Zoological Journal of the Linnean Society, 2008, 152, 409–458. With 24 figures
Disentangling the subgeneric division of Arenopontia
Kunz, 1937: resurrection of Psammoleptastacus Pennak,
1942, re-examination of Neoleptastacus spinicaudatus
Nicholls, 1945, and proposal of two new genera and a
new generic classification (Copepoda, Harpacticoida,
Arenopontiidae)
SERDAR SAK1, RONY HUYS
FLS2*
and SÜPHAN KARAYTUĞ3
1
Balıkesir Üniversitesi, Fen-Edebiyat Fakültesi, Biyoloji Bölümü, Çağış Kampüsü, 10145, Balıkesir,
Turkey
2
Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK
3
Mersin Üniversitesi, Fen-Edebiyat Fakültesi, Biyoloji Bölümü, Çiftlikköy Kampüsü, 33343, Mersin,
Turkey
Received 10 November 2006; accepted for publication 18 June 2007
A new generic classification is proposed for the 32 valid species of the interstitial marine family Arenopontiidae
(Copepoda, Harpacticoida), primarily based on new observations of type species and reliable descriptions from the
literature. The subgeneric division of Arenopontia Kunz, 1937 is abolished, and both Arenopontia and Neoleptastacus
Nicholls, 1945 are upgraded to full generic rank. Arenopontia is restricted to the subterranea group, comprising
Arenopontia subterranea Kunz, 1937 (type), Arenopontia problematica Masry, 1970, Arenopontia nesaie Cottarelli,
1975, and Arenopontia riedli Lindgren, 1976. The doubtful status of both Arenopontia pontica Apostolov, 1969 and
recent Egyptian records of A. nesaie is discussed, and the alleged cosmopolitanism of A. subterranea is reviewed.
Arenopontia is characterized by the unique morphology of the P1 (prehensile endopod, armature of distal segments
of exopod and endopod). The genus Psammoleptastacus Pennak, 1942 is reinstated to accommodate Psammoleptastacus arenaridus Pennak, 1942 (type), Arenopontia stygia Noodt, 1955 and Psammoleptastacus barani sp. nov. The
latter is described from the Turkish Black Sea coast, and had previously been identified as A. stygia in Bulgarian
waters. The species identified as A. subterranea by Rao & Ganapati in 1969 is considered species inquirenda in
Psammoleptastacus. Neoleptastacus is resurrected to accommodate all arenopontiids that have an inner spinous
process on the P5. The Chilean species Arenopontia clasingi Mielke, 1985, Arenopontia pacifica Mielke, 1985, and
Arenopontia spicata Mielke, 1985 are transferred to Neoleptastacus. The genus Pararenopontia Bodiou & Colomines,
1986 is considered a junior synonym of Neoleptastacus, with its type species Pararenopontia breviarticulata (Mielke,
1975) being relegated to species incertae sedis in this genus. The monotypic genus Mesopontia gen. nov. is
established to accommodate Arenopontia dillonbeachia Lang, 1965, which holds an intermediate position between
Arenopontia and Neoleptastacus. Material from Puget Sound identified as Arenopontia spinicaudata (Nicholls, 1945)
by Chappuis in 1958 is attributed to Mesopontia dillonbeachia comb. nov. Psammoleptastacus orientalis
Krishnaswamy, 1957, Arenopontia intermedia Rouch, 1964, and Arenopontia peteraxi Mielke, 1982 are transferred
to a new genus, Onychopontia gen. nov., together with Onychopontia nichollsi sp. nov. (type), which was
discovered among type material of Neoleptastacus spinicaudatus Nicholls, 1945. Redescriptions are given for
A. nesaie, P. arenaridus, N. spinicaudatus, and M. dillonbeachia. A key to the five arenopontiid genera as well as keys
(or comparative tables) to the species of Arenopontia, Onychopontia, Mesopontia, and the spinicaudatus lineage of
Neoleptastacus are provided. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society,
2008, 152, 409–458.
ADDITIONAL KEYWORDS: distribution – generic diagnoses – revision – sexual dimorphism – taxonomy.
*Corresponding author. E-mail: r.huys@nhm.ac.uk
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
409
410
S. SAK ET AL.
INTRODUCTION
In one of the pioneering papers on the interstitial
fauna of coastal groundwater (‘Küstengrundwasser’)
in north-western Europe, Kunz (1937) proposed the
genus Arenopontia for a new species Arenopontia
subterranea from Schilksee in the Kiel Bay, Germany.
Pennak (1942a) erected the genus Psammoleptastacus for a new species, Psammoleptastacus arenaridus,
from two sandy beaches in the Woods Hole area, USA.
Nicholls (1945) established the genus Neoleptastacus
for its type and only species Neoleptastacus spinicaudatus, from Australia. All three genera were
originally placed in the Canthocamptidae by their
respective authors. Lang (1948) transferred Arenopontia to the subfamily Leptopontiinae in the
Cylindropsyllidae, and Noodt (1955a) suggested
Psammoleptastacus should sink as a synonym of this
genus. Chappuis (1955) believed the separate generic
status of Neoleptastacus was not warranted, and also
relegated the genus to a junior subjective synonym of
Arenopontia. Subsequent authors accepted Chappuis’
course of action, with the exception of Krishnaswamy
(1957) who continued using Neoleptastacus as a valid
genus, and Wells (1967) who preferred a subgeneric
division of Arenopontia into the nominate subgenus
and Neoleptastacus, reflecting the distinct difference
in P5 morphology. This subdivision gained wide
acceptance (e.g. Kunz, 1971; Mielke, 1975; Lindgren,
1976; Itô, 1978; Bodin, 1979, 1988; Bodiou & Colomines, 1986; Wells & Rao, 1987; Cottarelli, Bruno &
Venanzetti, 1994; Karanovic, 2000), but was not universally accepted (Masry, 1970; Cottarelli, 1973, 1975;
Mielke, 1982a, b, 1985, 1987). Bodiou & Colomines
(1986) proposed a new genus Pararenopontia for two
Arenopontia species with reduced leg segmentation:
Arenopontia breviarticulata Mielke, 1975 and Arenopontia trisetosa Mielke (1982a).
Mielke (1982a) questioned the significance attributed to the P5 morphology as a subgeneric discriminant, as some species exhibit a transitionary
condition between the Arenopontia and Neoleptastacus types of P5. Martínez Arbizu & Moura (1994)
argued that the subgenera of Arenopontia (Arenopontia and Neoleptastacus) are not sustainable on
grounds of potential paraphyly and/or polyphyly, and
that Pararenopontia should be synonymized with
Arenopontia. This amalgamation was disputed by
Huys, Bodiou & Bodin (1996a), who resurrected
Pararenopontia, and by Huys et al. (1996b: 35), who
maintained the subgeneric classification (although
they did not explicitly list the subgenus Neoleptastacus). Bodin (1997) offered a compromise by adopting Wells’ (1967) original subdivision and adding
Pararenopontia as a third subgenus (this new rank
was erroneously attributed to Martínez Arbizu &
Moura, 1994). Finally, Wells (2007) abandoned the
subgeneric classification altogether, and maintained
Pararenopontia as a valid genus.
In this paper we have set out to: (1) redefine the
generic boundaries of Arenopontia; (2) provide arguments for the resurrection of Psammoleptastacus as a
valid genus; (3) upgrade Neoleptastacus to its original
generic rank; (4) propose two new genera for species
previously allocated to Arenopontia, and (5) describe
two new species from the Turkish Black Sea coast and
Western Australia, respectively.
MATERIAL AND METHODS
Samples in Turkey were collected using the Karaman–
Chappuis method (Delamare Deboutteville, 1953).
Specimens were cleared in lactic acid and dissected in
lactophenol. Dissected parts were mounted on slides in
lactophenol mounting medium. Broken glass fibres
were added to prevent the animal and appendages
from being compressed by the coverslip, and to facilitate rotation and manipulation, allowing observation
from all angles. Preparations were sealed with
Entellan® (Merck). All drawings have been prepared
using a camera lucida on an Olympus BX-50 or Leica
DMR differential interference contrast microscope.
Measurements were made with an ocular micrometer.
Total body length was measured from the anterior
margin of the rostrum to the posterior margin of the
caudal rami. The scale bars in the illustrations are in
mm. The descriptive terminology is adopted from Huys
et al. (1996b). Abbreviations used in the text are as
follows: ae, aesthetasc; enp, endopod; exp, exopod;
exp-1 (enp-1), exp-2 (enp-2), and exp-3 (enp-3) to
denote the proximal, middle, and distal segment of a
ramus; P1–P6, for swimming legs 1–6. The type material was deposited in the Natural History Museum,
London (NHM) and Balıkesir University Zoology
Museum (BUZM), and was borrowed from the
National Museum of Natural History, Smithsonian
Institution, Washington D.C. (NMNH) and the
Swedish Museum of Natural History, Stockholm
(SMNH).
RESULTS AND DISCUSSION
FAMILY ARENOPONTIIDAE MARTÍNEZ ARBIZU &
MOURA, 1994
Based on the arguments outlined below, a new
generic classification is proposed for the family,
resulting in the upgrade of the subgenera Arenopontia and Neoleptastacus, in the resurrection of Psammoleptastacus, in the rejection of Pararenopontia as a
valid genus, and in the proposal of two new genera,
Mesopontia and Onychopontia. Table 1 summarizes
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
Arenopontia
Mesopontia gen. nov.
Neoleptastacus
Psammoleptastacus
Onychopontia gen. nov.
P1 exp-3 armature
2 pinnate spines + 1
geniculate seta + 1
penicillate seta
2 pinnate spines + 2
geniculate setae
1–2 pinnate spines + 2
geniculate setae
2 pinnate spines + 2
geniculate setae
2 naked setae + 2
geniculate setae
P1 endopod
prehensile
longer than exopod
not prehensile
as long as exopod
not prehensile
as long or longer than
exopod
not prehensile
shorter than exopod
not prehensile
longer than exopod
P1 enp-2 armature
1 spine + 1
geniculate claw
1 spine + 1
geniculate seta
1 spine + 1
geniculate seta*
2 geniculate setae
2 geniculate setae
P2–P3 endopods 씸
two-segmented
two-segmented
one- or two-segmented
two-segmented
two-segmented
P2 enp-2 inner serrate
seta
present
present
usually present†
present
absent
P3 endopod 씹
segmentation
not modified
as in 씸
not modified
as in 씸
not modified
as in 씸
modified
two-segmented
modified
one-segmented
P4 exp-3 inner serrate
seta
absent
present
usually present‡
absent
absent
P5 with outer basal
seta +
3 or 4 discrete
elements
4 discrete elements
(innermost one
spiniform)
1–3 discrete elements +
inner spinous
process
4 discrete elements
innermost smaller in
씹
3 or 4 discrete
elements in 씸
3 discrete elements in
씹 (with innermost
smaller or fused)
*Except speluncae lineage (N. speluncae and N. phreaticus), which has two geniculate setae.
†Except N. ornamentus and N. reductaspina.
