JOURNAL OF CRUSTACEAN BIOLOGY, 21(1): 231–242, 2001
REVISION OF THE ENDEMIC AUSTRALIAN GENUS SANCHO
(AMPHIPODA, EUSIRIDAE)
J. K. Lowry and J. L. Barnard
Division of Invertebrate Zoology, Australian Museum, 6 College Street, Sydney, 2010,
NSW, Australia (e-mail: [email protected])
A B S T R A C T
The endemic Australian eusirid amphipod genus Sancho Stebbing, 1897, is revised based on new
material. A neotype is established for Sancho platynotus Stebbing, 1897, which is redescribed, and
the new species S. kuiteri is described. Members of Sancho are considered to be obligate commensals
of sponges. Their depressed body form and entire telson may be an adaptation to their commensal
life-style, but their cantilevered second gnathopods may indicate a phylogenetic relationship with
the Eusirus-like members of the Eusiridae.
In 1897 Stebbing described Sancho
platynotus from Port Jackson, Australia. The
description and illustrations were brief, there
was no habitat information, and the taxon was
not placed in a family. Stebbing later (1906)
placed Sancho in the Calliopiidae. So did J.
L. Barnard (1969), who carefully considered
the close relationship between calliopiids and
eusirids and commented specifically on the
eusirid-like gnathopods of Sancho. Barnard
and Karaman (1991) amalgamated the Calliopiidae and Eusiridae and placed Sancho in
the Eusiridae. They considered it to be a peculiar genus in the family mainly because of
its depressed body and urosome.
Before J. L. Barnard died in 1991, we were
studying a new taxon from Port Jackson
which appeared to be closely related to Sancho platynotus. Barnard was never convinced
that Sancho belonged in the Eusiridae (= Calliopiidae) and was in favour of establishing
a new family for S. platynotus and the new
taxon. A major problem for us was that Sancho platynotus had never been collected
again, and the original type material, placed
in the Copenhagen Museum, was lost. Finally,
after 101 years, Megan Huggett (Honours student, University of New South Wales) collected specimens of S. platynotus associated
with the sponge Holopsamma laminaefavosa
Carter, 1885, during a survey of amphipods
associated with sponges in Botany Bay. Study
of this new material from Botany Bay and the
new taxon from Port Jackson has resolved
many of the problems associated with the
original description of Sancho. In this paper
we redescribe S. platynotus and describe the
new species Sancho kuiteri. Because the types
of S. platynotus are lost and S. platynotus is
the type species of the genus, we are establishing a neotype. Sancho platynotus is a fragile species, and there were no complete specimens in Huggett’s collections. The neotype
is therefore a nearly complete female.
Species of Sancho appear to be eusirid-like
amphipods which have become morphologically adapted to living on sponges. The main
differences between Sancho and other eusirid
taxa are the shape of the head, the sexually
dimorphic second gnathopods, and the depressed body and urosome. The only seemingly similar taxon is the poorly described
Choroes incisus Stebbing, 1888, which also
has a depressed body and urosome, but the
shape of the head and the dimorphic nature
of the gnathopods are unknown. According to
K. H. Barnard (1931), who had many specimens (males and females) available for study,
C. incisus also lives on sponges. Although J.
L. Barnard was in favor of creating a new
family for Sancho, it now seems better to retain Sancho in the Eusiridae until a thorough
revision of the family and its sister taxa is
achieved.
Material used in this study is lodged in the
Australian Museum, Sydney (AM). The following abbreviations are used on the figures:
A, antenna; C, coxa; EP, epimeron; G,
gnathopod; H, head; LL, lower lip; MD,
mandible; MP, maxilliped; MPOP, maxilliped
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
Fig. 1. Sancho kuiteri, new species, holotype m, 4.1 mm, AM P40664, Botany Bay, New South Wales, Australia.
outer plate; MPIP, maxilliped inner plate;
MX, maxilla; P, pereiopod; PL, pleon; T, telson; U, uropod; UL, upper lip; UR, urosomite; l, left; r, right.
Eusiridae
Sancho Stebbing, 1897
Sancho Stebbing, 1897: 42.—Stebbing, 1906: 288.—J. L.
Barnard, 1969: 179.—Barnard and Karaman, 1991:
339.
Type Species.—Sancho platynotus Stebbing,
1897, by monotypy.
