanes Strait) that influences

water circulation at a regional scale (Miloslavich

et al. 2011). Thus, the confluence of Malvinas and

Brazil currents, together with the abundant terres-

trial runoff of Río de la Plata, and the relatively

shallow waters of the area, combine to produce a

singular hydrographic system (Acha et al. 2008;

Franco-Fraguas et al. 2014).

The Patagonian Shelf (PS) extends for about

5,649 km along the Atlantic coast of South Amer-

ica from northern Uruguay (33° 51'21"S, 53° 11'

43"W) to the southern tip of Argentina, bordering

Chile (54° 55'39"S, 64° 52'12"W). The area of

the PS extends more than 3 million km2in

Uruguayan and Argentine territories and compris-

es coastal environments, the continental shelf

itself, the slope, and ocean basins. Its continental

shelf is generally up to 100 m in depth, and is the

largest and one of the most productive ecosys-

tems in the southern hemisphere (Acha et al.

2004; Miloslavich et al. 2011). Most of the PS is

thus influenced by the Malvinas current, which

originates in the Antarctic circumpolar current

and carries a high nutrient load north along the

Argentine and Uruguayan coasts. Nutrient-poor

waters of the Brazil current meet the Malvinas

current as it moves southward along the edge of

the slope (Piola et al. 2010). In the confluence or

transition zone (from 30° S to 46° S), a series of

oceanographic phenomena (eddies, marine fronts,

etc.) allows for high biological production (Fran-

co-Fraguas et al. 2014; Burone et al. 2021).

Thus, the geographical coverage of WH expe-

ditions during the period 1966-1978 constitutes

an invaluable sampling design, allowing lines of

research and the analysis of biogeographic pat-

terns at an enormous spatial realm, among others.

For most taxonomic groups, however, species

determination of benthic invertebrates needs thor-

ough revision. The estimated number of taxono-

mists devoted to invertebrates in this region is

low, and most research is focused on molluscs

and crustaceans (Meier and Dikow 2004;

Miloslavich et al. 2011).

We reviewed the ZMH invertebrates collec-

tion database and checked material from WH

cruises from 1966 to 1978. The database was

provided as several electronic spreadsheets, sep-

arated by taxonomic groups, often including two

different spreadsheets for determined and unde-

termined material. We then analysed the number

of records (i.e., assigned inventory number that

may refer to a single or several individuals or

colonies) and quantified the ratio of determined

lower taxa (species and genus) to total samples.

Assigned names were provided as is, since it was

beyond our aim to review species concepts

and/or nomenclatural issues. Due to different for-

mat of associated data, information on geograph-

ic coordinates of records was extracted (when

available), very often by cross-checking the

information provided against station lists and

117

CARRANZA ET AL.: BENTHIC INVERTEBRATES FROM THE RV ‘WALTHER HERWIG I AND II’

geographic references provided in other databas-

es. Then, this georeferenced information was

incorporated in a Geographic Information Sys-

tem, further detecting, fixing or deleting ‘suspi-

cious’ coordinates due to error in data entering or

handling. These records were mapped. Statistics

on the spatial distribution of records were calcu-

lated, and particularly, an overview of the distri-

bution of sampling effort associated with the

Marine Ecoregions of the World (MEOWs)

framework (Spalding et al. 2007) was provided.

When available, depth-distribution of the sam-

ples was also registered.

The database herein assembled includes 3,305

records, comprising material from seven animal

Phyla, namely Porifera, Cnidaria, Brachiopoda,

Bryozoa, Echinodermata, Hemichordata and

Chordata. Eighty six percent of records (2,840)

were georeferenced and associated with 8

MEOWs, representing Eastern Brazil, South-east-

ern Brazil, Rio Grande, Uruguay-Buenos Aires

Shelf, Malvinas, Patagonian Shelf, North Patag-

onian Gulfs and Channels and Fjords of Southern

Chile. Nearly 80% of records were concentrated

in three MEOWs: Malvinas (N = 947), Patagonian

Shelf (N = 797) and Uruguay-Buenos Aires Shelf

(N = 458). Some 242 records fall outside the lim-

its of MEOWs (i.e., 200 NM) and are thus referred

to as collected from international waters (Figure

1). From the total, 2,853 records had associated

depth data, showing an overall depth range from

1,400 to 33 m. Nearly half of the records (1,507)

came from depths shallower than 200 m.

Cnidaria was the best represented phylum, with

1,518 records, including hydrozoans (417 records)

and anthozoans, comprising both hexacorallian

(Antipatharia, Scleractinia, Corallimorpharia and

Actiniaria) and octocorallians (Pennatularia and

Alcyonacea). From the total number of records,

204 differently determined lower taxa were found,

including determination of morphospecies (e.g.,

‘sp.’ and ‘sp. 1’), but more than 72% of the mate-

rial is still undetermined (and some probably

unsorted) at species level. Excluding morphos-

pecies (i.e., not counting preliminary species

determinations such as ‘antarctica group’, ‘aff.’

and ‘cf.’, etc.), only 163 species names are men-

tioned in the database. If we look at genus, 65% of

records still lack determination (Table 1).

Groups with better taxonomic coverage

include Anthozoa (Scleractinia, Corallimorpharia

and Actiniaria), with all samples present in the

database determined at species level. In particu-

lar, the high degree of taxonomic coverage in

Actiniaria is due to the extensive work of Rie-

mann-Zürneck (1973, 1975a, 1975b, 1978, 1980,

1986a, 1986b) on this material. However, there

are 524 samples labelled as ‘Anthozoa indet’, and

it is unclear whether all Scleractinia, Corallimor-

pharia and Actiniaria have been determined, or

whether unidentified Anthozoa may include fur-

ther records of these groups (Table 1). In this line,

for four decades new species of Anthozoa have

continued to be discovered from this material

(Cairns 2012).

