JOURNAL OF CRUSTACEAN BIOLOGY, 21(1): 13-27, 2001

AN APPRECIATION OF THE CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS

Rony Huy s and Geoff Boxshall

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK (e-mail: [email protected] ; [email protected] )

Arthur Humes had a long and distinguished each colony or fragment of coral is isolated career in biology. His first publication was in a plastic bag. In the laboratory the coral in 1938 and his first paper on copepods was and sea water are placed in a bucket to which in 1941. Since then he has produced an in­ sufficient 95% ethanol is added to make an credible 252 publications on copepods. Arthur approximately 5% solution. The coral is left is best known to the members of The Crus­ in this solution at ambient temperature for tacean Society as the founding editor of the several hours or over night. Then the coral Journal of Crustacean Biology but, in the is thoroughly rinsed by shaking well and the copepod research community, he is recog­ wash water is poured through afine net (120 nised as being responsible for opening up one holes per 2.5 cm. each hole approximately of the four main fields of copepod systemat- 100 pm square). The copepods are then ics: the taxonomy and biodiversity of cope­ picked from the sediment retained in the net. pods associated with, and parasitic on, ma­ It appears that the dilute alcohol, together rine invertebrates. Starting virtually from with the accumulating products of decompo­ scratch, building a new field has required an sition, stimulates the copepods to leave the immense volume of descriptive taxonomy. By polyps of the coral host, and they fall to the providing this he has established the basic bottom of the container. In comparison, rapid framework of knowledge on these groups— washing of the freshly collected corals usu­ the knowledge base, which has, in recent ally yielded very few xarifiids (Humes and years, provided phylogeneticists with an in­ Dojiri, 1982). This method can be applied credibly rich source of data. equally to many other host groups, such as Arthur Humes’s success in opening up the soft corals, echinoderms, sponges etc. field of invertebrate associates depended ini­ In addition to his extraction method, tially on his recognition that virtually any ma­ Humes also pioneered various improvements rine macroinvertebrate was a potential host to in methods for the microscopic study of cope­ copepods. In 1993, in the Maxilliped lecture pods. A much-cited methods paper is Humes given when he was president of the World As­ and Gooding (1964) in which the hanging- sociation of Copepodologists, Humes calcu­drop slide method is described in detail. In lated that associated copepods are known this short paper Humes and his brilliant stu­ from only 1.14% of the 151,400 species of dent, Richard Gooding, recommended the use potential marine invertebrate hosts worldwide of lactic acid as the best clearing agent for the (Humes, 1994). On these calculations, the preparation of temporary mounts for whole amount of descriptive taxonomy remaining to or dissected copepods. They also recom­ be done is enormous. mended that dissections be carried out in a drop of lactic acid on a 22-mm coverslip ce­ His M eth o d s mented to a wooden slide with a bored cen­ During his career, Humes himself surveyed tral hole 15 mm in diameter. The preparation a huge range of invertebrate host taxa and de­ is then ready for examination under the com­ veloped an extraction technique, which was pound microscope by inverting the slide. One to prove suitable for coaxing the copepods out of the major advances of this method is that from many hosts. This method, as applied to a single specimen can usually provide a full the extraction of xarifiid copepods from hard set of observations of all appendages. Humes corals (Humes and Dojiri, 1982) is repeated and Gooding explicitly stated another major here: immediately on collection in the fieldadvantage of this technique is that it causes

13 14 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001

Table 1. New copepod taxa authored or co-authored by Arthur Humes (*: includes one subspecies).

Decade Number of new species Number of new genera Number of new families 1941-1950 4 1 1951-1960 43 6 3 1961-1970 172* 23 1971-1980 166 59 4 1981-1990 197 29 4 1991-2000 119 33 7 Totals 701* 151 18 little or no compression of the copepods orresearch on copepods, Arthur has established their dissected parts—a problem that plagued 18 new families, 151 new genera, 700 new the taxonomic studies of many copepod re­ species, and 1 new subspecies (Table 1). This searchers before 1964 (and still does). They amazing contribution to copepod taxonomy also noted that staining is usually undesirable is one of the greatest taxonomic efforts ever in this kind of mount and that phase-contrast accomplished by a single individual in the microscopy may be used. In all these respects field of Crustacea. In years of highest pro­ Humes and Gooding pioneered the best mod­ ductivity his output of new species descrip­ ern practice, except that the development of tions would amount to one per week (Fig. 1) differential interference contrast microscopy as for example in 1973 when he published the has essentially replaced phase contrast mi­ first revision of the Lichomolgoidea with Jan croscopy. Stock (Humes and Stock, 1973), or in 1982 when he produced (alone or jointly) major His O u t p u t contributions to the systematics of the Tae­ Arthur Humes was a sterling systematist niacanthidae, Xarifiidae, and Rhynchomolgi­ and showed a truly astounding level of career dae (Dojiri and Humes, 1982; Humes, 1982a, productivity. In the course of nearly 60 years b; Humes and Dojiri, 1982). Arthur’s taxo­

New species

60 700

600

500

0)« o 400 a.o (0 o z 300

200

100

w.T-COtOKQiCVV-COCOOCvVCOCDOCVV-COQOOO/VCOCOOCVV-^COO w- Ts. ir\ /r\ fr\ fr\ Cr\ fr\ N N /V Av N rv-v rr\ rr\ /'Ts /'"Tn rrs rrv rt\ /—y

Year

Fig. 1. Number per year and cumulative number of new species described by Arthur Humes between 1941 and 2000. HUYS AND BOXSHALL: CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS 15