Diagnosis.—Body strongly depressed, with
urosome flexed under pleon. Antenna 1 accessory flagellum present, 1-articulate. Mandible molar triturative. Lower lip without inner lobes. Maxilla 1 inner plate with two apical setae. Maxilla 2 with outer plate slightly
to strongly broader than inner plate; inner
plate without oblique setal row. Maxilliped
with outer plate not enlarged. Gnathopod 2
sexually dimorphic, weakly to strongly cantilevered in male. Uropod 3 with outer ramus
one-third to three-fourths length of inner ramus. Telson entire or emarginate.
Species Composition.—Sancho kuiteri, new
species; S. platynotus Stebbing, 1897.
Remarks.—Sancho is apparently endemic to
southeastern Australia. Species in the genus
appear to be obligate commensals of sponges,
and as an adaptation to this life-style they
have dorsoventrally flattened bodies and depressed urosomes similar to other commensals such as iciliids and maxillipiids, which
are also endemic to the Indo-West Pacific area.
Species of Sancho have notched telsons
similar to the calliopiid-like members of the
Eusiridae, but Sancho platynotus, in particular, has cantilevered gnathopods similar to the
eusirid-like members of the family. If these
cantilevered gnathopods are homologous then
Sancho and Eusirus may have a common ancestor. It is interesting that Eusirus antarcticus Thomson, 1880 (sensu stricto), which
lives in southern New Zealand, has a heavy
body and broad legs like species of Sancho.
Sancho kuiteri, new species
Figs. 1–5
Type Material.—Holotype, m, 4.1 mm, AM P40664;
Paratypes: f, 3.6 mm, P40665; m, 3.9 mm, P40666; f,
LOWRY AND BARNARD: REVISION OF AMPHIPOD GENUS SANCHO
233
Fig. 2. Sancho kuiteri, new species, holotype m, 4.1 mm, AM P40664, f, 3.6 mm, AM P40665, Botany Bay, New
South Wales, Australia.
4.1 mm, P40667; juvenile, P40668; m, P40670; 133 specimens, P40669; off Green Point, Watsons Bay, Port Jackson, New South Wales, Australia, 33°50.6′S 151°16.55′E,
living on the sponge Holopsamma laminaefavosa (AM
Z5090), 10 m, R. Kuiter, 12 September 1981.
ovigerous ff, AM P58135; Bare Island, Botany Bay, New
South Wales, Australia, 33°59.53′S 151°13.83′E, living
on the sponge Holopsamma laminaefavosa, 12 m, M. J.
Huggett and R. De Nys, 13 August 1998.
Additional Material.—4 mm, 12 ff and juveniles, AM
P57935; 1 m, 2 juveniles, AM P57938; 2 ff, AM P58133;
7 mm, 12 ff and juveniles, AM P58134; 5 mm, 14 mostly
Description.—Based on holotype male, 4.1
mm. Head and body white. Head longer than
deep; lateral cephalic lobe poorly developed;
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
Fig. 3. Sancho kuiteri, new species, holotype m, 4.1 mm, AM P40664, f, 3.6 mm, AM P40665, Botany Bay, New
South Wales, Australia. Scale bars = 0.1 mm.
anteroventral margin rounded, anteroventral
corner rounded; rostrum present, moderate
length; eyes medium-sized, oval. Antenna 1
long, about 0.6 × body, 1.5 × as long as antenna 2; peduncular article 1 longer than (2
×) article 2; article 2 longer than (1.6 ×) article 3; article 3 shorter than (0.3 ×) article
1; accessory flagellum present, minute, 1-articulate; flagellum 36-articulate, with sparse
setae along posterior margin of most articles;
calceoli absent. Antenna 2 peduncular article
3 short, length 0.8 × breadth, without flange
on anterolateral margin; article 4 long, 1.7 ×
breadth; article 5 long, 4.5 × breadth; flagellum shorter than (0.8 ×) peduncle, 21-articulate; calceoli absent.
Upper lip separate, not produced.