‡Except N. australis and N. pacifica; unknown in N. accraensis.
CLASSIFICATION OF ARENOPONTIIDAE
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
Table 1. Salient features differentiating arenopontiid genera. Apomorphic character states are set in boldface
411
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S. SAK ET AL.
Table 2. New classification reflecting restricted taxonomic concept of Arenopontia and reallocation of remaining species
to Neoleptastacus, Psammoleptastacus, and two new genera
ARENOPONTIA KUNZ, 1937
A. subterranea Kunz, 1937*
A. problematica Masry, 1970
A. nesaie Cottarelli, 1975
A. riedli Lindgren, 1976
PSAMMOLEPTASTACUS PENNAK, 1942
P. arenaridus Pennak, 1942*
P. stygius (Noodt, 1955) comb. nov.
P. barani sp. nov.
ONYCHOPONTIA GEN. NOV.
O. orientalis (Krishnaswamy, 1957) comb. nov.
O. intermedia (Rouch, 1962) comb. nov.
O. peteraxi (Mielke, 1982) comb. nov.
O. nichollsi sp. nov.*
MESOPONTIA GEN. NOV.
M. dillonbeachia (Lang, 1965) comb. nov.*
NEOLEPTASTACUS NICHOLLS, 1945
N. spinicaudatus Nicholls, 1945*
N. australis (Chappuis, 1952) comb. nov.
N. acanthus (Chappuis, 1954) comb. nov.
N. longiremis (Chappuis, 1955) comb. nov.
N. secundus Krishnaswamy, 1957
N. africanus (Chappuis & Rouch, 1961) comb. nov.
N. accraensis (Lang, 1965) comb. nov.
N. indicus (Rao, 1967) comb. nov.
N. ishikarianus (Itô, 1968) comb. nov.
N. angolensis (Kunz, 1971) comb. nov.
N. gussoae (Cottarelli, 1973) comb. nov.
N. trisetosus (Mielke, 1982) comb. nov.
N. clasingi (Mielke, 1985) comb. nov.
N. pacificus (Mielke, 1985) comb. nov.
N. spicatus (Mielke, 1985) comb. nov.
N. chaufriassei (Bodiou & Colomines, 1986) comb. nov.
N. ornamentus (Mielke, 1987) comb. nov.
N. reductaspina (Mielke, 1987) comb. nov.
N. phreaticus (Cottarelli et al., 1994) comb. nov.
N. speluncae (Cottarelli et al., 1994) comb. nov.
N. huysi (Karanovic, 2000) comb. nov.
*Type species of respective genera.
Species inquirendae and species incertae sedis is not listed.
the main diagnostic characters for each genus. The
updated generic assignment of the 32 valid species in
the family is shown in Table 2.
GENUS ARENOPONTIA KUNZ, 1937
The genus Arenopontia currently contains 32 species
allocated to three subgenera (Bodin, 1997; Karanovic,
2000) (or 30 species when the subgenus Pararenopontia is attributed full generic rank; see Wells,
2007). The subgenus Arenopontia encompasses 13
species and possibly one subspecies [Apostolov (1973)
claimed that Arenopontia pontica Apostolov, 1969 is
a subspecies of A. subterranea], whereas Karanovic
(2000) listed 17 valid species in the subgenus Neoleptastacus [note that Arenopontia sakagamii Itô,
1978 was also listed by this author, but according to
Wells & Rao (1987) this species is synonymous with
Arenopontia indica Rao, 1967]. The subgeneric division first proposed by Wells (1967) fell into disuse
in the 1980s, when Mielke (1982a, b, 1985, 1987)
described several new species from Central and South
America without attributing them to either subgenus.
Unfortunately, Bodin (1988, 1997) erroneously listed
three of those species, Arenopontia clasingi, Arenopontia pacifica, and Arenopontia spicata (all described
by Mielke, 1985), under the nominate subgenus
Arenopontia (Arenopontia), as if Mielke (1985) had
originally intended such a subgeneric assignment. It
is obvious from Mielke’s (1985, 1987) descriptions,
however, that these species share the Neoleptastacus
type of P5, and should be assigned to this subgenus if
Wells’ (1967) subdivision bears any phylogenetic significance. Bodin’s (1988, 1997) error unfortunately
perpetuated in the literature, as exemplified by Karanovic’s (2000) recent key to the subgenus Neoleptastacus, which makes no reference to Mielke’s (1985)
species. Arenopontia clasingi, A. pacifica, and A. spicata are here formally transferred to Neoleptastacus,
which will be attributed full generic rank (see below).
The subgenus Arenopontia currently encompasses
the following species: A. subterranea; Arenopontia
arenarida (Pennak, 1942a); Arenopontia stygia Noodt,
1955b; Arenopontia orientalis (Krishnaswamy, 1957);
Arenopontia intermedia Rouch, 1962; Arenopontia
dillonbeachia Lang, 1965; Arenopontia problematica
Masry, 1970; Arenopontia nesaie Cottarelli, 1975;
Arenopontia riedli Lindgren, 1976; and Arenopontia
peteraxi Mielke, 1982a. Bodin (1979, 1988, 1997)
added A. subterranea Kunz? sensu Şerban & EitelLang (1957) as a species incertae sedis, but the latter
should be regarded as a nomen nudum. Various
Eastern European authors (e.g. Georgescu, Marcus &
Şerban, 1962) have repeatedly referred to Şerban &
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
CLASSIFICATION OF ARENOPONTIIDAE
Eitel-Lang’s (1957) paper as ‘Notes sur les Copépodes
de la Mer Noire. Izdanija, Skopje’; however consistently, no proper citation of volume number or pagination has been given. According to Şerban (1959), the
authors were at that time still in the process of
submitting the paper (‘Une description detaillée en
sera publiée par Şerban & Eitel-Lang’), and it has now
been confirmed (C. Pleşa, pers. comm. to RH, 1 August
1996) that the manuscript was never published. The
reference nevertheless mistakenly persisted in modern
literature (e.g. Apostolov & Marinov, 1988).
Morphological comparison revealed a core group of
closely related species within the (sub)genus Arenopontia, encompassing the type species A. subterranea,
A. pontica, A. problematica, A. nesaie, and A. riedli.
These five species differ from other members of the
family in their unique P1 morphology, including: (1)
the prehensile endopod with enp-1 being distinctly
elongate, and with enp-2 bearing an outer spine and
an inner geniculate claw, and (2) the modification of
the inner distal element of exp-3 into a penicillate
seta. Based on these autapomorphies, we here restrict
the generic concept of Arenopontia to this subterranea
group. Our unpublished studies based on sandy beach
samples from all over Europe revealed that many new
species await description (e.g. Sak, Karaytuǧ & Huys,
in press a), and that the five currently known species
only represent the tip of the iceberg. The genus is
primarily restricted to the Northern Hemisphere, the
only exception being Wells’ (1967) doubtful outlier of
A. subterranea in Mozambique.
Diagnosis: Arenopontiidae. Urosomites without conspicuous surface ornamentation. Anal somite without
paired dorsolateral spinous processes. Anal operculum not modified. Hyaline frills of abdominal somites
with rectangular digitate lappets. Caudal ramus with
dorsolateral spur or raised spinular row near medial
margin. P1 exopod: three-segmented; exp-1 with outer
spine; exp-3 with two spines, one outer distal geniculate seta, and one inner distal penicillate seta. P1
endopod: prehensile, longer than exopod; enp-2 with
one outer distal spine and one inner distal geniculate
claw. P2–P3 endopods: two-segmented. P3 enp-2 with
outer distal element defined at base or absent. P4
enp-2 with well developed outer distal element. Armature formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.021
Endopod
0.110 or 0.120
0.010 or 0.020
0.020
P3 endopod male: not sexually dimorphic, twosegmented. P5 with outer basal seta and three or four
discrete elements: innermost one distinctly smaller in
males. P6 male with one or two seta(e).
413
Type species: Arenopontia subterranea Kunz, 1937
(by monotypy).
Other species: Arenopontia problematica Masry, 1970;
Arenopontia nesaie Cottarelli, 1973; Arenopontia
riedli Lindgren, 1976.
Species inquirendae: Arenopontia pontica Apostolov,
1969; Arenopontia nesaie Cottarelli, 1975 sensu Mitwally & Montagna (2001).
Nomen nudum: Arenopontia subterranea Kunz, 1937?
sensu Şerban & Eitel-Lang (1957).
ARENOPONTIA
SUBTERRANEA
KUNZ, 1937
Arenopontia (Arenopontia) subterranea Kunz, 1937:
Wells (1967)
Original description: Kunz (1937): pp. 107–110; Abb.
8 (figs 38–42), 9 (figs 43–47), 10 (figs 48–51).
Type locality: Germany, Kieler Förde, Schilksee; ‘Küstengrundwasser’ (intertidal coastal groundwater).
Arenopontia subterranea has been reported from a
wide range of localities throughout Europe, from the
Baltic to the Black Sea basin. With additional records
from Madeira (Delamare Deboutteville, 1960b), India
(Rao, 1967, 1968, 1970, 1980, 1991; Rao & Ganapati,
1968, 1969; Rao & Misra, 1983), Mozambique
(Wells, 1967), and North Carolina (Lindgren, 1976) it
is not surprising that this species has been regarded as
potentially cosmopolitan (Wells, 1967, 1986; Lindgren,
1976). Lindgren (1976) claimed its range might be
extended with more investigation of sandy beaches in
the Pacific. Unfortunately, the great majority of these
records are not accompanied by illustrations, and
consequently their authenticity cannot be verified. The
discovery of a closely related species from the Isle of
Sylt (Sak, 2004) casts further doubt on the validity of
most north-western European, and even some
German, records. Arlt’s (1983) illustrations show that
his Baltic specimen does not belong to A. subterranea
either, raising the suspicion that not all records from
east of the Skagerrak necessarily pertain to the type
species. There is no doubt that many authors have
attributed their material to A. subterranea on the sole
basis that this species shows extensive intraspecific
variability. The true range of the species is as yet
unknown, and the only reliable records appear to be
restricted to German waters: (1) North Sea coast – Isle
of Sylt (Noodt, 1952, 1956, 1957; Mielke, 1975, 1976),
Amrum (Noodt, 1956, 1957), Sankt Peter-Ording
(Noodt, 1956), and Helgoland (Martínez Arbizu &
Moura, 1994); (2) Kieler Bucht – Schilksee (Kunz,
1937; Noodt, 1956), Bottsand, Gelting Birk, Weißenhaus, and Heiligenhafen (Noodt, 1956, 1957).
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
414
S. SAK ET AL.
The type material of A. subterranea, as well as the
remainder of Kunz’ earlier collections, were destroyed
during World War II when the Institut für
Meereskunde was heavily bombed in 1944 (Schriever,
1984). We have been unable to obtain topotype or
other material that could be attributed with confidence to A. subterranea, and instead we have selected
A. nesaie for the model description. Illustrations and
text are based on material collected from the Turkish
west coast (Marmara Sea), which represents a considerable extension of the range for the species.