New genera

160

140 30

120 25

co 20 co Ö)fl) o z

20

Year

Fig. 2. Number per year and cumulative number of new genera described by Arthur Humes between 1941 and 2000. nomic work was built on a solid base of mas­ 300,000 individual copepods! On average, sive collections which enabled him to study this accounts to 130 copepods per week over the variability within species and between geo­ a 45-year period (Fig. 3). graphical areas and hosts. It is therefore not Arthur remained extraordinarily energetic, surprising that nearly all of his species have even at a later age, and this is best demon­ successfully stood the test of time. In fact, strated by the remarkable fact that nearly half only one genus (.Metaxymolgus Humes and of his new species and genera were described Stock, 1972) and two species Sciaenophilus (, during his 20-year tenure as Editor of the inopinus Humes, 1957; Oncaea praeclara Journal of Crustacean Biology between 1980 Humes, 1988) have been synonymized since, and 1999 (Figs. 1, 2). the last two being representatives of groups Arthur was less acquainted with, the fish par­ T h e D iv er sit y o f H is R e se a r c h asites and the marine plankton. Sixteen phyla of invertebrates are utilized As the preeminent authority in the field as hosts for copepods (Huys and Boxshall, Arthur Humes received numerous collections, 1991). Arthur Humes almost covered this en­ foremost amongst these were the copepods tire spectrum and published on twelve of collected from hydrothermal vents and cold them (Fig. 4), including the first records of seeps. More importantly, Arthur himself ac­ copepods occurring on the lophophorate cumulated with unrelenting stamina a stu­ phyla Phoronida and Brachiopoda (Boxshall pendous amount of material during his field­ and Humes, 1988; Humes and Boxshall, work in West Africa, Madagascar, the Moluc­ 1988) and new species from bizarre host cas, Enewetak Atoll, the Great Barrier Reef, groups such as the flatworms (Humes, 1997) and New Caledonia. Although the actual fig­ and the vestimentiferans (Humes, 1973a; ure is undoubtedly higher, specimen counts Humes and Dojiri, 1980, 1981). Of all groups based on his published papers from 1955 on­ his descriptive work on the poecilostomatoids wards suggest that Arthur Humes (and his and siphonostomatoids associated with collaborators) sorted and examined over cnidarian hosts will remain as an enduring 16 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001

Individuals examined

70000 350000

60000 300000

50000 250000

« 40000 200000 "ö

30000 150000 >

20000 100000

10000 50000

Year

Fig. 3. Number per year and cumulative number of individuals examined by Arthur Humes between 1941 and 2000. monument. In 1986, Humes estimated the the impetus to extend his search for copepods number of described copepod species asso­ to both the sublittoral environment and to ciated with cnidarians at 416, but to his credit, other host groups such as the cnidarians and he forgot to mention that 282 (or 68%) had echinoderms. With the exception of the oc­ been described by himself (Humes, 1985b). casional paper on deep-sea copepods (Humes, In the remaining years of his career Arthur 1973a, 1974; Humes and Grassle, 1979), his added another 110 species, bringing his per­ career went full circle in the 1980s and 1990s sonal total for this host group to 392 species.when he received the copepods collected at The evolutionary success of the copepod- hydrothermal vents in the eastern Pacific and cnidarian association did not distract his at­ the mid-Atlantic and deeply immersed him­ tention from copepods occurring on other in­self in a totally unknown fauna. Over the vertebrate groups such as the echinoderms years, the examination of nearly 60,000 spec­ (137 species), molluscs (54 species), and to imens from these habitats culminated in the a lesser extent the crustaceans (20 species) description of three new families, 18 new and the polychaetes (14 species) (Fig. 4). genera, and 61 new species (Humes and Although the great majority of Arthur’s Segonzac, 1998; Humes, 1999a). material was collected by snorkeling and Copepods occur on vertebrate as well as SCUBA diving in shallow subtidal habitats, invertebrate hosts, but Arthur rarely published his fascination for the group led him to cover on associates of vertebrates. In over five the entire depth range of associated copepods. decades he published only five taxonomic pa­ His early research in the 1940s and 1950s fo­ pers on vertebrate associates (Humes, 1957, cused primarily on crustacean and molluscan 1964, 1965; Humes and Rosenfield, 1960; hosts from easily accessible habitats. Surveys Gooding and Humes, 1963), the most unusual of marsh crabs, mud shrimps and even the ed­ of these being the description of a new ible mussel in North America and West Africa harpacticoid species, Harpacticus pulex resulted in the unexpected discovery of sev­ Humes, 1964, found on the sloughed skin of eral harpacticoids and provided Arthur with a porpoise and a manatee in Florida. HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD S Y STEMATICS 17

CEPHALOPODA [4] OPHIUROIDEA [15] 11%

HOLOTHUROIDEA [45] ECHINOIDEA [32] GASTROPODA [12] 33% 23% 22%

BIVALVIA [38] 71%

SCYPHOZOA [3] 1% CRINOIDEA [14] 10%

ASTEROIDEA [31] 23% HYDROZOA [8] 2%

ANTHOZOA [381] 97% POLYCHAETA [14]

CNIDARIA [392] VESTIMENTIFERA [4] PLATYHELMINTHES

MOLLUSCA [54]

ECHINODERMATA [137]

PORIFERA [6] BRACHIOPODA [1]

HOST UNKNOWN [5] PHORONIDA [1]

FREE-LIVING [2] ECHIURA [1]

MAMMALS [1] CRUSTACEA [20] HYDROTHERMAL VENTS [57] FISH [1] UROCHORDATA [4]

Fig. 4. Pie chart (below) displaying contribution of each host category/habitat in Arthur Humes’s research. Rela­ tive importance of individual classes in most important host phyla (Cnidaria, Echinodermata, Mollusca) displayed in separate pie charts (upper). All pie charts based on number of new copepod species described (indicated in square brackets). 18 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001

Table 2. List of copepod orders and families containing new species established by Arthur Humes. Family names in parentheses indicate the two species established by Humes that were subsequently synonymised. The inclusion of the Ascidicolidae is based on recognition of the affinities of the previously unplaced genus Gomphopodarion Humes, 1974. [*: Includes one subspecies.]

C a la n o id a P oecilostomatoida S iphonostomatoida Diaptomidae 2 Anchimolgidae 84 Asterocheridae 41 Ridgewayiidae 1 Anthessiidae 15 B rychiopontiidae 1 Catiniidae 1 (Caligidae) (1) C yc l o po id a Clausidiidae 22 Cancerillidae 3 Ascidicolidae 1 Erebonasteridae 4 Coralliomyzontidae 6 Cyclopidae 3 Lamippidae 1 Dirivultidae 52 Cyclopinidae 2 Lichomolgidae 41 Ecbathyriontidae 1 Lernaeidae 1 Lubbockiidae 1 Megapontiidae 1 Notodelphyidae 4 Macrochironidae 9 Micropontiidae 1 Myicolidae 7 Nanaspididae 11 H arpacticoida Mytilicolidae 2 Nicothoidae 1 Ameiridae 1 Octopicolidae 4* Stellicomitidae 9 Cancrincolidae 4 (Oncaeidae) (1) Canuellidae 2 Pseudanthessiidae 24 U n pla ced G e n er a Canthocamptidae 2 Rhynchomolgidae 187 Parangium Humes, 1985 1 Diosaccidae 1 Sabelliphilidae 9 Bythocheres Humes, 1988 1 Harpacticidae 1 Serpulidicolidae 1 Peltidiidae 1 Synapticolidae 28 Porcellidiidae 1 Synaptiphilidae 1 Tegastidae 5 Taeniacanthidae 12 Tisbidae 4 Thamnomolgidae 4 Vahiniidae 2 M isophrioida Xarifiidae 76 Misophriidae 1