Mandibles incisors symmetrical, left and right
with 4 serrations; lacinia mobilis asymmetri-
cal, left with 4 serrations, right with 2 serrations; accessory setal row, left with 2 serrate
setae, right with 1 serrate seta; molars columnar, fully triturating, each with 4 long pappose setae; palp well developed, article 2
long, 1 × article 3; article 3 long. Lower lip
inner lobes absent. Maxilla 1 inner plate
small, subovate, with 2 apical setae; outer
plate, with 11 strong setal-teeth; palp 2-articulate with about 8 apical or apicomedial setae. Maxilla 2 inner and outer plates broad;
inner plate with apical slender setae, without
medial slender setae, without oblique row of
slender setae; outer plate with apical slender
setae, without medial slender setae. Maxilliped inner plate large, nearly reaching end of
merus, subrectangular, without nodular setae,
without apicomedial setae, oblique setal row
with 14 long plumose setae; outer plate sub-
LOWRY AND BARNARD: REVISION OF AMPHIPOD GENUS SANCHO
235
Fig. 4. Sancho kuiteri, new species, holotype m, 4.1 mm, AM P40664, f, 3.6 mm, AM P40665, Botany Bay, New
South Wales, Australia. Scale bars = 0.1 mm.
ovate, reaching to end of merus, with 6 pappose setae along apical margin, without robust setae along medial margin, without submarginal setae; palp large, 4-articulate; carpus broad, length 1.1 × propodus, with setose
inner margin; propodus long, length 3.1 ×
breadth, without setose inner margin; dactylus cone-like, unguis absent.
Pereion.—Gnathopod 1 not sexually dimorphic, subchelate, smaller than gnathopod
2; coxa deeper than wide, anteroventral mar-
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
Fig. 5. Sancho kuiteri, new species, holotype m, 4.1 mm, AM P40664, f, 3.6 mm, AM P40665, Botany Bay, New
South Wales, Australia.
gin slightly produced, subquadrate; basis
without anterodistal lobe; carpus subrectangular, long, length 3 × breadth, subequal to
(1.1 ×) propodus, posterior margin without
lobe; propodus narrow, length 2 × breadth,
palm transverse, straight, without seta near
inner base of dactylus, with several posterodistal robust setae; dactylus slender, falcate. Gnathopod 2 sexually dimorphic, subchelate; coxa deeper than wide; basis without
anterodistal lobe; carpus/propodus cantilevered on narrow hinge; carpus subquadrate, short, length 0.9 × breadth, shorter
than (0.3 ×) propodus, not produced; propodus broad, length 1.7 × breadth, slightly produced posterodistally, palm acute, straight,
crenate, without seta near inner base of dactylus, with several posterodistal robust setae;
dactylus slender, falcate, inner margin
smooth.
Pereiopod 3 coxa deeper than wide; merus
long, length about 1.5 × breadth, anterior margin slightly expanded; carpus long, about 1.5
× breadth; propodus length nearly 3 ×
breadth, with 2 setae along posterior margin.
Pereiopod 4 coxa as wide as deep, with a subquadrate posteroventral lobe. Pereiopod 5
coxa wider than deep, equilobate; basis
strongly expanded posteriorly, with long slender setae along anterior margin; merus short,
length about 1 × breadth; carpus long, subrectangular, length about 1.7 × breadth, covered in minute denticles; propodus subrectangular, length about 2.5 × breadth, with 3
slender setae and 5 robust setae along anterior margin, not expanded distally, with about
20 setae along posterior margin; dactylus
short, slightly curved, inner margin smooth.
Pereiopod 6 coxa as wide as deep, with welldeveloped posteroventral lobe; basis expanded, with slender setae along anterior margin, posterior margin slightly rounded, crenate; merus short, length 1 × breadth; carpus
long, length 1.2 × breadth; propodus subrectangular, length about 2.8 × breadth, covered
in minute denticles, without 4 slender setae
along anterior margin, without distal robust
setae; dactylus slightly curved, with subterminal seta on posterior margin. Pereio
pod 7 coxa wider than deep, without pos-
LOWRY AND BARNARD: REVISION OF AMPHIPOD GENUS SANCHO
teroventral lobe; basis with posterior margin
straight, crenate; carpus and propodus covered in minute denticles.
Pleon.—Epimeron 3 posteroventral corner
subquadrate, with minute acute spine. Uropod 1 peduncle with 8 dorsolateral robust setae and 6 dorsomedial robust setae; outer ramus shorter than inner ramus; outer ramus
with 2 lateral, 1 medial, and 2 apical robust
setae; inner ramus with 4 medial, 2 lateral,
and 2 apical robust setae. Uropod 2 peduncle with 5 dorsolateral robust setae, and 1 dorsomedial robust seta; outer ramus shorter than
inner; with 3 lateral, 2 medial, and 2 apical
robust setae; inner ramus with 4 medial, 6 lateral, and 2 apical robust setae. Uropod 3 peduncle short, without dorsolateral or dorsomedial robust setae; outer ramus shorter than
inner; outer ramus with 2 lateral and 2 medial robust setae; inner ramus with 4 medial
and 2 lateral robust setae. Telson notched, as
long as broad, length 1 × breadth, without
dorsal robust setae, with sparse dorsal slender setae, distal margins rounded, without apical penicillate setae, with 1 apical slender seta
on each lobe, without apical robust setae.