ARENOPONTIA
NESAIE
COTTARELLI, 1975
Arenopontia (Arenopontia) nesaie Cottarelli, 1975
Arenopontia nesiae Cottarelli, 1975:
Martínez Arbizu
calami)
&
Moura
(1994:
57)
(lapsus
63)
(lapsus
Arenopontia nessiae Cottarelli, 1975:
Martínez Arbizu
calami)
&
Moura
(1994:
Arenopontia ciplaki Sak, 2004 (nomen nudum)
Original description: Cottarelli (1975): pp. 65–70;
figures 1–11, 13–16, 18–19, 21–23.
Type locality: Italy, Sardinia, near Cagliari, Bay of
Quartu S. Elena, Poetto beach.
Material examined: (1) one 씸 dissected on eight slides
(NHM reg. no. 2006. 1953), one 씹 mounted in toto on
slide (NHM reg. no. 2006. 1954), one 씹 dissected on
eight slides (NHM reg. no. 2006. 1955), 22 씸씸 and 22
씹씹 preserved in alcohol (NHM reg. no. 2006. 1956–
1965); (2) > 50 씸씸 and > 50 씹씹 preserved in alcohol
(deposited in BUZM). All material was collected
from Dutlimanı Beach (Marmara Sea), 40°22.479′N,
28°03.080′E, Balıkesir Province, Turkey; leg. S.
Karaytuğ and S. Sak, 18 September 2001.
Redescription
Female: Total body length from tip of rostrum to
posterior margin of caudal rami: 341–396 mm
(mean = 366 mm, n = 25). Maximum width: 38 mm
(mean of 20 individuals = 41 mm), measured at posterior margin of cephalothorax. Body: slender and cylindrical, without clear distinction between prosome and
urosome (Fig. 1A, B). Hyaline frills of thoracic somites
weakly developed and crenulated; those of genital
double-somite and free abdominal somites strongly
developed, and consisting of rectangular digitate
lappets (Figs 1A, B, 2A, B). Genital double-somite
(Figs 1A, B, 2A): slightly longer than wide; without
chitinous ribs marking original segmentation; with
two mid-dorsal, two lateral, and two ventral pores.
Anal somite (Fig. 3A, B): with two dorsal and two
lateral pores. Anal operculum: with minute pinnules
along free distal margin (Fig. 3A). Anus: positioned
subterminally between caudal rami. Rostrum
(Fig. 1C): small, broadly subtriangular, tapering distally, with two delicate sensillae.
Caudal rami: approximately twice longer than wide
(measured in dorsal view), tapering posteriorly; with
a proximal pore dorsally (Fig. 3A), one pore near the
ventral proximal margin (Fig. 2A), and one pore laterally near the insertion site of seta III (Fig. 3B);
outer distal corner produced into posteriorly directed
recurved spinous process, accompanied by outer
spinular row at base (Fig. 3A, B); dorsal surface with
flagellate spur-like process near inner margin, accompanied by a few tiny spinules near base (Figs 1D, 3B).
Armature consisting of seven setae: seta I, small;
setae II and III, long and naked; seta IV, short,
sparsely pinnate, located between seta V and spinous
process; seta V, long and with fracture plane; seta VI,
small, naked, and located at inner distal corner;
seta VII, foliaceous and triarticulate at base.
Antennule (Fig. 3C): long, six-segmented. Segment 1 with a tiny seta near the anterodistal margin.
Segment 2 longest, about 3.5 times longer than wide.
Segment 4 with long aesthetasc (32-mm long) fused at
base with seta. Distal segment: with seven naked setae
(two of which are spatulate) and apical acrothek,
consisting of short aesthetasc (20-mm long) and two
slender setae. Armature formula: 1-[1], 2-[7 + 1
plumose], 3-[4], 4-[(1 + ae)], 5-[1], 6-[7 + acrothek].
Antenna (Fig. 3D, E): coxa small, without ornamentation. Allobasis: about 2.7 times as long as maximum
width; original segmentation marked by partial transverse surface suture; with two spinular rows, as illustrated. Exopod one-segmented, elongate, with a naked
apical seta (about 3.3 times longer than exopod). Free
endopod with two spinular rows on anterior surface,
and with finer spinules at outer distal corner; lateral
armature consisting of two short spines; apical armature consisting of two spines and three geniculate
setae, the longest of which with spinules around
geniculation, and fused basally to tiny accessory
seta.
Mandible: with two-segmented palp (Fig. 2D); basis
elongate with one lateral seta; endopod with one
inner, one outer, and three apical setae; all armature
elements naked. Gnathobase: with coarse teeth distally, and with one naked seta at dorsal corner.
Maxillule (Fig. 1E): with praecoxal arthrite bearing
two setae and five spines around distal margin. Coxal
endite: with two long naked setae. Basis with rami
entirely incorporated; palp represented by nine naked
setae.
Maxilla (Fig. 2E): syncoxa with two cylindrical
endites; proximal endite with three setae; distal
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CLASSIFICATION OF ARENOPONTIIDAE
415
Figure 1. Arenopontia nesaie Cottarelli, 1975 (씸). A, habitus, dorsal view. B, habitus, lateral view. C, rostrum, dorsal
view. D, left caudal ramus, inner lateral view. E, maxillule.
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416
S. SAK ET AL.
Figure 2. Arenopontia nesaie Cottarelli, 1975. A, urosome 씸, ventral view. B, urosome 씹, ventral view. C, genital field
씸. D, mandible. E, maxilla. F, maxilliped.
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CLASSIFICATION OF ARENOPONTIIDAE
417
Figure 3. Arenopontia nesaie Cottarelli, 1975 (씸). A, caudal rami, anal somite, and posterior margin of penultimate somite,
dorsal view. B, left caudal ramus, outer lateral view. C, antennule. D, antenna, outer lateral view. E, antenna, inner lateral
view.
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418
S. SAK ET AL.
endite with two setae. Allobasis: drawn out into long
claw; with one accessory setae. Endopod onesegmented , and with three setae. All elements naked.
Maxilliped (Fig. 2F): syncoxa small and unarmed.
Basis: elongate and unarmed. Endopod with small
accessory seta, and with slightly curved claw bearing
subterminal spinule.
P1 (Fig. 4A): intercoxal sclerite long and rectangular. Praecoxa: triangular and naked. Coxa:
without ornamentation. Basis: with spinular row
near bases of endopod and exopod; anterior surface
with a proximal pore and a small inner seta.
Exopod: three-segmented; exp-1 and exp-2 with
spinules around outer margin; exp-1 longest, with
long unipinnate outer spine; exp-2 without outer
element; exp-3 with short unipinnate outer spine, a
long curved unipinnate spine, and one geniculate
seta distally, and one inner, apically penicillate seta
subdistally. Endopod: two-segmented, prehensile;
enp-1 9.3 times longer than wide, and about twice
longer than exopod; with a serrate inner seta in
proximal third, and a subdistal spinule along outer
margin; enp-2 slightly longer than wide, with a
short unipinnate spine, a geniculate claw, and a
small inner spinule.
P2–P4 (Fig. 4B–D): intercoxal sclerites naked,
wider in P2, but more deeply concave in P3–P4.
Praecoxae: small and naked. Coxae: squarish and
without ornamentation. Bases: smaller than coxae,
with a spinular row near base of endopod (P3–P4);
anterior surface with a pore; outer basal seta absent
(P2), plumose (P3), or naked (P4). Exopods: threesegmented; segments with spinular ornamentation,
as illustrated; inner distal seta of exp-3 sparsely
bipinnate, all other elements unipinnate; P3–P4 exp-3
with anterior pore. Endopods: two-segmented; P2–P4
enp-1 about 1.5, 2.2, and 3.0 times longer than their
respective distal segments, with few spinules, as illustrated. P2: enp-1 with a long, apically serrate, backwardly directed seta near proximal inner corner.
P2–P3: enp-2 with a long, bipinnate, apical seta. P4:
enp-2 with apically serrate seta, fused at base, and
long unipinnate seta at outer distal corner. Armature
formula as follows: P2, exopod, 0.0.021, endopod,
0.110; P3, exopod, 0.0.021, endopod, 0.010; P4,
exopod, 0.0.021, endopod, 0.020.
Fifth legs (Fig. 2A) closely set together, but not
touching in ventral midline. Baseoendopod and
exopod: fused, forming a rectangular plate; distal
margin with three pinnate setae, middle one markedly shorter than the others, but not vestigial; outer
basal seta, long and plumose.
Genital field: positioned near anterior margin of
genital double-somite (Fig. 2A). Genital apertures
(Fig. 2C): fused forming median common slit; closed
off by fused P6 forming operculum with three minute
spinous processes on either side; copulatory pore
located midventrally, close to genital slit; seminal
receptacles difficult to discern.
Male: Total body length from tip of rostrum to
posterior margin of caudal rami: 320–374 mm
(mean = 346 mm; N = 25). Maximum width: 40 mm
(mean = 38, N = 20), measured at cephalothorax.
Body ornamentation (Fig. 5A): essentially as in
female. Sexual dimorphism: in antennule, genital segmentation, and P5 and P6. Spermatophore length:
approximately 35 mm.
Antennule (Fig. 5B, C): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 2
longest, and about 2.7 times longer than wide;
segment 4 an incomplete sclerite with one modified
(fused at base) and one tiny element; segment 5 with
three setae plus long aesthetasc (42-mm long) fused
basally to a small slender seta; segment 6 with a
spinulose spine and long distal seta; segment 7 with
three modified spines and a seta; segment 8 with a
modified spine; distal segment with seven naked
setae (two of which spatulate) and apical acrothek.
Setal formula: 1-[1], 2-[7 + 1 plumose], 3-[4 + 2
spines], 4-[1 + 1 modified], 5-[3 + (1 + ae)], 6-[1 + 1
modified],
7-[1 + 3
modified],
8-[1
modified],
9-[7 + acrothek]. Acrothek consisting of short aesthetasc (16-mm long) fused basally to two slender
setae.
P5 (Fig. 2B): with armature as in female, but with
middle and inner elements comparatively shorter.
Sixth legs (Fig. 2B): asymmetrical, with smallest P6
closing off functional gonopore; each with a long
plumose seta.
Remarks: Arenopontia nesaie was originally described from Poetto Beach in the Bay of Quartu S.
Elena near Cagliari, Sardinia (Italy). Our material
differs from Cottarelli’s (1975) description in some
aspects, but these are most likely attributable to
deficiencies in the original figures. In the type material the marginal spines on the P5 of both sexes
appear shorter; however, the flagellate distal parts of
these elements are usually difficult to discern, and it
seems conceivable that they were not illustrated correctly in the original description. Similarly, Cottarelli
(1975) did not illustrate the spinules at the base of
the spur and around the terminal process of the
caudal ramus, but such morphological minutiae were
generally overlooked prior to the advent of differential
interference contrast microscopy. The female antennule has fewer setae on the proximal segments than
in the Turkish material, but this can be attributed to
the fact that Cottarelli viewed the appendage in
dorsal aspect and hence overlooked various setae
arising from the ventral surface. The three-
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CLASSIFICATION OF ARENOPONTIIDAE
419
Figure 4. Arenopontia nesaie Cottarelli, 1975 (씸). A, P1, anterior view. B, P2, anterior view. C, P3, anterior view. D, P4,
anterior view.