Altogether Humes published on six of the Tisbidae in the Harpacticoida (Humes, 1955, ten currently recognised orders of copepods 1960, 1986c). Similarly, his survey of nu­ (Tables 2, 3) including describing a new ma­ merous hosts led him to demonstrate another rine calanoid, Ridgewayia fosshageni Humes phenomenon, that of the multiple associa­ and Smith, 1974. This species was observed tions. Humes showed that one host individ­ forming free-swimming aggregations in the ual or colony may frequently support more immediate vicinity of the sea anemone than one copepod. A striking example, but not Bartholomea annulata Lesueur, although they unique in its kind (Humes, 1994), is the hard were never observed resting on or feeding on coralAcropora hyacinthus (Dana), which har­ the anemone (Humes and Smith, 1974). These bours nine poeciiostomatoid species. Arthur’s aggregations might now be categorised as research revealed that each of the 12 cnidar- swarming behaviour, and the possibility of as­ ian hosts in New Caledonia and 13 hosts in sociation with the anemone requires further the Moluccas had at least five species of as­ experimental verification. His recent work on sociated copepods. This observation illus­ hydrothermal vent copepods also led Arthur trates that, particularly in tropical and sub­ into describing new free-living members of tropical areas, copepod diversity is likely to the marine families Cyclopinidae (order Cy­ exceed invertebrate host diversity, often by a clopoida) and Misophriidae (order Misophri­ factor five or higher. oida), a project that was completed just be­ fore his death (Humes, 1999b). H arpacticoids , o r H o w It A l l S tarted While the majority of Arthur’s contribu­ Arthur’s interest in associated copepods tions to copepods were purely taxonomic, the was stimulated by the discovery of a new collection and processing of vast numbers of harpacticoid in the gili chamber of marsh copepods also enabled him to study their de­ crabs when he was studying their nemertean velopment in a variety of families such as parasites as part of his Ph.D. at Lousiana State the Temoridae in the Calanoida, the Clau­ University (Humes, 1941). This very first new sidiidae in the Poecilostomatoida, and the copepod, Cancrincola plumipes, was a rep- HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS 19

Table 3. List of copepod orders and families containing new genera established by Arthur Humes. The inclusion of the Ascidicolidae is based on recognition of the affinities of the previously unplaced genus Gomphopodarion Humes, 1974.

C y clo po id a P oecilostomatoida S iphonostomatoida Ascidicolidae 1 Anchimolgidae 27 Asterocheridae 12 Cyclopinidae 1 Catiniidae 1 Brychiopontiidae 1 Notodelphyidae 1 Clausidiidae 1 Cancerillidae 2 Erebonasteridae 2 Coralliomyzontidae 4 H arpacticoida Lichomolgidae 6 Dirivultidae 13 Cancrincolidae 1 Lubbockiidae 1 Ecbathyriontidae 1 Canthocamptidae 1 Macrochironidae 1 Nanaspididae 1 Myicolidae 1 Nicothoidae 1 Octopicolidae 1 Stellicomitidae 5 Pseudanthessiidae 3 Rhynchomolgidae 38 U n pla ced G enera Sabelliphilidae 6 Parangium Humes, 1985 Synapticolidae 5 Bythocheres Humes, 1988 Taeniacanthidae 3 Thamnomolgidae 3 Vahiniidae 1 Xarifiidae 4

resentative of the Ameiridae (currently clas­ and hermit crabs. The latter group offers a sified as the family Cancrincolidae), and good example of independent colonization by Arthur returned to publish on this group on three different families of harpacticoids, the several subsequent occasions when surveying Tisbidae, Porcellidiidae, and Canuellidae terrestrial crabs along both sides of the cen­ (Humes and Ho, 1968a, b). tral Atlantic. The genera Cancrincola Wilson, 1913, andAntillesia Humes, 1958, utilize pri­ C o pepo d s a n d C n id a r ia - marily Gecarcinidae (true land crabs) as hosts H is L ife t im e O b sessio n and are restricted to the Atlantic basin. Stim­ Although Arthur published on a plethora of ulated by his initial discovery, Arthur ex­ host groups, it was the Cnidaria that was to tended his search for harpacticoids to another occupy him for most of his scientific career. group of terrestrial crabs, the Grapsidae, and Copepods are associated with all three classes discovered that at least in the Indo-Pacihc a of Cnidaria, the Hydrozoa, the Scyphozoa, different genus, Pholetiscus Humes, 1947, and the Anthozoa, and Humes published on was associated with this host group (Humes, all three of them (Fig. 4). Humes’s studies 1956). Arthur’s recognition of the cantho- of the poecilostomatoid family Macrochi­ camptid affinities of this new genus was sig­ ronidae contributed significantly to the sys- nificant because it demonstrated that the pri­ tematics of copepods associated with the marily free-living Harpacticoida had entered polyp stages of hydroids (e.g., Humes, 1966, into association with land crabs at least twice 1977; Humes and De Maria, 1969) and the during their evolutionary history. scyphozoan medusae (e.g., Humes, 1969, During his career Arthur Humes described 1970), but it is his work on the Anthozoa that 23 species of harpacticoids belonging to ten deserves special mention. His first paper on different families (Table 2) and utilizing eight this group was not until 1958 when he de­ different host groups: Crustacea, Hydrozoa, scribed a new species of Lamippe Bruzelius, Scyphozoa, Scleractinia, Alcyonacea, Bi­ 1858, from West African pennatulaceans, but valvia, Holothuroidea and Echinoidea. what followed after his fieldwork in Nosy Bé Harpacticoids have generally been considered was a real deluge of new taxa, representing to be only rarely associated with inverte­ 251 new species associated with hexacoral- brates, but Humes’s studies have firmly es­ lian hosts and 130 new species with octoco- tablished that at least in the modified families rallian hosts. Except for the Ceriantharia and Tegastidae, Peltidiidae, and Porcellidiidae the Corallimorpharia, Arthur Humes surveyed several lineages have entered into symbiotic hosts of all major anthozoan groups (Fig. 5). relationships with cnidarians, echinoderms, No copepodologist before him has dominated 20 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001