Female.—Based on paratype female.
Gnathopod 1 subequal to gnathopod 2; carpus long, length 3 × breadth, longer than (1.2
×) propodus; palm slightly obtuse, convex.
Gnathopod 2 carpus subrectangular, very
long, length 4 × breadth, longer than (1.2 ×)
propodus; propodus narrow, length 3.1 ×
breadth, not produced posterodistally, palm
slightly obtuse, straight, smooth, inner margin setose. Pereiopod 5 coxa with posteroventral lobe.
Etymology.—Named for the collector, Rudi
Kuiter, in recognition of his contribution to
the knowledge of Australian marine animals.
Habitat.—Living mainly in association with
the sponge Holopsamma laminaefavosa and
occasionally with the sponge Phoriospongia
cf. kirki (Bowerbank, 1841).
Remarks.—Sancho kuiteri and S. platynotus
are closely related sister taxa which live together in the same sponge gardens, although
S. kuiteri appears to be restricted to fewer
species of sponges than S. platynotus. Sancho
kuiteri was collected almost exclusively on
Holopsamma laminaefavosa whereas S.
237
platynotus was collected on eight species of
sponges and only rarely on Holopsamma laminaefavosa.
Both species show remarkable similarities
in the depressed body form, the unusual
gnathopods, unusual coxae 4, and pereiopods
5 to 7. The gnathopods, for instance, show a
similar development pattern, which only differs in adult males, where the carpus becomes
strongly cantilevered and the propodus becomes ovate in Sancho platynotus. Apparently neither species has calceoli.
Distribution.—Port Jackson and Botany Bay,
New South Wales, Australia.
Sancho platynotus Stebbing, 1897
Figs. 6–9
Sancho platynotus Stebbing, 1897: 42, pl. 9A.—Stebbing,
1906: 288, figs. 72, 73.
Type Material.—Neotype f, 3.9 mm, AM P58144. Bare
Island, Botany Bay, New South Wales, Australia,
33°59.53′S 151°13.83′E, 12 m, sponge, M. J. Huggett and
R. De Nys, 13 August 1998.
Additional Material Examined.—1 m, 4.5 mm, AM
P58136 on sponge; 1 f, 3.5 mm, AM P58143 on sponge;
1 m, 3.5 mm, AM P58146 on sponge; 1 f, 3.5 mm, AM
P58147 on sponge; 2 specimens AM P 57910 on sponge;
1 specimen AM P 57933 on the sponge Antho
(Isopenectya) chartacea (Whitelegge, 1907); 1 specimen
AM P 57934 on the sponge Callyspongia sp.; 2 specimens P 57936 on the finger sponge Phorbas sp.; 1 specimen AM P57937 on the sponge Mycale (Carmia) sp.; 2
specimens AM P57938 on the sponge Phoriospongia cf.
kirki; Bare Island, Botany Bay, New South Wales, Australia 33°59.53′S 151°13.83′E, 12 m, sponge, M. J.
Huggett and R. De Nys, 13 August 1998.
Description.—Based on the neotype female,
3.9 mm, P58144, plus female, 3.5 mm
P58143 and female, 3.5 mm P58147. Head
and body white. Head as long as deep; lateral cephalic lobe poorly developed; anteroventral margin rounded, anteroventral
corner rounded; rostrum present, short; eyes
medium-sized, round. Antenna 1 medium
length, about 0.3 × body; peduncular article
1 longer than (1.6 ×) article 2; article 2 longer
than (2 ×) article 3; article 3 shorter than (0.3
×) article 1; accessory flagellum present,
minute, 1-articulate; flagellum about 45-articulate; with sparse setae along posterior
margin of most articles; calceoli absent. Antenna 2 about 35-articulate; calceoli absent.