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420
S. SAK ET AL.
Figure 5. Arenopontia nesaie Cottarelli, 1975 (씹). A, habitus, dorsal view. B, antennule, ventral view. C, antennule,
anterior view.
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CLASSIFICATION OF ARENOPONTIIDAE
segmented mandibular palp, which was considered
diagnostic for A. nesaie, was not observed in our
material, and requires confirmation. The extra
segment boundary indicates a two-segmented
endopod, which has thus far not been reported for any
other oligoarthran harpacticoid (but see Mitwally &
Montagna, 2001; cf. below). It is also noteworthy that
Cottarelli (1975) had accidentally rotated exp-3 in his
drawing of the P2. His drawing of the female genital
field superimposes external and internal structures;
for a more accurate interpretation see Fig. 3C and
Martínez Arbizu & Moura (1994: fig. 2c, as A. nessiae). The Sardinian specimens are somewhat smaller
[334 mm (씸), 285–300 mm (씹)] than the Marmara
population [341–396 mm (씸), 320–374 mm (씹)], but it
is questionable whether this size discrepancy has
any significance beyond the range of intraspecific
variability.
Arenopontia nesaie appears to be widely distributed
in the Mediterranean, with confirmed intertidal
records from El Saler (Valencia, Spain) by Martínez
Arbizu & Moura (1994, as A. nesiae), Sardinia
(Cottarelli, 1975), the mouth of the Trigno River on the
Adriatic coast (Molise, Italy) by Bruno, Cottarelli &
Berrera (1998), and Dutlimanı beach (Sea of
Marmara, Turkey) by Sak (2004, as A. ciplaki; present
account). Mitwally & Montagna (2001) provided a
redescription of A. nesaie based on specimens collected
from three beaches (Bir Masoud, El Mamoura, and El
Shatby) near Alexandria, Egypt, but the many deficiencies in their illustrations make it difficult to validate their identification. According to Wells (2007),
Mitwally & Montagna (2001) make statements about
the setation of P1–P4 that, if true, mean that their
material cannot belong to Arenopontia. It is obvious
that their atypical setal formula results from a failure
to distinguish between ornamentation elements (such
as long spinules) and genuine setae/spines. Their
reports of an outer seta on P1 enp-1 and P3–P4 enp-1,
as well as their claim of four elements on P2 exp-3, are
false and do not reflect deficiencies in Cottarelli’s
(1975) original description, as claimed by the authors.
The elements on the female P5 are distinctly longer
than in A. nesaie (but are similar to our specimens),
and the caudal ramus appears shorter. The variability
illustrated for the male P5 suggests that Mitwally &
Montagna (2001) had an amalgam of Arenopontia
species in their samples. No information was given on
the number of setae on the male P6. The distal
segment of the P4 exopod appears rotated in their
Fig. 11G. Finally, the mandibular palp is erroneously
illustrated as three-segmented (see above). Pending
re-examination of more material of the Egyptian
populations, A. nesaie Cottarelli (1975) sensu Mitwally & Montagna (2001) is considered species
inquirenda in Arenopontia.
ARENOPONTIA
PONTICA
421
APOSTOLOV, 1969
Original description: Apostolov (1969): pp. 125–127;
Abb. 36–45.
Type locality: Bulgaria, south of Lozenetz, Düni
Beach; 5 m from low-tide mark.
Remarks: Apostolov’s (1969) description of A. pontica,
from the Bulgarian Black Sea coast, is a taxonomic
nightmare because of several internal inconsistencies
between the text and illustrations. Apostolov (1969:
111) claimed to have found two females (although on
p. 125 he stated that three females were recorded),
but for some inexplicable reason provided a brief
diagnosis of the male. He referred to Figure 46 in his
description of the male P5, but this figure is not
printed. His illustrations of the female show several
extraordinary features not found in any other
member of the Arenopontiidae: (1) the antennary
exopod is bisetose – in all species of Arenopontia this
ramus displays only one apical seta; (2) P1 exp-2
bears an outer spine – the absence of this spine is a
high-level diagnostic, being a synapomorphy linking
the Parastenocarididae, Leptopontiidae, and Arenopontiidae (Martínez Arbizu & Moura, 1994); (3) P1
enp-1 lacks an inner seta – this seta is present in all
species, except for the inadequately described A. problematica and Arenopontia accraensis Lang, 1965 – we
have been able to confirm its presence in the types of
A. problematica; (4) P2–P3 exp-3 with four elements,
i.e. with two outer spines and two terminal setae – all
Arenopontiidae have only one outer spine and share a
[021] setal formula on the distal exopod segment –
note that Apostolov (1969) contradicts himself in the
setal formula table on p. 125 (three elements), his
Figure 42 (four elements), and the comparative table
on p. 127 (four elements); (5) P2–P3 exp-2 bears a
long inner seta – the latter seta is absent in all other
arenopontiids, except for Arenopontia angolensis
Kunz, 1971, which according to Kunz’ (1971) setal
formula possesses a seta on P2 exp-2. However, as
Kunz neither illustrated the P2 nor mentioned this
character in the text or the table comparing Arenopontia africana f. africana and A. africana f. angolensis
(he does state that the P2 is as in the nominate
subspecies, apart from the ornamentation of the inner
seta on enp-2), we strongly suspect that his report is
based on a slip of the pen in his table, rather than on
an observational error.
Apostolov (1969) recognized a close relationship
with A. subterranea, A. indica and A. sp. sensu Griga
(1964) [the latter was later identified as conspecific
with Stenocaropsis valkanovi (Marinov, 1974), family
Cylindropsyllidae]. In our opinion it is impossible to
make any positive statement on the identity and
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S. SAK ET AL.
Table 3. Diagnostic characters of Arenopontia species
A. riedli
A. nesaie
A. subterranea
A. problematica
P1 enp-1:exp
P2 enp
P3 enp
P5씸씹
P6씹
CR
An Op
1.4
2.0
1.5
1.5†
0.120
0.110
0.110
0.110‡
0.020
0.010
0.010
0.010‡
5
4
4
4
2
1
2?*
2?*
Spur
Spur
Spinules
Spinules
Smooth
Pinnate
Smooth
Smooth
*According to Kunz (1937) and Masry (1970) the P6 is a minute plate bearing three elements, but it is likely that these
claims are based on observational errors.
†Based on Sak’s (2004) redescription.
‡Masry (1970) claimed there are two distal elements on P2–P3 enp-2 but this has been corrected by Sak (2004).
CR = caudal ramus; An Op = anal operculum.
possible relationships of A. pontica other than that
this species can be assigned to the genus Arenopontia
as diagnosed herein. Pending redescription, A. pontica is considered here as species inquirenda. This
course of action is in contrast to Marinov’s (1971)
suggestion to relegate A. pontica to a junior subjective
synonym of A. subterranea. Marinov rightly pointed
out some of the weaknesses in Apostolov’s (1969)
description, but it remains a mystery how he reconciled the many differences between the latter and his
own illustrations of A. subterranea from the Bulgarian coast.
Inspired by the variability reported for French
mediterranean (Chappuis, 1954a) and Romanian
populations (Şerban, 1959) of A. subterranea, but
apparently unaware of Marinov’s (1971) paper,
Apostolov (1973) claimed that A. pontica may well
be a synonym of the latter. He further proposed that
the Black Sea specimens represent a new subspecies
of A. subterranea, but refrained from formally
naming it. Apostolov stated that considerable variability was found in the caudal rami, the P1 exopod,
and the P5, but it is conceivable that this is at least
partly attributable to his failure to discriminate
between two or more coexisting species. His drawings of the female P5 clearly refer to two different
species: his Figure 5 shows a fifth leg of the subterranea type, whereas Figure 6 was almost certainly
based on the species previously identified by
Marinov (1971) as A. stygia (and described below as
Psammoleptastacus barani sp. nov.). In accordance
with Şerban’s (1959) observations, Apostolov (1973)
maintained that his material did not display the
foliaceous seta VII, or the penicillate seta on P1
exp-3.
Key to species: A simple dichotomous identification
key is difficult to construct; however, species can be
reliably identified by considering the salient diagnostic characters summarized in Table 3.
GENUS PSAMMOLEPTASTACUS PENNAK, 1942A
Pennak (1942a) proposed this genus for a new species,
P. arenaridus, collected from two sandy beaches near
Woods Hole, and placed it without any further
comment in the Canthocamptidae. He remarked on the
superficial resemblance with other interstitial genera
(Leptastacus, Paraleptastacus, and Arenopontia), but
considered the differences in the antenna, maxillipeds,
P5, and caudal rami sufficient for generic distinction.
Pennak’s (1942a) paper remained largely unnoticed
until Noodt (1955b) synonymized Psammoleptastacus
with Arenopontia, a course of action that was endorsed
by Lang (1965) but was overlooked by Krishnaswamy
(1957), who added a second species, Psammoleptastacus orientalis Krishnaswamy, 1957 from the Madras
coast. As noted by Wells (1967), Lang’s (1965) statement that P. orientalis belongs to Arenocaris (Leptastacidae) is obviously a slip of the pen.
Noodt (1955b) considered A. stygia to be most closely
related to A. arenarida, recognizing some subtle differences in the caudal rami and P2–P4, whereas Lindgren
(1976) suggested A. stygia is potentially ‘. . . an
intraspecific variation of A. arenarida’. The genus
Psammoleptastacus is reinstated herein for the latter
two species and a new species, P. barani sp. nov., from
the Turkish Black Sea coast, which had previously
been misidentified as A. stygia by Marinov (1971).
Arenopontia subterranea Kunz, 1937 sensu Rao &
Ganapati (1969) is regarded as a species inquirenda in
Psammoleptastacus, and P. orientalis is transferred to
Onychopontia gen. nov. Psammoleptastacus differs
from Arenopontia and other arenopontiid genera in the
small size of the P1 endopod, which is shorter than the
exopod. It is most closely related to Onychopontia, with
which it shares the sexual dimorphism on the P3
endopod (apomorphic) and the presence of two geniculate setae on P1 enp-2.
Diagnosis: Arenopontiidae. Urosomites: without conspicuous surface ornamentation. Anal somite: without
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CLASSIFICATION OF ARENOPONTIIDAE
paired dorsolateral spinous processes. Anal operculum: not modified. Hyaline frills of abdominal somites
with rectangular digitate lappets. Caudal ramus:
with dorsolateral spur near medial margin. P1
exopod: three-segmented; exp-1 with outer spine;
exp-3 with two spines and two geniculate setae. P1
endopod: not prehensile, shorter than exopod; enp-2
with two geniculate setae. P2–P3 endopods: twosegmented. P3 endopod: with outer distal element
defined at base. P4 endopod: with well-developed
outer distal element. Armature formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.021
Endopod
0.120
0.020
0.020
P3 endopod male: sexually dimorphic, twosegmented; enp-1 unarmed; enp-2 minute, with
strong spinule on outer margin, curved spine distally
(sometimes fused at base), and fine seta on inner
margin. P5: with outer basal seta and four discrete
elements; innermost one distinctly smaller in male.