Table 4. Number of specimens examined by Arthur group. The vast majority of these associates Humes for each host category. belong to the Anchimolgidae, Rhynchomol­ gidae, and Xarifiidae. Members of the An­ Porifera 357 chimolgidae are exclusively associated with Cnidaria 103,127 scleractinian corals and are currently accom­ Anthozoa 96,367 modated in 28 genera—27 of which having Alcyonacea 34,344 been described by Humes and the remaining Gorgonacea 17,031 Pennatulacea 384 one being named after him by Sebastian and Stolonifera 844 Pillai (1973) Humesiella [ ]. Arthur named no Telestacea 1,896 less than 84 species of Anchimolgidae which Actiniaria 3,900 represents nearly 95% of the total number de­ Antipatharia 2,966 scribed. Scleractinia 33,734 Zoanthidea 1,268 The Xarifiidae is a good example of the Hydrozoa 5,592 progress stimulated by Humes’s research. Scyphozoa 1,168 Xarifiids are internal parasites of both her- matypic and ahermatypic scleractinian corals Platyhelminthes 9 and had never been reported before 1960. Polychaeta 2,590 They inhabit the gastrovascular cavities of the coral polyps and can best be extracted using Vestimentifera 236 the techniques described above (Humes and Dojiri, 1982). This family had humble be­ Mollusca 33,069 Bivalvia 30,080 ginnings: in 1960 Humes described just two Cephalopoda 382 species, which he placed in a new genus, Xar­ Gastropoda 2,607 ifia, the type genus of his new family. Cur­ rently the Xarifiidae comprises 84 species in Echinodermata 86,408 Asteroidea 18,196 four genera, and Humes is author or co-au­ Crinoidea 7,684 thor (with either Ju-shey Ho or Masahiro Do­ Echinoidea 10,703 jiri) of all four genera and the great majority Holothuroidea 19,244 (76) of the 84 species (Humes, 1960b, 1962, Ophiuroidea 30,581 1985a; Humes and Ho, 1967, 1968; Humes Brachiopoda 11 and Dojiri, 1982, 1983). The family is in­ completely known, for Humes (1985a) men­ Phoronida 3 tioned an additional 16 new species in his possession that were represented by too few Echiura 5 specimens to allow their description. The dis­ Crustacea 18,572 tribution of xarifiids is limited by the eco­ logical requirements of their coral hosts, but Urochordata 860 they occur from the Red Sea-Madagascar area eastward to an arc formed by Japan-Enewe- Fish 49 tak Atoll-New Caledonia (Humes, 1985a). Mammals 158 They are absent from the Caribbean and from the Pacific east of 166°W (e.g., from Hawaii, Hydrothermal vents 57,922 Moorea and Panama). Free-living 1,628 The Xarifiidae is a classic example of a family of associated copepods that, once their Host unknown 859 habitat was discovered and a method to ex­ tract them was developed, proved to be both Total 305,863 widespread and common. In stark contrast is the family Vahiniidae, also associated with cnidarian hosts, in this case antipatharians, this field as Arthur did, and we suspect none and also established by Humes (Humes, ever will again. 1967). The family was based on a monotypic Among the various groups of the Hexaco­ new genus Vailima Humes, 1967 and a sec­ rallia, the Scleractinia, or hard corals, have ond species was added to the genus in 1979 more copepod associates than any other (Humes, 1979) but, despite extensive sam- HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD S Y STEMATICS 21

ANTHOZOA

Zoanthidea [5] Alcyonacea [99]

o Antipatharia [14] Gorgonacea [19] no Stolonifera [4] Actiniaria [22] o

F Pennatulacea [4]

Scleractinia [210] Telestacea [4]

Fig. 5. Pie chart displaying detailed host utilization of new species described by Arthur Humes from hexacorallian and octocorallian anthozoans. Number of new species indicated in square brackets.