Upper lip separate. Mandibles incisors
symmetrical, left and right each with 5 ser-
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
Fig. 6. Sancho platynotus Stebbing, 1897, m, 4.5 mm, AM P58136, neotype f, AM P58144, urosome. Botany Bay,
New South Wales, Australia.
rations; lacinia mobilis asymmetrical, left
with 3 serrations, right with 4 serrations; accessory setal row absent; molars columnar,
fully triturating, with 5 long pappose setae on
left side only; palp well developed, article 2
long, 1 × article 3; article 3 long. Lower lip
inner lobes absent. Maxilla 1 inner plate
longer than broad, subrectangular, with 2 apical setae; outer plate with 11 strong setalteeth; palp 2-articulate, with 8 apical or apicomedial setae. Maxilla 2 inner plate narrow,
outer plate broader; inner plate with apical
slender setae, without medial slender setae,
without oblique row of slender setae; outer
plate with apical slender setae, without medial slender setae. Maxilliped inner plate
medium size, reaching about halfway along
merus, subrectangular, without nodular setae,
without apicomedial setae, oblique setal row
with about 20 long plumose setae; outer plate
subovate, reaching to end of merus, with 3
pappose setae along apical margin, with 10
robust setae along medial margin, without
submarginal setae; palp large, 4-articulate;
carpus broad, length 1.1 × propodus, with setose inner margin; propodus long, length 3.1
× broad, without setose inner margin; dactylus cone-like, unguis absent.
Pereion.—Gnathopod 1 not sexually dimorphic, slightly chelate, subequal to gnatho-
pod 2; coxa deeper than wide, anteroventral
margin produced, rounded; basis without anterodistal lobe; carpus subrectangular, long,
length 3 × breadth, longer than (1.4 ×) propodus, posterior margin without lobe; propodus
narrow, length 1.8 × breadth, palm slightly
obtuse, convex, without seta near inner base
of dactylus, with several posterodistal robust
setae; dactylus slender, falcate. Gnathopod 2
sexually dimorphic, chelate; coxa deeper than
wide; basis without anterodistal lobe; carpus/propodus cantilevered on narrow hinge;
carpus subrectangular, long, length 3.2 ×
breadth, longer than (1.2 ×) propodus, not
produced; propodus narrow, length 2.9 ×
breadth, not produced posterodistally, palm
slightly obtuse, slightly convex, smooth,
without seta near inner base of dactylus, with
several posterodistal robust setae; dactylus
slender, falcate, inner margin with long setae.
Pereiopod 3 coxa deeper than wide; merus
long, length about 1.5 × breadth, anterior margin slightly expanded; carpus long, length
about 2 × breadth; propodus length about 2.5
× breadth, with 2 setae along posterior margin. Pereiopod 4 coxa slightly deeper than
wide, with a subquadrate posteroventral lobe;
merus long, length about 1.8 × breadth, anterior margin slightly expanded. Pereiopod 5
coxa wider than deep, with posteroventral
LOWRY AND BARNARD: REVISION OF AMPHIPOD GENUS SANCHO
239
Fig. 7. Sancho platynotus Stebbing, 1897, neotype f, 3.9 mm, AM P58144, f, 3.5 mm, AM P58143, A1, Botany Bay,
New South Wales, Australia. Scale bars = 0.1 mm.
lobe; basis strongly expanded posteriorly,
with robust setae along anterior margin;
merus short, length about 1 × breadth; carpus long, subrectangular, length about 1.5 ×
breadth; propodus subrectangular, length
about 3 × breadth, with 3 rows of slender setae and 5 robust setae along anterior margin,
not expanded distally, with about 5 rows of
setae along posterior margin; dactylus short,
slightly curved, inner margin smooth. Pereio-
pod 6 coxa as wide as deep, with well-developed posteroventral lobe; basis expanded,
with slender setae along anterior margin, posterior margin slightly rounded, smooth; merus
long, length 1.2 × breadth; carpus long, length
3 × breadth; propodus subrectangular, length
about 3 × breadth, with 3 rows of slender setae along anterior margin, without distal robust setae; dactylus slightly curved, with subterminal seta on posterior margin. Pereiopod
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
Fig. 8. Sancho platynotus Stebbing, 1897, neotype f, 3.9 mm, AM P58144, m, 3.5 mm, AM P58146, G1, 2, Botany
Bay, New South Wales, Australia. Scale bars = 0.2 mm.
7 coxa wider than deep, with broad, shallow
posteroventral lobe; basis with posterior margin straight, smooth.
