Hall of Fame - Arthur Humes (2)

His Output

Arthur Humes was a stirling systematist and showed a truly astounding level of career productivity. In the course of nearly 60 years research on copepods, Arthur has established 18 new families, 151 new genera, 700 new species and 1 new subspecies. This amazing contribution to copepod taxonomy is one of the greatest taxonomic efforts ever accomplished by a single individual in the field of Crustacea. In years of highest productivity his output of new species descriptions would amount to one per week as for example in 1973 when he published the first revision of the Lichomolgoidea with Jan Stock (Humes and Stock, 1972), or in 1982 when he produced (alone or jointly) major contributions to the systematics of the Taeniacanthidae, Xarifiidae and Rhynchomolgidae (Dojiri and Humes, 1982; Humes, 1982a-b; Humes and Dojiri, 1982). Arthur's taxonomic work was build on a solid base of massive collections which enabled him to study the variability within species and between geographical areas and hosts. It is therefore not surprising that nearly all of his species have successfully stood the test of time. In fact, only one genus (Metaxymolgus Humes and Stock, 1972) and two species (Sciaenophilus inopinus Humes, 1957; Oncaea praeclara Humes, 1988) have been synonymized since, the last two being representatives of groups Arthur was less acquainted with - the fish parasites and the marine plankton.

As the preeminent authority in the field Arthur Humes received numerous collections, foremost amongst these were the copepods collected from hydrothermal vents and cold seeps. More importantly, Arthur himself accumulated with unrelenting stamina a stupendous amount of material during his fieldwork in West Africa, Madagascar, the Moluccas, Enewetak Atoll, the Great Barrier Reef and New Caledonia. Although the actual figure is undoubtedly higher, specimen counts based on his published papers from 1955 onwards suggest that Arthur Humes (and his collaborators) sorted and examined over 300,000 individual copepods! On average, this accounts to 130 copepods per week over a 45-year period.

Arthur remained extraordinarily energetic, even at a later age, and this is best demonstrated by the remarkable fact that nearly half of his new species and genera were described during his 20-year tenure as Editor of the Journal of Crustacean Biology between 1980 and 1999.

The Diversity of his Research

Sixteen phyla of invertebrates are utilized as hosts for copepods (Huys and Boxshall, 1991). Arthur Humes almost covered this entire spectrum and published on twelve of them, including the first records of copepods occurring on the lophophorate phyla Phoronida and Brachiopoda (Boxshall and Humes, 1988; Humes and Boxshall, 1988) and new species from bizarre host groups such as the flatworms (Humes, 1997) and the vestimentiferans (Humes, 1973a; Humes & Dojiri, 1980, 1981). Of all groups his descriptive work on the poecilostomatoids and siphonostomatoids associated with cnidarian hosts will remain as an enduring monument. In 1986, Humes estimated the number of described copepod species associated with cnidarians at 416, but to his credit, he forgot to mention that 282 (or 68%) had been described by himself (Humes, 1985b). In the remaining years of his career Arthur added another 110 species, bringing his personal total for this host group to 392 species. The evolutionary success of the copepod-cnidarian association did not distract his attention from copepods occurring on other invertebrate groups such as the echinoderms (137 species), molluscs (54 species) and to a lesser extent the crustaceans (20 species) and the polychaetes (14 species).

Although the great majority of Arthur's material was collected by snorkeling and SCUBA diving in shallow subtidal habitats, his fascination for the group led him to cover the entire depth range of associated copepods. His early research in the 1940s and 1950s focused primarily on crustacean and molluscan hosts from easily accessible habitats. Surveys of marsh crabs, mud shrimps and even the edible mussel in North America and West Africa resulted in the unexpected discovery of several harpacticoids and provided Arthur with the impetus to extend his search for copepods to both the sublittoral environment and to other host groups such as the cnidarians and echinoderms. With the exception of the occasional paper on deep-sea copepods (Humes, 1973a, 1974; Humes & Grassle, 1979), his career went full circle in the 1980s and 1990s when he received the copepods collected at hydrothermal vents in the eastern Pacific and the mid-Atlantic and deeply immersed himself in a totally unknown fauna. Over the years, the examination of nearly 60,000 specimens from these habitats culminated in the description of three new families, 18 new genera and 61 new species (Humes and Segonzac, 1998; Humes, 1999a).

Copepods occur on vertebrate as well as invertebrate hosts but Arthur rarely published on associates of vertebrates. In over five decades he published only five taxonomic papers on vertebrate associates (Humes 1957, 1964, 1965; Humes and Rosenfield, 1960; Gooding and Humes, 1963), the most unusual of these being the description of a new harpacticoid species, Harpacticus pulex Humes, 1964, found on the sloughed skin of a porpoise and a manatee in Florida.

Altogether Humes published on six of the ten currently recognized orders of copepods including describing a new marine calanoid, Ridgewayia fosshageni Humes and Smith, 1974. This species was observed forming free-swimming aggregations in the immediate vicinity of the sea anemone Bartholomea annulata Lesueur, although they were never observed resting on or feeding on the anemone (Humes and Smith, 1974). These aggregations might now be categorized as swarming behaviour and the possibility of association with the anemone requires further experimental verification. His recent work on hydrothermal vent copepods also led Arthur into describing new free-living members of the marine families Cyclopinidae (order Cyclopoida) and Misophriidae (order Misophrioida), a project that was completed just before his death (Humes, 1999b).

While the majority of Arthur's contributions to copepods were purely taxonomic, the collection and processing of vast numbers of copepods also enabled him to study their development in a variety of families such as the Temoridae in the Calanoida, the Clausidiidae in the Poecilostomatoida and the Tisbidae in the Harpacticoida (Humes, 1955, 1960, 1986c). Similarly, his survey of numerous hosts led him to demonstrate another phenomenon, that of the multiple associations. Humes showed that one host individual or colony may frequently support more than one copepod. A striking example, but not unique in its kind (Humes, 1994), is the hard coral Acropora hyacinthus (Dana) which harbours nine poecilostomatoid species. Arthur's research revealed that each of the 12 cnidarian hosts in New Caledonia and 13 hosts in the Moluccas had at least five species of associated copepods. This observation illustrates that, particularly in tropical and subtropical areas, copepod diversity is likely to exceed invertebrate host diversity, often by a factor five or higher.

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