pling of antipatharians, more vahiniids have Indo-Pacific alcyonaceans, 96 of which were not been collected. The expenditure of suffi­ described by Humes, solely or jointly with M. cient search effort by Arthur Humes now al­ Dojiri, B. W. Frost, J.-s. Ho or J. H. Stock lows us to conclude that these animals are (Humes, 1990a, 1996). genuinely rare. Within the Octocorallia, the Alcyonacea, or T h e L ichomolgoid C o m p l e x soft corals, have the greatest number of cope­ Over his long career Humes exhibited a pod associates, and as with the scleractinians, particular affinity for one family-group above our knowledge on this group bears the strong all others, the Lichomolgidae. The enormous stamp of Arthur Humes’s approach to sys- numbers of new species and genera added to tematics. The first unequivocal association of this family-group by Humes and his collab­ a copepod with an alcyonacean was reported orators over the years stimulated two major by Stock and Kleeton (1963) who discovered revisionary works. The first revision was pub­ Critomolgus bulbipes (Stock and Kleeton, lished in 1973 in collaboration with Jan Stock 1963) on two soft corals in the vicinity of (Humes and Stock, 1973), although the new Banyuls. At that time nothing was known genera and families were validly published a about such associations in the tropical Indo- year earlier (Humes and Stock, 1972). In this Pacific. However, Humes’s extensive field­ work three new families, Pseudanthessiidae, work in Madagascar, New Caledonia, the Rhynchomolgidae, and Urocopiidae, were es­ Moluccas and Enewetak Atoll triggered a se­ tablished and placed in a new superfamily, the ries of papers on soft coral associates, and Lichomolgoidea, together with the two ex­ these served to demonstrate that copepods are isting families, the Lichomolgidae and Sabel­ particularly abundant on alcyonaceans and liphilidae. These families were distinguished gorgonaceans. Three decades after the initial largely on the basis of differences in the seg­ discovery of a copepod on a soft coral, 98 mentation and setation of the first to fifth copepod species are now known to occur onswimming legs. However, by the mid-1990s 22 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001 more than 40 new genera had been added to myicolids are serious pests of commercially the lichomolgoid complex, and the bound­ cultured bivalves and have been responsible aries between the five constituent families had for mass mortality in cultured bivalves (Ho become blurred. Humes and Boxshall (1996) and Yoosukh, 1994; Ho and Zheng, 1994). As undertook a further revision of the lichomol­ the economic importance of aquaculture in­ goid complex, excluding the Urocopiidae, and creases world wide, the basic descriptive tax­ recognized six new families, the Anchimol­ onomy of these potential pests generated by gidae, Kelleriidae, Octopicolidae, Macrochi­ Humes will provide the platform on which ronidae, Synapticolidae, and Thamnomolgi­ subsequent life cycle and biological research dae. Together with his numerous co-authors, can be built. Humes’s cumulative contribution to the ex­ pansion of knowledge of the lichomolgoid E c h in o d e r m A sso cia tes - complex is a total of eight new families (ex­ A L a s t in g A ffiliatio n cluding the Urocopiidae), 90 new genera, and All five classes of echinoderms serve as an amazing 385 new species. These absolute hosts for copepods, and Arthur made major numbers represent 80% of the known fami­ contributions across all five, totalling 137 new lies, 67% of the genera, and 68% of the species (Fig. 4). He published his first paper species, respectively. on the group in 1958 and continued working on echinoderm associates throughout his ca­ T h e M o l lu sc Pa r a sites reer until his death. In fact, his last paper that During the 1950s Arthur almost exclu­ went to press dealt with the biology and tax­ sively wrote on copepods associated with onomy of species ofOphiopsyllus Stock, molluscs and echinoderms. It was during this Humes, and Gooding, 1963, and Pseudan­ period that his first disciple, Roger Cressey, thessius Claus, 1889, associated with ophi- entered his laboratory and jointly worked up uroids in Belize (with Gordon Hendler). the material collected from West African bi­ Arthur’s research on echinoderm associates valves (Humes and Cressey, 1958a). Mean­ serves to demonstrate the sheer abundance while, Arthur concentrated also on the mol­ of copepods associated with marine inverte­ lusc associates from Madagascar which re­ brate hosts. Despite his own attempt to esti­ sulted in a major contribution the following mate the number of copepods (Humes, 1994), year (Humes, 1959). Over his career Humes their absolute abundance appears to be be­ returned many times to the study of parasites yond imagination and certainly beyond cal­ of marine molluscs, paying particular atten­ culation. One classic example of the tremen­ tion to the heterogeneous family, the Myi­ dous carrying capacity of these hosts is colidae, established without diagnosis by Ya­ demonstrated in his first paper dealing with maguti (1936). In 1986 Humes restricted the the large tropical basket star Astroboa nuda concept of the family by excluding five gen­ (Lyman) (Humes, 1973b). From three basket era which he placed in a newly designated stars (approximately 20 cm in diameter) ex­ family, the Anthessiidae. Humes (1986a) re­ amined, Arthur recovered a staggering 27,209 tained only the genera Myicola, Pseudomyi­individuals of Collocherides astroboae Stock, cola, and Ostrincola in the Myicolidae and 1971, and Doridicola micropus (Humes, provided detailed diagnoses of both families. 1973). Another example is the siphonosto- Later Humes and Boxshall (1988) added matoid family Stellicomitidae, the first new Parostrincola, unique within the family in its family recognised by Humes (Humes and utilization of an intertidal brachiopod, Lin­ Cressey, 1958b). These tiny associated cope­ gula anatina Lamarck, as host, and Ho and pods inhabit starfishes and can reach enor­ Kim (1992) established two further new gen­ mous population densities on their hosts. era, one of which,Exostrincola, was based on Humes (1971), for example, reported 1,420 Ostrincola simplex Humes, 1958. The myi- individuals of Stellicomes supplicans Humes, colids are typically parasites of marine bi­ 1971, on just two individual starfishes only valve molluscs, occurring on the gills, in the 10 cm in diameter. mantle cavity, and in the intestine of their In addition to describing four new fami­ hosts. Describing these taxa and creating a lies associated with echinoderms (Humes and more robust classification system represents Cressey, 1958b, 1959; Humes, 1974; Humes a real contribution to societal needs. Some and Boxshall, 1996), Arthur also contributed HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS 23 significantly to our knowledge of the Taeni­ (1987). The dirivultids are the dominant acanthidae. This is a very unusual family in group of copepods at most hydrothermal sites that it utilizes both invertebrates and verte­ in the eastern Pacific, in the Marianas Back- brates as hosts. The 91 species and 14 gen­ Arc Basin, and on the mid-Atlantic ridge, era of taeniacanthids recognized as valid by and, given that Tunnicliffe et ai. (1998) listed