Pleon.—Epimeron 3 posteroventral corner
acute with minute notch. Uropod 1 peduncle
with 6 dorsolateral robust setae and 5 dorsomedial robust setae; outer ramus shorter than
inner ramus; outer ramus with 3 lateral and
2 apical robust setae; inner ramus with 1 medial, 4 lateral, and 4 apical robust setae. Uropod 2 peduncle with 5 dorsolateral robust setae and 1 dorsomedial robust seta; outer ramus shorter than inner; outer ramus with 4
lateral, 4 medial and 2 apical robust setae; inner ramus with 4 medial, 6 lateral and 2 apical robust setae. Uropod 3 peduncle short
LOWRY AND BARNARD: REVISION OF AMPHIPOD GENUS SANCHO
241
Fig. 9. Sancho platynotus Stebbing, 1897, neotype f, 3.9 mm, AM P58144, m, 3.5 mm, AM P58146, *f, 3.5 mm, AM
P58147, Botany Bay, New South Wales, Australia. Scale bars for P1–P7 = 0.2 mm, for U1–U3 and T = 0.1 mm.
without dorsolateral and robust setae; outer
ramus shorter than inner; outer ramus with 4
lateral and 2 medial; inner ramus with 7 medial and 5 lateral robust setae. Telson notched,
as long as broad, length about 1 × breadth,
without dorsal robust setae, with sparse dor-
sal slender setae, distal margins truncated,
without apical penicillate setae, with 1 apical slender seta on each lobe, without apical
robust setae.
Male.—Based on adult male, 3.5 mm.
Gnathopod 1 subchelate, smaller than gnatho-
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 21, NO. 1, 2001
pod 2; anteroventral margin slightly produced, subquadrate; length 2.6 × breadth, (1.2
×) propodus; length 1.4 × breadth, palm transverse, straight. Gnathopod 2 subchelate; carpus subovate, length 2.8 × breadth, shorter
than (0.6 ×) propodus; propodus broad, length
1.4 × breadth, palm slightly acute, slightly
concave, smooth; inner margin with short setae. Pereiopod 5 with long slender setae along
anterior margin; with 3 rows of robust setae
along anterior margin. Pereiopod 6 with slender setae along crenate anterior margin.
Habitat.—Living in association with the
sponges Antho (Isopenectya) chartacea,
Callyspongia sp., Phorbas sp., Mycale
(Carmia) sp., Phoriospongia cf. kirki, and
Holopsamma laminaefavosa.
Remarks.—See comments under S. kuiteri.
Distribution.—Port Jackson and Botany Bay,
New South Wales, Australia.
ACKNOWLEDGEMENTS
At the time of Jerry Barnard’s death (1991), he and I
had illustrated and described Sancho kuiteri. We could
not finish the paper because we had no material of S.
platynotus. After Megan Huggett collected S. platynotus
(1998), I finished the paper more or less as we had
planned. I would like to thank Rudi Kuiter and Megan
Huggett for collecting the animals that form the basis of
this paper. I would like to thank Jerry Barnard, who initially recognised S. kuiteri as a close relative of S.
platynotus and initiated this study. I thank Helen Stoddart, who made the whole animal drawing of S. platynotus and also made many useful comments on the manuscript; Rachael Peart, who illustrated the rest of S.
platynotus; and John Hooper, who identified the sponges.
Finally, this paper is dedicated to Arthur Humes, who
did not study amphipods but for many years provided us
with inspirational copepod taxonomy and with the wonderful Journal of Crustacean Biology.
LITERATURE CITED
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gammaridean Amphipoda.—United States National
Museum Bulletin 271: 1–535.
———, and G. Karaman. 1991. The families and genera
of marine gammaridean Amphipoda (except marine
gammaroids).—Records of the Australian Museum,
Supplement 13(2): 419–866.
Barnard, K. H. 1931. Amphipoda.—Discovery Reports 5:
1–326.
Stebbing, T. R. R. 1888. Report on the Amphipoda collected by H.M.S. Challenger during the years
1873–1876.—Report on the Scientific Results of the
Voyage of H.M.S. Challenger during the years
1873–76, Zoology 29: 1–1737, pls. 1–210.
———. 1897. Amphipoda from the Copenhagen Museum
and other sources.—Transactions of the Linnean Society of London, (Zoology) 7: 25–45.
———. 1906. Amphipoda I. Gammaridea.—Das Tierreich 21: 1–806.
RECEIVED: 10 April 2000.
ACCEPTED: 5 June 2000.