P6 male: with two setae.
Type species: Psammoleptastacus arenaridus Pennak,
1942a (by monotypy).
Other species: Arenopontia stygia Noodt, 1955b = P.
stygius (Noodt, 1955b) comb. nov.; P. barani sp. nov.
Species inquirenda: Arenopontia subterranea Kunz,
1937 sensu Rao & Ganapati (1969)
PSAMMOLEPTASTACUS
ARENARIDUS
PENNAK, 1942A
Psammoleptastacus arenardius Pennak, 1942a: Coull
(1977) (lapsus calami)
Arenopontia
(1955a)
arenarida
(Pennak,
Arenopontia (Arenopontia)
1942a): Wells (1967)
1942a)
arenarida
Noodt
(Pennak,
Arenopontia arenardia (Pennak, 1942a): Coull (1971,
1977) (lapsus calami)
Arenopontia stygia Noodt (1955b) sensu Coull (1971)
and Lindgren (1976)
Original description: Pennak (1942a): pp. 275–278;
plate I, figures 1–11.
Type locality: USA, Massachusetts, Woods Hole.
Pennak (1942a) collected material from both Nobska
and north Cape Cod beaches, but did not specify the
type locality; sand washings in vicinity of high tide
mark.
423
Material examined: NMNH: one 씹 syntype mounted
in toto on slide, and partly remounted by one of us
(RH); erroneously labelled ‘Paraleptastacus arenaridus n.g. n. sp.’; Cat. no. 81982; leg. R.W. Pennak, September 1939.
Partial redescription
Male: Total body length from tip of rostrum to posterior margin of caudal rami: 325 mm. Body: slender
and cylindrical, without clear distinction between
prosome and urosome. Hyaline frills of thoracic
somites weakly developed and crenulated (Fig. 6A,
B); those of abdominal somites strongly developed
and consisting of rectangular digitate lappets
(Fig. 6A).
Caudal rami (Fig. 6A, C): approximately 2.8 times
longer than basal width, tapering posteriorly; with
one pore dorsally, one pore near ventral proximal
margin, and two pores laterally near outer spinules;
outer distal corner produced into posteriorly directed
recurved spinous process, accompanied by ventral
spinular row at base; dorsomedial surface with posteriorly directed spinous process. Armature consisting
of seven setae: seta I, small; setae II and III, long and
naked; seta IV, short, sparsely pinnate, located
between seta V and distal spinous process; seta V,
long and with fracture plane; seta VI, small, naked,
and fused at base to seta V; seta VII, weakly foliaceous and triarticulate at base.
Rostrum (Fig. 6D): small, broadly subtriangular,
tapering distally, with two delicate sensillae and subapical pore.
Antennule (Fig. 7A): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 2
longest; segment 4 an incomplete sclerite with one
modified (fused at base) and one tiny element; segment 5 with long aesthetasc fused basally to seta;
segments 6–8 with one seta and one basally fused
spiniform element. Setal formula: 1-[1], 2-[7 + 1
plumose], 3-[4 + 1 pinnate spine], 4-[1 + 1 modified],
5-[2 + (1 + ae)], 6-[1 + 1 modified], 7-[1 + 1 modified],
8-[1 + 1 modified], 9-[7 + acrothek]. Acrothek consisting of short aesthetasc fused basally to two slender
setae.
P1 (Fig. 7B): coxa without ornamentation. Basis:
with spinular row near bases of endopod and exopod;
anterior surface with inner naked seta. Exopod:
three-segmented; about 1.3 times the length of
endopod; all segments with spinules along outer
margin; exp-1 longest, with long unipinnate outer
spine; exp-2 without outer element; exp-3 with two
unipinnate spines and two geniculate setae of different lengths. Endopod: two-segmented, not prehensile;
enp-1 slightly longer than exp-1, with a serrate seta
at about halfway along the length of the inner
margin, and with two subdistal spinules along outer
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424
S. SAK ET AL.
Figure 6. Psammoleptastacus arenaridus Pennak, 1942a (씹). A, urosome, ventral view. B, P5 and P6, ventral view.
C, posterior portion of penultimate somite, anal somite, and caudal rami, dorsolateral view. D, rostrum.
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CLASSIFICATION OF ARENOPONTIIDAE
425
Figure 7. Psammoleptastacus arenaridus Pennak, 1942a (씹). A, antennule, ventral view. B, P1, anterior view. C, P2
endopod, anterior view. D, P3 endopod, anterior view. E, P4 endopod, anterior view.
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426
S. SAK ET AL.
margin; enp-2 about half the size of enp-1, with two
geniculate setae and a few spinules.
P2 endopod (Fig. 7C): two-segmented; enp-1 with
few spinules along outer margin; enp-2 short, with a
long, apically serrate, backwardly directed seta near
proximal inner corner, and a long bipinnate inner seta
and a short bare outer spine around distal margin.
P3 endopod (Fig. 7D): two-segmented; enp-1 with
few strong spinules along outer margin; enp-2
minute, with strong spinule at outer distal corner,
short thin seta arising from inner distal corner
(homologous with long inner distal seta of female),
and naked curved apical spine, fused at base
(homologous with outer distal spine of female).
P4 endopod (Fig. 7E): two-segmented; enp-1 with
five strong spinules along outer margin; distal margin
of enp-2 with long, basally fused, serrate seta and
long, unipinnate outer seta.
P2–P4: spine and seta formula as for the genus.
P5 (Fig. 6B): forming subrectangular plate; outer
basal seta sparsely plumose. Free distal margin: with
three short bipinnate spines and one long bipinnate
outer seta; inner spine longer than the other two.
Sixth legs (Fig. 6B): asymmetrical, with smallest P6
closing off functional gonopore; each with a short
inner and a long plumose outer seta.
Remarks: Pennak’s (1942a) illustrations of the male
are restricted to the antennule, and no reference was
made to the sexual dimorphism on the P3 endopod.
His erroneous description of the caudal ramus,
showing a basally swollen seta V and a triangular
process at the distal outer corner, has no doubt been
a source of confusion for subsequent identifications.
Re-examination of a male paratype proved: (1) caudal
ramus seta V to be normally developed, and the terminal spinous process to be much longer and more
sharply pointed; and (2) P1 endopod to be markedly
shorter than the exopod. Both aspects contradict Pennak’s (1942a) description, but agree with Lindgren’s
(1976) observations based on North Carolina specimens. Lindgren also found that the proximal third of
seta V was modified in approximately 60% of the
population; individuals with unmodified setae were
provisionally identified as A. stygia. Given this
intraspecific variability, in conjunction with both
‘populations’ having the same distribution within the
beach, we attribute all North Carolina records of
A. stygia to P. arenaridus. These include Lindgren’s
(1976) intertidal record from west of the Iron Steamer
Pier near Morehead City, and, although provisionally,
Coull’s (1971) subtidal record north of Cape Hatteras
(at a depth of 100 m!). Psammoleptastacus arenaridus
appears to be restricted to the north-eastern Atlantic
seaboard of the USA, from the Woods Hole area in the
north (Pennak, 1942a, b, 1952; Lindgren, 1976) to at
least North Inlet, South Carolina (Coull & Dudley,
1985) in the south. Pennak (1942b) provides data on
the horizontal distribution and relative abundance.
PSAMMOLEPTASTACUS
STYGIUS
(NOODT, 1955B)
COMB. NOV.
Arenopontia stygia Noodt (1955b)
Arenopontia (Arenopontia) stygia Noodt (1955b):
Wells (1967)
Original description: Noodt (1955b): pp. 101–102;
Tafel 35 (figs 75–82) (씸 only).
Type locality: France,
medium coarse sand.
Landes,
Mimizan-Plage;
Remarks: The type material of A. stygia (a single
female) is no longer extant. Noodt’s (1955b) dorsal
view of the caudal ramus shows three setae inserting
at about the same level; the short inner one (adjacent
to seta VII) is not a setal element but the dorsolateral
spur. In addition to the type locality (Delamare
Deboutteville, Gerlach & Siewing, 1955; Noodt,
1955b, c; Delamare Deboutteville, 1960a), P. stygius
has been recorded from the Bassin d’Arcachon,
Gironde (Renaud-Debyser, 1963a, b) and the Portuguese coast (Francelos, south of Porto) (Galhano,
1970). Marinov’s (1971) record from Bulgaria is
attributable to P. barani sp. nov. (see below).
PSAMMOLEPTASTACUS
BARANI SP. NOV.
Arenopontia stygia Noodt, 1955b sensu Marinov
(1971)
Type locality: Turkey, Black Sea coast, Istanbul,
Sahilköy (east of Bosporus); sandy beach.
Material examined: Holotype 씸 (dissected on eight
slides) (BUZM). Paratypes are one 씸 and one 씹 in
alcohol (NHM reg. nos. 2006. 1966–1967), and two
씹씹 dissected on two and seven slides, respectively
(NHM reg. nos. 2006. 1968–1969); all collected at type
locality; leg. S. Karaytuğ and S. Sak, 01 May 2001.
Description
Female: Total body length from tip of rostrum to
posterior margin of caudal rami: 330–380 mm
(mean = 361 mm; N = 7). Maximum width measured
at P5-bearing somite. Body: slender and cylindrical,
without clear distinction between prosome and
urosome (Fig. 8A, B). Hyaline frills of thoracic somites
weakly developed and crenulated; those of genital
double-somite and free abdominal somites strongly
developed, and consisting of rectangular digitate
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CLASSIFICATION OF ARENOPONTIIDAE
427
Figure 8. Psammoleptastacus barani sp. nov. (씸). A, habitus, dorsal view. B, habitus, lateral view. C, anal somite and
caudal rami, dorsal view. D, posterior part of penultimate somite, anal somite, and left caudal ramus, lateral view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
428
S. SAK ET AL.
lappets (Figs 8A, B, 9A, 8D). Genital double-somite
(Fig. 9A): slightly longer than wide; without chitinous
ribs marking original segmentation; with one middorsal, two lateral, and two ventral pores. Anal somite
(Fig. 8C, D): with two dorsal and two lateral pores.
Anal operculum: with minute spinules along free
distal margin (Fig. 8C). Anus: positioned subterminally between caudal rami.
Caudal rami (Fig. 8C, D): approximately 2.75
longer than basal width, tapering posteriorly; with
one pore dorsally, one near ventral proximal margin
(Fig. 9A), and one laterally near outer spinules; outer
distal corner produced into posteriorly directed
recurved spinous process, accompanied by ventral
spinular row at base; dorsomedial surface with
posteriorly directed spinous process arising from base
of seta VII. Armature: as in P. arenaridus, but with a
more foliaceous seta VII.