Dojiri and Cressey (1987) fall into two eco­ 443 hydrothermal vent species, dirivultids logical groups according to host preference. constitute over 10 percent of the faunal di­ Three genera, Echinosocius Humes and versity at hydrothermal sites. They have been Cressey, 1961, Echinirus Humes and Cressey, found in the washings of tubicolous poly­ 1961, and Clavisodalis Humes, 1970, are as­ chaetes, gastropods, bivalves, and in the gili sociates of sea urchins (Echinoidea). These chambers and around the oral region of three genera comprise a total of 14 species shrimps and crabs, as well as attached to the which occur only in the Indo-west Pacific and tentacular crown of vestimentiferan worms. Red Sea and inhabit the oesophagus of their Dirivultidae can be hyperabundant at vent echinoderm hosts. Arthur was author or co­ sites as illustrated by Humes (1990b) who author of 12 of these 14 known species. The found over 15,000 individuals of Stygiopon­ remaining 11 genera are widely distributed tius quadrispinosus Humes, 1987, in 210 ml parasites of marine fishes, both elasmo- of flocculent material collected at the Gorda branchs and teleosts. This family has been ex­ Ridge in the Eastern Pacific. Of particular sig­ tensively studied and was the subject of two nificance was Arthur’s discovery of eight revisions. Dojiri and Humes (1982) reviewed pairs in amplexus of this species which sub­ the taeniacanthids parasitic on sea urchins in sequently provided the first information on the southwestern Pacific and provided a key the functional morphology of the digeniculate to the known species of urchin parasites antennules and their role in precopulatory worldwide. Dojiri and Cressey (1987) revised mate guarding in siphonostomatoid copepods the entire family and provided a comprehen­ (Huyí and Boxshall, 1991). sive diagnosis of the family, a key to genera, A second family of siphonostomatoid cope­ and keys to the species parasitic on fishes. pods, the Ecbathyriontidae, was established by Humes in 1987, based on a single new H ydrothermal V e n t s , C old S e e p s , an d species found in the deep-sea hydrothermal THE DIRIVULTID EXPLOSION vent fauna on the Galapagos Rift. This fam­ The discovery of a specialized fauna as­ ily remains monotypic and its host group is sociated with the larger invertebrates living unknown. It is of particular phylogenetic in­ at hydrothermal vents and cold seeps pro­ terest in that, in addition to the large aes- vided Arthur Humes with a new challenge in thetasc derived from ancestral antennulary the early 1980s. The family Dirivultidae was segment XXI, the male of Ecbathyrion pro­ established in 1981 (Humes and Dojiri, 1981) lixicauda Humes, 1987, retains an excep­ based on a single species of a new genus, tionally high number of aesthetascs, those de­ Dirivultus Humes and Dojiri, 1981, collected rived from ancestral segments I to IV, VII, IX from a vestimentiferan worm off the Califor­ to XI, XIV, XVI, and XVIII (Huys and nia coast. Because of different in-press times, Boxshall, 1991). in the preceding year (1980) these authors Another hydrothermal vent family estab­ added a second monotypic genus, Ceutho­ lished by Humes, the Erebonasteridae, is the ecetes Humes and Dojiri, 1980, also associ­ only family of the order Poecilostomatoida in ated with a vestimentiferan host. During two which ventral copulatory pores are found that decades Arthur served as the main recipient are separate from the genital apertures on the for new copepod collections from hydrother­ dorsal surface of the female genital double­ mal vents and cold seeps worldwide which somite. It is also the only family in the order enabled him to publish extensively on the to retain a discrete mandibular palp (Humes, subject, including no less than 15 papers deal­ 1987). The family is characterised by these ing with the Dirivultidae. Due to his single- two extreme plesiomorphic states and this handed efforts this family has now grown to family, together with another family—the comprise twelve genera and 53 species, if we Fratiidae (cf. Ho et al., 1998)—of inverte­ include Fissuricola Humes, 1987, which was brate associates, will be of pivotal importance not originally placed in this family by Humes in the emerging concept of the Cyclopoida 24 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001 and Poecilostomatoida as a monophyletic turelle in Paris. His collections frequently taxon. comprised hundreds of paratypes of which he would normally deposit the vast majority in H is L eg a c y a major museum and the remainder in his own In this short appreciation we cannot do jus­ wonderfully arranged reference collection. In tice to the breadth and depth of Arthur addition, he deposited material of numerous Humes’s contributions to copepod taxonomy. other species, previously described by others We have attempted to pick out a few high­ but rediscovered during his own fieldwork. lights that seem, to us, to be of particular sig­ His meticulously curated personal collection nificance. We wish to stress that the accuracy was bequeathed to the National Museum of and detail of Arthur Humes’s taxonomic de­ Natural History and together with his earlier scriptions and those of his students, Roger donations constitutes the largest copepod col­ Cressey, Richard Gooding, Ju-shey Ho, and lection accumulated ever during an individ­ Masahiro Dojiri, are exemplary. He set a con­ ual’s lifetime. The new fourth edition of the sistently high standard in his illustrations, International Code of Zoological Nomencla­ which ensured he was providing data that ture has introduced a mandatory requirement would be of lasting value in biodiversity stud­ for the availability of new species-group taxa ies and phylogenetic analyses long into the fu­ published after 1999, i.e., a statement naming ture. His students continued this tradition, and the collection in which the type is or will be Arthur’s far-reaching influence will be of ben­ deposited (Art. 16.4.2). As with his methods efit to copepodology far into the 21st Century. Humes was well ahead of his time and en­ Arthur was a bom systematist who loved sured a lasting legacy. to synthesize information. His synoptic treat­ Arthur undoubtedly left many works un­ ments made it possible to become familiar finished. The fact that interest in the sipho- with the wide diversity of copepods associ­ nostomatoids did not equal interest in the poe­ ated with particular host groups such as the cilostoma toids is largely due to Arthur’s Holothuroidea (Humes, 1980), Actiniaria influence and contribution. On several occa­ (Humes, 1982a), Asteroidea (Humes, 1986b), sions he referred to the many collections of and Alcyonacea (Humes, 1990a), and to iden­ siphonostomatoid copepods, acquired during tify them without having to possess the scat­ his nearly four years of fieldwork in the 1960s tered original literature. Arthur always went and 1970s, and still awaiting study in his lab. to great effort to identify the host in order to Similarly, he felt that his collections of cope­ promote studies on host specificity and geo­ pods associated with ascidian hosts had not graphical distribution. He believed that yet received the attention they deserved. Dur­ knowledge of the exact host name could po­ ing the Fifth International Copepod Confer­ tentially shortcut the identification process ence in Baltimore in 1993, the last of its kind significantly, and therefore he regularly up­ he attended, Arthur voiced his intention to dated records on host-copepod associations in continue describing copepods until the turn a synthetic cross-referenced format. of the millennium. In view of Arthur’s sus­ From the start of his career Humes recog­ tained output in recent years, which showed nized that name-bearing types are the sole in­ no sign of decline, and the prospect of more ternational standards of reference and should research time following his retirement as Ed­ be deposited in an institution that maintains itor ofJournal of Crustacean Biology, it is a research collection with proper facilities for unlikely that this turning point would have preserving them and making them available marked the end of his extraordinarily pro­ for study to others. More than just being a ductive career. tireless collector, Arthur deposited type ma­ In recognition of his monumental work on terial of each of the 700+ species he described copepod systematics Arthur Humes was a re­ in international museums such as the National cipient of the Research Excellence Award Museum of Natural History in Washington, given by The Crustacean Society. Uniquely, the Natural History Museum in London, the the Board of The Crustacean Society voted to Rijksmuseum van Natuurlijke Historie in Lei­ bestow this honour on Arthur Humes posthu­ den, the Zoölogisch Museum in Amsterdam mously, since the nomination process had and the Muséum National d’Histoire Na­ been completed before his death. At the win­ HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS 25 ter meeting of The Crustacean Society in At­ . 1956. Pholetiscus rectiseta n. sp. des cavités lanta in January 2000, the award was pre­ branchiales d'un crabe à Madagascar (Copepoda, Harpacticoida).—Mémoires de l'Institut Scientifique sented by the President, Dr. Joel Martin, to de Madagascar, sér. A, 11: 79-84. Dr. Charles Derby who accepted it on behalf . 1957. Two new caligoid copepods from Egyp­ of Arthur Humes. This award has since been tian fishes.—Journal of Parasitology 43: 201-208. renamed the Arthur G. Humes Award for Re­ . 1959. Copépodes parasites de Mollusques à Madagascar.—Mémoires de l'Institut Scientifique de search Excellence. Madagascar, sér. F, 2: 285-342. It is difficult to imagine the held of asso­ . 1960a. The harpacticoid copepod Sacodiscus ciated copepods without Arthur Humes. In (=UnicaIteutha) ovalis (C. B. Wilson, 1944) and its these times of decreasing support for, and in­ copepodid stages.—Crustaceana 1: 279-294. terest in, taxonomic research, the shear vol­ . 1960b. New copepods front ntadreporarian corals.—Kieler Meeresforschungen 16: 229-235. ume of his work may never be surpassed. To . 1962. Eight new species ofXarifia (Copepoda, paraphrase the title of his own past-presi­ Cyclopoida), parasites of corals in Madagascar.—Bul­ dential address presented at the Annual Meet­ letin of the Museum of Comparative Zoology (Harvard ing of the American Microscopical Society in University) 128: 35-63. Denver in December 1984 (Humes, 1985b), . 1964. Harpacticus pulex, a new species of cope­ pod front the skin of a porpoise and a manatee in any biographical sketch of the man, who Florida.—Bulletin of Marine Science of the Gulf and never lost his fascination with his beloved an­ Caribbean 14: 517-528. imals, could simply be condensed as “Arthur . 1965. Sciaenophilus inopinus Huntes, 1957, a Humes and Copepods: A Success Story.” synonym of Sciaenophilus pharaonis (Nordnrann, 1832) contb. n. (Copepoda: Caligidae).—Journal of Parasitology 51: 1009, 1010. L itera tur e C ited . 1966. New species ofMacrochiron (Copepoda, Boxshall, G. A., and A. G. Humes. 1988. A new genus Cyclopoida) associated with hydroids in Madagascar.— of Lichomolgidae (Copepoda: Poecilostomatoida) as­ Beaufortia 14: 5-28. sociated with a phoronid in Hong Kong.—Bulletin of . 1967. Vahinius petax n. gen., n. sp., a cyclopoid the British Museum (Natural History) 54: 301-307. copepod parasitic in an antipatharian coelenterate in Dojiri, M., and R. F. Cressey. 1987. Revision of the Tae­ Madagascar.—Crustaceana 12: 233-242. niacanthidae (Copepoda: Poecilostomatoida) parasitic . 1969. A cyclopoid copepod.Sewellochiron fi­ on fishes and sea urchins.—Smithsonian Contributions delis n. gen., n. sp., associated with a medusa in Puerto to Zoology 447: i-iv, 1-250. Rico.—Beaufortia 16: 171-183. ------, and A. G. Humes. 1982. Copepods (Poe­ . 1970. Paramacrochiron japonicum n. sp., a cy­ cilostomatoida: Taeniacanthidae) from sea urchins clopoid copepod associated with a medusa in Japan.— (Echinoidea) in the southeast Pacific.—Zoological Publications of the Seto Marine Biological Laboratory Journal of the Linnean Society 74: 381-436. 18: 223-232. Gooding, R. U., and A. G. Humes. 1963. External . 1971. Cyclopoid copepods (Stellicomitidae) anatomy of the female Haemobaphes cyclopterina, a parasitic on sea stars front Madagascar and Eniwetok copepod parasite of marine fishes.—Journal of Para­ Atoll.—Journal of Parasitology 57: 1330-1343. sitology 40: 663-677. . 1973a. Tychidion guyanense n. gen., n. sp. (Co­ Ho, J.-s., M. Conradi, and P. J. López-González. 1998. pepoda, Cyclopoida) associated with an annelid off A new family of cyclopoid copepods (Fratiidae) sym­ Guyana.—Zoologische Mededelingen 46: 189-196. biotic in the ascidian ( Clavelina dellavallei) from Cádiz, . 1973b. Cyclopoid copepods associated with the Spain.—Journal of Zoology, London 246: 39-48. ophiuroid Astroboa nuda in Madagascar.—Beaufortia ------, and I.-H. Kim. 1992. A new genus of poe- 21: 25-35. cilostome copepod of the family Myicolidae parasitic . 1974. New cyclopoid copepods associated with in a commercial clam from Malaysia.—Journal of Nat­ an abyssal holothurian in the eastern North Atlantic.— ural History 26: 303-309. Journal of Natural History 8: 101-117. ------, and W. Yoosukh. 1994.Ostrincola humesi sp. . 1977. Cyclopoid copepods (Lichomolgidae) as­ nov., a myicolid copepod (Poecilostomatoida) parasitic sociated with hydroids in the tropical western Pacific in rock oysters cultured in the Gulf of Thailand.—Hy- Ocean.—Pacific Science 31: 335-353. drobiologia 288: 151-155. . 1979. Poecilostonre copepods associated with , and G.-X. Zheng. 1994. Ostrincola koe (Co­ antipatharian coelenterates in the Moluccas.—Beau­ pepoda, Myicolidae) and mass mortality of cultured fortia 28: 113-120. hard clam ( Meretrix meretrix) in China.—Hydrobiolo- . 1980. A review of the copepods associated with gia 284: 169-173. holothurians, including new species from the Indo-Pa- Humes, A. G. 1941. A new harpacticoid copepod from cific.— Beaufortia 30: 31-123. the gili chambers of a marsh crab.—Proceedings of . 1982a. A review of Copepoda associated with the United States National Museum 90(3110): 379-386. sea anemones and anemone-like forms (Cnidaria, An­ ------. 1955. The postembryonic developmental stages thozoa).—Transactions of the American Philosophical of a fresh-water calanoid copepod. Epischura massa- Society 72: 1-120. chusettsetisis Pearse.—Journal of Morphology 96: . 1982b. Copepoda (Poecilostomatoida, Li­ 441-471. chomolgidae) associated with alcyonacean genus Sar- 26 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001