Amongst Echinodermata, Argentine re-

searchers such as Bernasconi (1972, 1973) stud-

ied the ophiuroids, echinoids and asteroids from

the 1966 cruise, and Hernández (1982) studied

the holothuroids. According to Brogger and

O’Hara (2015), WH ophiuroids from 1971 were

studied by Bartsch (1982). In spite of this, virtu-

ally all Asteroidea and 96% of Ophiuroidea sam-

ples are undetermined, as are roughly half of

Holothuroidea and Crinoidea. Thus, the current

status of the taxonomic coverage of the Echino-

dermata is far from complete. This, however,

does not reflect insufficient taxonomic knowl-

edge, but rather a simple lack of time and/or

opportunity to cross-check determinations made

by Argentine researchers using essentially the

same material. Voucher specimens seem to be

deposited both in Argentina (Museo Argentino de

Ciencias Naturales –MACN) and in the ZMH.

For example, and concerning Ophiuroidea, Brog-

ger and O’Hara reported material identified by

Bernasconi (1973) from Walther Herwig Stn. 277

(1966), as Ophiacantha vivipara Ljungman, 1871

118 MARINE AND FISHERY SCIENCES 35 (1): 115-122 (2022)

deposited in Argentina (MACN 27263). No Ophi-

uroids from this station are stored in the ZMH,

although there are Cnidarians and Bryozoans

from the same station. However, material for O.

vivipara is present in the ZMH collection, but

labelled generically concerning the procedence

(i.e., ‘WH 1966, Patagonischer Schelf, 36°52'-

54°51'S 54°01'-63°51'W, 95-800 m’).

This database is thus a ‘Gold mine’ of unreport-

ed PBD, gathered using a standardised methodol-

ogy and thus avoiding one of the most common

problems of museum collections: a skewed abun-

dance distribution due to collectors’ potentially

self-selecting underrepresented species over com-

mon ones (Guralnick and Cleve 2005). This col-

lection further provides an excellent coverage

119

CARRANZA ET AL.: BENTHIC INVERTEBRATES FROM THE RV ‘WALTHER HERWIG I AND II’

Figure 1. Surveyed stations by RV ‘Walther Herwig I and II’ in the Southwestern Atlantic Ocean from 1966 to 1978. It must be

noticed that since we were unable to retrieve location data for a number of stations, the map underestimates the spatial

coverage of expeditions.

40° 0 00' "

30° 0 00' "

50° 0 00' "

20° 0 00' "

10° 0 00 S' "

40° 0 00' "50° 0 00' "60° 0 00' "70° 0 00 W' "

Southwestern

Atlantic Ocean

from the study region, and it is hard to provide an

exact figure of how costly it will be to implement

the same sampling effort nowadays, but probably

ranging in the order of tens of millions in current

currency.

Once taxonomic work progresses, these data

based on qualitative sampling can be rapidly

assessed to produce biogeographic or macroeco-

logical studies. Meanwhile, entire regions may

remain poorly or completely uninventoried, forc-

ing conservation decisions to be based in incom-

plete data (Guralnick and Cleve 2005). Since

there is an urgent need for the availability of this

information, we must encourage support and

funding for comprehensive inventories of marine

invertebrates by trained researchers in taxonomy.

In this context, the decreasing number of taxono-

mists worldwide should be enhanced by means of

global initiatives such as NSF-PEET and

DIVERSITA. NSF-funded Planetary Biodiversi-

ty Inventories (PBI) targets the global coverage

of a given Taxon, while the Partnerships for

Enhancing Expertise in Taxonomy (PEET) focus

in training the next generation of taxonomists.

Yet even so, taxonomy seems to be facing the

same chronic issues: inadequate funding and the

lack of training and recruitment of taxonomists

(Britz et al. 2020). This trend has to be quickly

reversed. Otherwise, and with little or no special-

ists available worldwide for a number of taxo-

nomic groups, we will not be able to exploit the

information stored in NHCs.

120 MARINE AND FISHERY SCIENCES 35 (1): 115-122 (2022)

Table 1. Summary of taxonomic coverage of samples stored in the Invertebrates II Collection of the Zoological Museum of

Hamburg (ZMH). See text for details.

Phylum Original taxon Total Determined Determined % determined

name in the samples (species) (genera) (species)

Brachiopoda Brachiopoda 104 0

Bryozoa Bryozoa 142 0

Chordata Ascidiacea 239 22 9

Cnidaria Anthozoa 528 0

Cnidaria Antipatharia 3 0

Cnidaria Pennatularia 108 2 2 2

Cnidaria Alcyonacea 196 93 103 47

Cnidaria Hydrozoa 417 302 115 72

Cnidaria Scleractinia 92 92 1

Cnidaria Corallimorpharia 2 2 1

Cnidaria Actiniaria 172 172 1

Echinodermata Asteroidea 530 1 0

Echinodermata Ophiuroidea 137 6 4

Echinodermata Crinoidea 9 4 44

Echinodermata Holothuroidea 39 20 3 51

Echinodermata Echinoidea 211 208 3 99

Hemichordata Pterobranchia 17 0

Porifera Porifera 359 19 0

Total 3,305 924 245 28

ACKNOWLEDGEMENTS

This work is part of the MSc. Thesis of P.

Limongi (PEDECIBA-CAP). Alvar Carranza

wishes to thank the ZMH staff, in particular

Helma Roggenbuck, Dr Bernahard Hausdorf and

Dr Martin Schwentner for their kind assistance

during a sabbatical leave in Hamburg.

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