Rostrum (Figs 8A, 10A): broadly subtriangular,
tapering apically, with two delicate sensillae and one
subapical ventral pore.
Antennule (Fig. 10A, B): six-segmented. Segment 1:
with one seta near anterodistal margin. Segment 2:
longest, about 3.5 times longer than wide. Segment 4:
with long aesthetasc (30-mm long) fused at base with
seta. Distal segment: with seven naked setae (one of
which spatulate), and apical acrothek consisting of
short aesthetasc (13-mm long) and two slender setae.
Armature formula: 1-[1], 2-[7 + 1 plumose], 3-[4],
4-[(1 + ae)], 5-[1], 6-[7 + acrothek].
Antenna (Fig. 10C, D): coxa small, without ornamentation. Original segmentation of allobasis marked
by partial transverse surface suture; with one spinular row along exopodal margin. Exopod: onesegmented, elongate, with long naked seta apically.
Free endopod: with two spinular rows on anterior
surface and finer spinules at outer distal corner;
lateral armature consisting of two short spines; apical
armature consisting of two spines and three geniculate setae, the longest of which with spinules around
geniculation and fused basally to tiny accessory seta.
Mandible, maxillule, maxilla, and maxilliped: as in
A. nesaie.
P1 (Fig. 11A): intercoxal sclerite long and rectangular. Praecoxa: subtriangular and naked. Coxa:
without ornamentation. Basis: with spinular row near
bases of endopod and exopod; anterior surface with a
proximal pore and a small inner basal seta. Exopod:
three-segmented; about 1.2 times longer than
endopod; all segments with spinules along outer
margin; exp-1 longest, with long unipinnate outer
spine; exp-2 without outer element; exp-3 with two
unipinnate outer spines and two geniculate apical
setae. Endopod: two-segmented, not prehensile; enp-1
distinctly longer than exp-1, with a serrate seta at
about two-thirds of the length of the inner margin,
and several spinules along outer margin; enp-2 less
than half the length of enp-1, with two geniculate
setae (inner one about twice as long as outer).
P2–P4 (Fig. 11B–D): intercoxal sclerites naked.
Praecoxae: small and naked. Coxae: rectangular and
without ornamentation. Bases: smaller than coxae,
with a spinular row near base of endopod (P3–P4);
anterior surface with a proximal pore; outer basal seta
absent (P2), plumose (P3), or naked (P4). Exopods:
three-segmented; segments with spinular ornamentation, as illustrated; inner distal seta of exp-3 sparsely
bipinnate, all other elements unipinnate; P3–P4 exp-3
with anterior pore. Endopods: two-segmented, with
few spinules, as illustrated. P2 enp-2: with a long,
apically serrate, backwardly directed seta. Distal
margin of P3 enp-2: with naked outer spine and long
bipinnate inner seta. P4 enp-1: slightly shorter than
exp-1; distal margin of enp-2 with long, basally fused,
serrate seta, and long, unipinnate outer seta. P2–P4:
spine and seta formula as for the genus.
Fifth legs (Fig. 9A): closely set together but not
touching in ventral midline. Baseoendopod and
exopod fused, forming a semicircular plate; distal
margin with two short bipinnate spines flanked by
two long bipinnate setae; outer basal seta long and
sparsely plumose.
Genital field (Fig. 10E): with genital apertures
fused forming median common slit; closed off by fused
P6 forming operculum, with one minute spinous
process on either side; copulatory pore located midventrally, close to genital slit; seminal receptacles
difficult to discern.
Male: Total body length from tip of rostrum to
posterior margin of caudal rami: 290–335 mm
(mean = 315 mm;
N = 6).
Body
ornamentation
(Figs 9B, 12A): essentially as in female. Sexual dimorphism: in antennule, genital segmentation, P3
endopod, P5, and P6. Spermatophore length: approximately 45 mm.
Antennule (Fig. 12C, D): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 2
longest, about 2.2 times as long as wide; segment 4 an
incomplete sclerite with one modified (fused at base)
and one tiny element; segment 5 with long aesthetasc
(45-mm long) fused basally to very small seta;
segments 6–8 with one seta and one basally fused
spiniform element. Segment 9: with one spatulate
seta. Setal formula: 1-[1], 2-[7 + 1 plumose], 3-[4 + 1
pinnate spine], 4-[1 + 1 modified], 5-[2 + (1 + ae)],
6-[1 + 1 modified], 7-[1 + 1 modified], 8-[1 + 1 modified], 9-[7 + acrothek]. Acrothek consisting of short
aesthetasc (13-mm long) fused basally to two slender
setae.
P3 endopod (Fig. 12B): two-segmented; enp-1 with
few strong spinules along outer margin; enp-2
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CLASSIFICATION OF ARENOPONTIIDAE
Figure 9. Psammoleptastacus barani sp. nov. A, urosome 씸, ventral view. B, urosome 씹, ventral view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
429
430
S. SAK ET AL.
Figure 10. Psammoleptastacus barani sp. nov. (씸). A, rostrum and antennule, dorsal view. B, antennule, ventral
view. C, antenna. D, free antennary endopod. E, genital field.
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CLASSIFICATION OF ARENOPONTIIDAE
431
Figure 11. Psammoleptastacus barani sp. nov. (씸). A, P1, anterior view. B, P2, anterior view. C, P3, anterior view.
D, P4, anterior view.
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432
S. SAK ET AL.
Figure 12. Psammoleptastacus barani sp. nov. (씹). A, habitus, dorsal view. B, P3, anterior view. C, antennule,
ventral view. D, antennule, dorsal view.
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CLASSIFICATION OF ARENOPONTIIDAE
433
Table 4. Diagnostic features of Psammoleptastacus species
P1 exopod:endopod*
P3 enp-2 씹 distal spine
P4
Caudal ramus length : basal width
P. arenaridus
P. stygius
P. barani sp. nov.
1.33
fused
exp-1 < enp-1
2.75
1.28
?
exp-1 < enp-1
3.85
1.15
discrete
exp-1 > enp-1
2.75
*Length of rami calculated as sum of segment lengths.
minute, with strong spinule at outer distal corner,
short fine seta arising from inner distal corner
(homologous with long inner distal seta of female),
and naked curved apical spine, discrete at base
(homologous with outer distal spine of female).
P5 (Fig. 9B): with armature as in female, but with
innermost element of distal margin much shorter and
bare.
Sixth legs (Fig. 9B): asymmetrical, with smallest P6
closing off functional gonopore; each with two pinnate
setae.
Etymology: The species is named after Prof. İbrahim
Baran, Dokuz Eylül University, in recognition of his
contributions to herpetology in Turkey.
Remarks: Psammoleptastacus barani sp. nov. differs
from its two known congeners in the longer P1
endopod, and the length of P4 enp-2, which is shorter
than, or at most as long as, the proximal exopod
segment, instead of being distinctly longer. It is
similar to P. arenaridus in the length of the caudal
ramus, but deviates from it in the detailed morphology of the male P3 endopod. Marinov (1971) recorded
a few specimens from the beaches of the Arkutino
region (south of Sozopol) along the Bulgarian coast,
which he attributed to A. stygia. He noted the difference in the relative length of the P1 endopod between
his specimens and Noodt’s (1955b) type population of
P. stygius. The similarity in the relative length of the
P1 endopod, P4 endopod, and caudal ramus, the P3
endopodal sexual dimorphism, and the female P5,
leave little doubt that Marinov (1971) was dealing
with P. barani sp. nov. The only discrepancy between
both descriptions is found in the male P5, which has
a much longer innermost seta in the Bulgarian material. Apostolov (1973) claimed that considerable variability was found in the P5 of his Black Sea
A. subterranea (no localities specified), but it is conceivable that his drawings of the female P5 were
based on two different species: his Figure 18-5 shows
a P5 of the Arenopontia type, whereas his Figure 18-6
was almost certainly based on the species previously
identified by Marinov (1971) as A. stygia. Note that
the type locality of P. barani sp. nov. is in close proximity to the Bulgarian collecting sites.
ARENOPONTIA SUBTERRANEA KUNZ, 1937
RAO & GANAPATI (1969)
SENSU
?Arenopontia subterranea Kunz (1937) sensu Rao
(1967)
Original description: Rao & Ganapati (1969): pp. 268–
269; figure 5.
Type locality: India, Andhra Pradesh, Vishakhapatnam, Waltair; sandy beach, medium sand.
Remarks: Rao & Ganapati (1969) stated that only
minor variations occurred between their specimens
from Waltair (India) and Kunz’ 1937 type material of
A. subterranea, listing examples such as the caudal
ramus seta VII, which is not foliaceous, the inner seta
on P1 exp-3, which is not modified, and the serrate
nature of the inner setae on enp-2 of P2 and P4.
However, their figure of the P1, showing a nonprehensile endopod and the absence of the penicillate
seta on exp-3, unequivocally excludes the Indian
specimens from the genus Arenopontia. The presence
of two geniculate setae on P1 enp-2, in conjunction
with the presence of the inner serrate seta on P2
enp-2, suggests a relationship with the genus Psammoleptastacus. Pending examination of new material,
Rao & Ganapati’s (1969) species is considered species
inquirenda in the latter genus. Provided their figures
of the fifth legs are correct, the presence of only four
elements on this limb is thus far unique within the
genus. The presence of two setae on the antennary
exopod and the absence of the outer distal element on
P2 enp-2 require confirmation. Rao’s (1967) record of
A. subterranea from Palm Beach, Waltair, conceivably
refers to the same species.
Key to species
A straightforward dichotomous key is impossible to
construct. Differences between species are subtle at
best (Table 4), and any identification should be
checked against the relevant descriptions.
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434
S. SAK ET AL.
GENUS NEOLEPTASTACUS NICHOLLS, 1945
The genus Neoleptastacus has had an intricate taxonomic history since its proposal by Nicholls (1945). It
remained monotypic until Chappuis (1955), and not
Kunz (1954), as claimed by Noodt (1955a), relegated
it to a junior synonym of Arenopontia. This course of
action was forecasted by Chappuis’ (1954b) statement
that Nicholls’ new genus Paraleptastacus [sic] should
be united with Arenopontia. Krishnaswamy (1957)
argued against this inclusion, and maintained the
validity of Neoleptastacus as a distinct genus; he also
added a second species, Neoleptastacus secundus,
from the Madras coast. Wells (1967) dismissed Krishnaswamy’s (1957) decision, and retained Neoleptastacus as a subgenus of Arenopontia. Following some
initial criticism by Mielke (1982a), the subgeneric
classification was abolished by Martínez Arbizu &
Moura (1994), and Neoleptastacus was synonymized
with Arenopontia. Huys et al. (1996b) and Bodin
(1997) reinstated both subgenera, but Wells (2007)
preferred to amalgamate them, and consequently
synonymized Neoleptastacus for the third time. The
genus is resurrected here and redefined to accommodate: (1) all species that have previously been allocated to the subgenus Neoleptastacus (see Karanovic,
2000); (2) Mielke’s (1985) Chilean species A. clasingi,
A. pacifica, and A. spicata; and (3) the two species
formerly included in the genus Pararenopontia
[Pararenopontia breviarticulata (Mielke, 1975) and
Pararenopontia trisetosa (Mielke, 1982a)]. A detailed
review of the genus, including new species and an
updated key, will be published elsewhere (Sak, Huys
& Karaytuǧ, in press b).