eophyton in the Indo-Pacific.—Publications of the Seto ------, and A. De Maria. 1969. The cyclopoid copepod Marine Biological Laboratory 27: 25-76. genus Macrochiron from hydroids in Madagascar.— . 1985a. A review of the Xarifiidae (Copepoda, Beaufortia 16: 137-155. Poecilostomatoida), parasites of scleractinan corals in the ------, and M. Dojiri. 1980. A siphonostome copepod Indo-Pacific.—Bulletin of Marine Science 36: 467-632. associated with a vestimentiferan from the Galapagos . 1985b. Cnidarians and copepods: a success rift and the East Pacific rise.—Proceedings of the Bi­ story.—Transactions of the American Microscopical ological Society of Washington 93: 697-707. Society 104: 313-320. ------, a n d ------. 1981. A new siphonostome family . 1986a. Myicola metisiensis (Copepoda: Poe­ (Copepoda) associated with a vestimentiferan in deep cilostomatoida), a parasite of the bivalveMya arenaria water off California.—Pacific Science 34: 143-151. in eastern Canada, redefinition of the Myicolidae, and ------, a n d ------. 1982. Xarifiidae (Copepoda) par­ diagnosis of the Anthessiidae n. fani.—Canadian Jour- asitic in Indo-Pacific scleractinian corals.—Beaufortia nal~of Zoology 64: 1021-1033. 32: 139-228. . 1986b. Synopsis of copepods associated with ------, a n d ------. 1983. Copepoda (Xarifiidae) par­ asteroid echinoderms, including new species from the asitic in scleractinian corals from the Indo-Pacific.— Moluccas.—Journal of Natural History 20: 981-1020. Journal of Natural History 17: 257-307. . 1986c. Copepodids and adults of Leptinogaster ------, and R. U. Gooding. 1964. A method for study­ major (Williams, 1907), a poecilostomatoid copepod ing the external anatomy of copepods.—Crustaceana 6: living in Mya arenaria L. and other marine bivalve 238-240. mollusks.—Fishery Bulletin, U.S. 85: 227-245. ------, and J. F. Grassle. 1979.Serpulidicola josephel­ . 1987. Copepoda from deep-sea hydrothermal lae sp. nov. (Copepoda, Cyclopoida) from a deep-wa­ vents.—Bulletin of Marine Science 41: 645-788. ter polychaete west of Ireland.—Crustaceana 36: . 1990a. Synopsis of lichomolgid copepods (Poe­ 309-315. cilostomatoida) associated with soft corals (Alcy­ ------, and G. Hendler. (In press.) Biology and taxon­ onacea) in the tropical Indo-Pacific.—Zoologische Ver­ omy of species ofOphiopsyllus and Pseudanthessius handelingen 266: 1-201. (Copepoda) associated with brittle stars (Ophiuroidea) . 1990b.Aphotopontius probolus, sp. nov., and in Belize.—Bulletin of Marine Science. records of other siphonostomatoid copepods from deep------, andJ.-s. Ho. 1967. New cyclopoid copepods as­ sea vents in the eastern Pacific.—Scientia Marina 54: sociated with the coral Psammocora contigua (Esper) 145-154. in Madagascar.—Proceedings of the United States Na­ . 1994. How many copepods? Pp. 1-7in F. D. tional Museum 122: 1-32. Ferrari and B. P. Bradley, eds. Ecology and Morphol­ ------, a n d ------. 1968. Xarifiid copepods (Cy­ ogy of Copepods. Proceedings of the Fifth International clopoida) parasitic in corals in Madagascar.—Bulletin Conference on Copepoda.—Hydrobiologia 292/293. of the Museum of Comparative Zoology (Harvard Uni­ . 1996. Orecturus amplus, a new species (Co­ versity) 136: 415-459. pepoda: Siphonostomatoida: Asterocheridae) from an ------, a n d ------. 1969a. The genusSunaristes (Co­ alcyonacean in New Caledonia.—Proceedings of the pepoda, Harpacticoida) associated with hermit crabs Biological Society of Washington 109: 112-117. in the western Indian Ocean.—Crustaceana 17: 1-18. . 1997.Pseudanthessius newmanae, new species ------, a n d ------. 1969b. Harpacticoid copepods of (Copepoda: Poecilostomatoida: Pseudanthessiidae) the genera Porcellidium and Paraidya associated with from marine turbellarians in Australia.—Memoirs of hermit crabs in Madagascar and Mauritius.—Crus­ the Queensland Museum 42: 227-231. taceana 17: 113-130. . 1999a. Copepoda (Siphonostomatoida) from------, and D. C. Rosenfield. 1960. Anchistrotos occi­ Pacific hydrothermal vents and cold seeps, including dentalis C. B. Wilson, 1924 (Crustacea, Copepoda), a Dirivultus spinigulatus sp. nov. in Papua New parasite of the orange filefish.— Crustaceana 1: 179-187. Guinea.—Journal of the Marine Biological Association ------, and M. Segonzac. 1998. Copepoda from deep- of the United Kingdom 79: 1053-1060. sea hydrothermal sites and cold seeps: description of . 1999b. Copepoda (Cyclopinidae and Misophri­ a new species of Aphotopontius from the East Pacific idae) from a deep-sea hydrothermal site in the north­ Rise and general distribution.—Cahiers de Biologie eastern Pacific.—Journal of Natural History 33: 961- Marine 39: 51-62. 978. ------, and W. L. Smith. 1974 Ridgewayia fosshageni n. , and G. A. Boxshall. 1988. Poecilostome copepods sp. (Copepoda: Calanoida) associated with an actiniar- associated with bivalve molluscs and a brachiopod at ian in Panama, with observations on the nature of the Hong Kong.—Journal of Natural History 22: 537-544. association.—Caribbean Journal of Science 14: 125-139. , a n d ------. 1996. A revision of the lichomol­------, and J. H. Stock. 1972. Preliminary notes on a goid complex (Copepoda: Poecilostomatoida), with the revision of the Lichomolgidae, cyclopoid copepods recognition of six new families.—Journal of Natural mainly associated with marine invertebrates.—Bulletin History 30: 175-227. Zoologisch Museum, Universiteit van Amsterdam 2: , and R. F Cressey. 1958a. Copepod parasites of 121-133. mollusks in West Africa.—Bulletin de l'Institut , and . 1973. A revision of the family Li­ Français d'Afrique Noire 20, ser. A, no. 3: 921-942. chomolgidae Kossmann, 1877, cyclopoid copepods , a n d ------. 1958b. A new family containing mainly associated with marine invertebrates.—Smith­ two new genera of cyclopoid copepods parasitic on sonian Contributions to Zoology 127: i-v, 1-368. starfishes.—Journal of Parasitology 44: 395-408. Huys, R., and G. A. Boxshall. 1991. Copepod Evolution.— , a n d ------. 1959. A new family and genus of The Ray Society, London, England. 159. Pp. 1-468. cyclopoid copepods parasitic on a holothurian.—Jour­ Sebastian, M. J., and N. Krishna Pillai. 1973. Humesiella nal of Parasitology 45: 209-216. corallicola n. g., n. sp., a cyclopoid copepod associ­ HUYS AND BOXSHALL! CONTRIBUTION OF ARTHUR HUMES TO COPEPOD SYSTEMATICS 27

ated with coral on the south east coast of India.—Hy- drothermal vent fauna.—Advances in Marine Biology drobiologia 42: 143-152. 34: 353-442. Stock, J. H., and G. Kleeton. 1963. Copépodes associés Yamaguti, S. 1936. Parasitic copepods from mollusks of aux invertébrés des côtes du Roussillon. 2. Lichomol­ Japan, I.—Japanese Journal of Zoology 7: 113-127. gidae ecto-associés d'Octocoralliaires.—Vie et Milieu 14(2): 245-261. Tunnicliffe, V., A. G. McArthur, and D. McHugh. 1998. R e c eiv e d : 28 September 2000. A biogeographical perspective of the deep-sea hy­ A c c ept e d : 29 September 2000.