Diagnosis: Arenopontiidae. Urosomites: occasionally
with conspicuous surface ornamentation (Neoleptastacus clasingi, Neoleptastacus ornamentus, and
Neoleptastacus reductaspina). Anal somite: with
(acanthus lineage) or without (all other lineages)
paired dorsolateral spinous processes. Anal operculum: sometimes with median extension. Hyaline frills
of abdominal somites with rectangular digitate or
nondigitate lappets. Caudal ramus: usually with dorsolateral spur near medial margin. P1 exopod: two- or
three-segmented; exp-1 with/without outer spine;
exp-3 (or exp-2 when exopod two-segmented) with one
or two spine(s) and two geniculate setae. P1 endopod:
not prehensile, at least as long as exopod; enp-2 with
two geniculate setae (Neoleptastacus speluncae and
Neoleptastacus phreaticus) or outer spine plus inner
geniculate seta (all other species). P2–P3 endopods:
one- or two-segmented. P3 endopod: with outer distal
element usually fused at base. P4 endopod: with welldeveloped outer distal element (except in trisetosus
lineage). Armature formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.(0–1)21
Endopod
0.(0–1)(1–2)0 or 110
0.0(1–2)0 or 010
0.020
P3 endopod 씹: not sexually dimorphic. P5 with
outer basal seta and between one and three discrete
elements; innermost element fused to segment
forming spinous process (weakly delimited in Neoleptastacus trisetosus); length of process sometimes
sexually dimorphic. P6 씹 with one or two seta(e).
Type species: Neoleptastacus spinicaudatus Nicholls
(1945) (by monotypy).
Other species: Arenopontia australis Chappuis, 1952 =
Neoleptastacus australis (Chappuis, 1952) comb.
nov.; Arenopontia acantha Chappuis, 1954b =
Neoleptastacus acanthus (Chappuis, 1954b) comb.
nov.; Arenopontia longiremis Chappuis, 1955 = Neoleptastacus longiremis (Chappuis, 1955) comb. nov.;
N. secundus Krishnaswamy, 1957; Arenopontia
africana Chappuis & Rouch, 1961 = Neoleptastacus
africanus (Chappuis & Rouch, 1961) comb. nov.;
Arenopontia accraensis Lang, 1965 = Neoleptastacus
accraensis (Lang, 1965) comb. nov.; Arenopontia
indica Rao, 1967 = Neoleptastacus indicus (Rao,
1967) comb. nov.; Arenopontia ishikariana Itô, 1968 =
Neoleptastacus ishikarianus (Itô, 1968) comb. nov.;
A. (Neoleptastacus) africana f. angolensis Kunz, 1971 =
Neoleptastacus angolensis (Kunz, 1971) comb. nov.;
Arenopontia gussoae Cottarelli, 1973 = Neoleptastacus
gussoae (Cottarelli, 1973) comb. nov.; Arenopontia
trisetosa Mielke, 1982a = Neoleptastacus trisetosus
(Mielke,
1982a) comb. nov.;
Arenopontia clasingi
Mielke, 1985 = Neoleptastacus clasingi (Mielke,
1985) comb. nov.; Arenopontia pacifica Mielke, 1985 =
Neoleptastacus pacificus (Mielke, 1985) comb. nov.;
Arenopontia spicata Mielke, 1985 = Neoleptastacus
spicatus (Mielke, 1985) comb. nov.; Arenopontia chaufriassei Bodiou & Colomines, 1986 = Neoleptastacus
chaufriassei (Bodiou & Colomines, 1986) comb. nov.;
Arenopontia ornamenta Mielke, 1987 = Neoleptastacus
ornamentus (Mielke, 1987) comb. nov.; Arenopontia
reductaspina Mielke, 1987 = Neoleptastacus reductaspina (Mielke, 1987) comb. nov.; Arenopontia
(Neoleptastacus) phreatica Cottarelli et al., 1994 =
Neoleptastacus phreaticus (Cottarelli et al., 1994)
comb. nov.; Arenopontia (Neoleptastacus) speluncae
Cottarelli
et al.,
1994 = Neoleptastacus speluncae
(Cottarelli et al., 1994) comb. nov.; Arenopontia
(Neoleptastacus) huysi Karanovic, 2000 = Neoleptastacus huysi (Karanovic, 2000) comb. nov.
Species
inquirendae: Arenopontia
?
gussoae
Cottarelli, 1973 sensu Mielke (1982b); Arenopontia ?
ishikariana Itô, 1968 sensu Mielke (1987).
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
CLASSIFICATION OF ARENOPONTIIDAE
Species incertae sedis: Arenopontia breviarticulata
Mielke, 1975 (see below).
NEOLEPTASTACUS
SPINICAUDATUS
NICHOLLS, 1945
Original description: Nicholls (1945): pp. 22–23; textfigure 3.
Type locality: Nicholls (1945) collected the species
from two sandy beaches some 500 km apart on the
coast of Western Australia; the first, Leighton Beach
(in the region of Perth), was sampled in October 1939;
samples from the second site, locally known as ‘Back
Beach’ at Dongarra, were taken in March 1940.
Nicholls did not specify a type locality; however, all
NHM syntypes came from sand washings at the
Dongarra site, which is regarded here as the locus
typicus.
Material examined: Syntype series consisting of:
(1) one 씸 dissected on one slide (NHM reg. no.
1947.10.3.8); (2) one 씹 dissected on one slide (NHM
reg. no. 1947.10.3.9); and (3) one vial containing 25
씸씸 and one 씹 in alcohol [besides a mixture of Psammopsyllus operculatus Nicholls, 1945 (one 씸 and one
씹), Ectinosomoides longipes Nicholls, 1945 (one 씸),
and a new species of arenopontiid described below as
Onychopontia nichollsi (ten 씸씸 and one 씹)]; leg. A.G.
Nicholls on 25–26 March 1940.
Redescription
Female: Total body length from tip of rostrum to
posterior margin of caudal rami: 274–295 mm
(mean = 281 mm, N = 15). Maximum width: 33 mm
(mean of 15 individuals = 32 mm), measured at posterior margin of cephalothorax. Body: slender and cylindrical, without clear distinction between prosome and
urosome (Fig. 13A, B). Hyaline frills of thoracic
somites weakly developed and crenulated; those of
genital double-somite and free abdominal somites
strongly developed and consisting of rectangular digitate lappets (Figs 13A, B, 14A, B, E). Genital doublesomite (Figs 13A, B, 14A) as long as wide; without
chitinous ribs marking original segmentation; with
one middorsal, two lateral, and six ventral pores
(Fig. 15F). Anal somite (Fig. 14A, C, E): with two
ventral and two lateral pores; ventral posterior
margin with medial spinule rows. Anal operculum:
smooth (Fig. 14F). Anus positioned subterminally
between caudal rami.
Caudal rami (Fig. 14C–F): approximately 2.2 times
longer than maximum width (measured in dorsal
view), tapering posteriorly; with one pore near ventral
proximal margin (Fig. 14E), and two pores laterally
near insertion site of seta II, and at base of distal
spinous process (Fig. 14C); outer distal corner pro-
435
duced into posteriorly directed recurved spinous
process; mediodorsal surface with small spur-like
process at base of seta VII, accompanied by minute
accessory process near inner margin (Fig. 14F). Armature consisting of seven setae: seta I, small; setae II
and III, long and naked, the latter displaced dorsally;
seta IV, short, sparsely pinnate, located between
seta V and spinous process; seta V, long, and with
fracture plane; seta VI small, naked, and located at
inner distal corner; seta VII, not foliaceous and triarticulate at base.
Rostrum (Fig. 16B): small, broadly subtriangular,
apical part lobate and offset, with two delicate sensillae and one midventral pore.
Antennule (Fig. 16A): long, six-segmented. Segment 1: with a tiny seta near anterodistal margin.
Segment 2: longest, about 2.5 times longer than wide.
Segment 4: with long aesthetasc (25-mm long), fused at
base with seta. Distal segment: with seven naked setae
(two of which are spatulate), and with apical acrothek
consisting of short aesthetasc (10-mm long) and two
slender setae. Armature formula: 1-[1], 2-[7 + 1
plumose], 3-[4], 4-[(1 + ae)], 5-[1], 6-[7 + acrothek].
Antenna (Fig. 17D): coxa, small and bare. Basis and
proximal endopod segment incompletely separated by
surface suture, without armature or ornamentation.
Exopod: minute, one-segmented, with one long bare
seta. Free endopod: with distal hyaline frill; abexopodal margin with two spinular rows and two bare,
curved spines; apical margin with two geniculate
setae, two bare spines and one composite element
consisting of spinulose, geniculate seta and small,
basally fused seta.
Labrum (Fig. 16C): sclerotized and wide; with blunt
spinules bilaterally around distal margin.
Mandible (Fig. 17E): gnathobase elongate, about as
long as palp; with several curved, minute teeth and
one tiny recurved seta at dorsal corner. Palp consisting of elongate, unisetose basis and one-segmented
endopod with one inner, two outer, and two apical
setae.
Maxillule (Fig. 16D): praecoxal arthrite with one
surface seta; distal margin with five spines and two
setae. Coxal endite: cylindrical, with two recurved
spines. Basis: elongate, with rami completely incorporated; basal armature consisting of three apical
setae. Exopod and endopod: represented by one and
three setae, respectively.
Maxilla (Fig. 17F): syncoxa with two cylindrical
endites; proximal endite with three setae (one fused
at base); distal endite with two setae (one fused at
base). Allobasis: drawn out into long claw with one
accessory seta. Endopod: one-segmented; with three
setae. All elements are naked.
Maxilliped (Fig. 16E): syncoxa longer than wide,
unarmed, with few spinules. Basis: elongate and
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S. SAK ET AL.
Figure 13. Neoleptastacus spinicaudatus Nicholls, 1945 (씸). A, habitus, dorsal view. B, habitus ovigerous specimen,
lateral view.
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CLASSIFICATION OF ARENOPONTIIDAE
437
Figure 14. Neoleptastacus spinicaudatus Nicholls, 1945. A, urosome 씸, ventral view. B, urosome 씹, lateral view. C, anal
somite and right caudal ramus 씸, outer lateral view. D, left caudal ramus 씸, inner lateral view (dorsal spur indicated
by an arrow); E, anal somite and left caudal ramus 씸, ventral view. F, right caudal ramus 씸, dorsal view.
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S. SAK ET AL.
Figure 15. Neoleptastacus spinicaudatus Nicholls, 1945 (씸). A, P1, anterior view. B, P2, anterior view. C, P3, anterior
view. D, P4, anterior view. E, P5, anterior view. F, genital field (copulatory pore indicated by arrow).
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CLASSIFICATION OF ARENOPONTIIDAE
unarmed. Endopod: with small accessory seta and
slightly curved claw, bearing strong, subterminal
spinule.
P1 (Fig. 15A): intercoxal sclerite long and rectangular. Praecoxa: triangular and naked. Coxa: with few
spinules on posterior surface. Basis: with spinules
around base of endopod, and at inner and outer distal
corners; anterior surface with a pore and a small
inner seta. Exopod: three-segmented; exp-1 and exp-2
with spinules around outer margin; exp-1 longest,
with short, bare outer spine; exp-2 without outer
element; exp-3 with short, bare outer spine, a curved
unipinnate spine, and two geniculate setae distally.
Endopod: two-segmented, not prehensile, as long as
exopod; enp-1 with a serrate inner seta at about
halfway along the segment length, and coarse
spinules along outer margin; enp-2 slightly shorter
than enp-1, with a short bare outer spine and a
geniculate inner seta.
P2–P4 (Fig. 15B–D): intercoxal sclerites naked.
Praecoxae: very small and bare. Coxae: squarish
and without ornamentation. Bases: smaller than
coxae, with a spinular row near base of endopod,
and a few spinules around outer corner in P2 and
P4; anterior surface with a pore in P2 and P4; outer
basal seta absent in P2, but present and bare in
P3–P4. Exopods: three-segmented; segments with
coarse spinular ornamentation, as illustrated; inner
distal seta of exp-3 sparsely bipinnate, all other elements unipinnate or bare, except for inner seta of
P4 exp-3 being serrate. Endopods: two-segmented;
P2–P4 enp-1 about 1.1, 2.3, and 3.8 times longer
than their respective distal segments, with coarse
spinules along outer margin, and a few tiny
spinules near inner distal corner. P2 enp-2: with a
long, apically serrate, backwardly directed seta near
proximal margin. P3 enp-2: with a short bipinnate
spine apically and outer distal spine fused to
segment forming bare spinous process. P4 enp-2:
with large apically serrate seta, fused at base, and
short unipinnate seta at outer distal corner. Spine
and seta formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.121
Endopod
0.120
0.020
0.020
Fifth legs (Fig. 15E): closely set together, but not
touching in ventral midline. Baseoendopod and
exopod fused, forming a rectangular plate; anterior
surface with two pores. Inner distal corner: with
strong spinous process (homologous to inner spine);
process minutely bipinnate, with subapical flagella,
and delimited at base on posterior surface. Distal
margin with plumose outer basal seta, one naked
seta, and two short, equally long, bipinnate spines.
439
Genital field positioned centrally on ventral surface
of genital double-somite (Fig. 14A). Genital apertures
(Fig. 15F) fused, forming median common slit; closed
off by fused P6 forming operculum with two minute
spinous processes on either side; copulatory pore
large (arrowed in Fig. 15F), leading to short copulatory duct, and surrounded by three pairs of pores;
seminal receptacles difficult to discern.
Male: Total body length from tip of rostrum to posterior margin of caudal rami: 272–277 mm (N = 2).
Maximum width: 30–31 mm (N = 2), measured at
cephalothorax. Body ornamentation (Fig. 14B): essentially as in female. Sexual dimorphism: in antennule,
genital segmentation, P5 (weak), and P6. Spermatophore length: approximately 30 mm.
Antennule (Fig. 17A–C): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 1
with few spinules on anterior surface; segment 2
longest and about 1.9 times longer than wide; segment 4 an incomplete sclerite with one tiny element;
segment 5 with two setae, plus long aesthetasc
(43-mm long), fused basally to a slender seta; segment 6 with two short setae; segment 7 with two
modified spines and a seta; segment 8 with three
modified spines and posterior corner produced into
lobate extension; distal segment with seven naked
setae (two of which spatulate) and apical acrothek.
Setal formula: 1-[1], 2-[6 + 1 plumose], 3-[4 + 2
pinnate spines], 4-[1], 5-[2 + (1 + ae)], 6-[2], 7-[1 + 2
modified], 8-[1 + 3 modified], 9-[7 + acrothek]. Acrothek consisting of short aesthetasc (15-mm long) fused
basally to two slender setae.
P5 (Fig. 16F): with armature as in female; inner
spinous process, without subapical flagella and leg
slightly more slender than in female.
Sixth legs (Fig. 16F) asymmetrical, with smallest
P6 closing off functional gonopore; each with a
long, plumose, outer seta and a short, naked, inner
spine.
Remarks: Nicholls (1945) overlooked the inner seta
on P4 exp-3, an error that perpetuated in various
comparative analyses (Noodt, 1955b; Bodiou & Colomines, 1986) and species keys (Lang, 1965; Karanovic, 2000). Within the genus, N. spinicaudatus
belongs to a lineage that is characterized by: (1) anal
somite without paired dorsolateral processes; (2) anal
operculum weakly developed, without rounded medial
extension; (3) P1 exp-1 with outer spine, exp-3 with
four setae/spines; (4) P1 enp-2 with outer spine and
inner geniculate seta distally; (5) P2 exp-2 with outer
spine of normal length (not extending far beyond
distal margin of exp-3); (6) endopod P2–P3, twosegmented; (7) P2 enp-2 with inner seta and two
distal spines; (8) P3 enp-2 with two distal spines
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S. SAK ET AL.
Figure 16. Neoleptastacus spinicaudatus Nicholls, 1945. A, antennule 씸, dorsal view. B, rostrum and proximal antennulary segments 씸, dorsal view. C, labrum 씸, anterior view. D, maxillule 씸, posterior view. E, maxilliped 씸. F, P5-bearing
and genital somites (씹), ventral view.
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CLASSIFICATION OF ARENOPONTIIDAE
441
Figure 17. Neoleptastacus spinicaudatus Nicholls, 1945. A and C, antennule 씹, anterior view (at different angles, with
armature largely omitted in C). B, antennule 씹, ventral view. D, antenna 씸. E, mandible 씸. F, maxilla 씸.
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442
S. SAK ET AL.
KEY
TO SPECIES OF SPINICAUDATUS LINEAGE
1. P4 exp-3: without inner seta ................................................................................................. N. pacificus.
P4 exp-3: with inner seta ................................................................................................................... 2.
2. Urosome (except anal somite): with distinct surface ornamentation consisting of elongate rectangular plates ......
.........................................................................................................................................N. clasingi.
Urosome: without conspicuous surface ornamentation ............................................................................. 3.
3. Caudal ramus: with dorsolateral spur near base of seta VII.....................................................................4.
Caudal ramus: without dorsolateral spur near base of seta VII..............................................N. ishikarianus.
4. Lappets of abdominal hyaline frills semi-incised obtusidigitate; inner seta of P2–P3 enp-2 longer than endopod; P5
3.0 times as long as wide, with naked spinous process...............................................................N. spicatus.
Lappets of abdominal hyaline frills denticulate; inner seta of P2–P3 enp-2 shorter than endopod; P5 about 2.5 times
as long as wide, with pinnate spinous process .................................................................. N. spinicaudatus.
(outer one fused to segment); and (9) P4 enp-2 outer
seta normally developed. In addition to the type
species, this spinicaudatus lineage includes N. ishikarianus, N. clasingi, N. pacificus and N. spicatus,
all of which assume a Pacific distribution. Neoleptastacus spinicaudatus is very similar to the
Chilean N. spicatus, but differs from it in the form of
the abdominal hyaline frills (lappets denticulate vs.
semi-incised obtusidigitate in N. spicatus), the relative lengths of the inner distal seta on P2–P3 enp-2,
and the shape and ornamentation of the spinous
process on P5 (which is still delimited at base on the
posterior surface).
The linear egg sac contains between two and four
large eggs; occasionally six eggs are found, in which
case they overlap. Nicholls (1945) also collected the
species from Leighton Beach (in the region of Perth).
Chappuis’ (1958) record from Puget Sound almost
certainly pertains to Mesopontia dillonbeachia (see
below). Chappuis (1954b: 269) claimed to have found
one female and one male in Annaba (= Bône), Algeria,
which closely resembled N. spinicaudatus.
GENUS PARARENOPONTIA BODIOU & COLOMINES,
1986
Bodiou & Colomines (1986) established this genus to
accommodate two unusual Arenopontia species with a
two-segmented P1 exopod, A. breviarticulata Mielke
(1975) (type species) and A. trisetosa Mielke (1982a);
the third species (Arenopontia biarticulata Wells,
1967) displaying this character was placed in the
genus Notopontia Bodiou in the Leptopontiidae. Both
Pararenopontia species have very few characters in
common, casting doubt on the monophyletic status of
the genus. Martínez Arbizu & Moura (1994) considered Pararenopontia an amalgam of species sharing
reduced leg segmentation, and synonymized it with
Arenopontia; however, some authors have suggested
that it should be maintained as a valid genus (Huys
et al., 1996a, b; Bruno et al., 1998; Wells, 2007) or
subgenus (Bodin, 1997).
Pararenopontia breviarticulata is known from a
single male collected from the Isle of Sylt (Germany)
(Mielke, 1975), and exhibits an interesting mosaic of
both apomorphic (two-segmented P1 exopod; reduced
P5) and plesiomorphic (armature of P2 endopod and
P4 exopod) characters. There is, however, some circumstantial evidence that Mielke’s (1975) description
is deficient in some aspects. The P3 appears remarkably similar to the P4, including the presence of an
inner seta on exp-3 (a feature not reported for any
other arenopontiid) and two very long setae on enp-2
(not recorded elsewhere in the family), raising the
suspicion that the author has not observed the real
P3, but may instead have duplicated observations of
P4. The male P5 is unique in possessing only three
elements; it is not clear whether the inner spine (or
spinous process) was overlooked, or whether the P5 is
genuinely underdeveloped as a result of paedomorphosis. Evidence for the latter is found in the congruence between Mielke’s illustration and the condition
observed in copepodid IV of N. indicus (cf. Rao, 1967:
fig. 3-22), at which stage the P1 exopod in arenopontiids is still two-segmented before adding a final
segment at the next moult. Mielke (1975) described
the anal operculum with two lateral ‘Zacken’ (prongs,
teeth), which are conceivably the positional homologues of the paired lateral spinous processes on the
anal somite in the acanthus group of Neoleptastacus.
This group includes N. acanthus, N. longiremis, N. secundus, N. indicus, N. gussoae, N. chaufriassei, N. ornamentus, N. reductaspina, and N. huysi. In all these
species, the P3 endopod has an inner distal seta; the
outer distal spine is either short, and fused to the
segment, or completely absent (longiremis, gussoae,
indicus, and reductaspina), but never setiform, and is
virtually as long as the outer distal seta [as illustrated by Mielke (1975) for P. breviarticulata]. The
extreme disparity in the length of the outer basal seta
between P3 and P4, and the very long outer spines on
P2–P4 exp-2 are additional characters unique to this
species. The armature formula of P1 enp-2 (one geniculate seta plus one outer distal spine) indicates
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CLASSIFICATION OF ARENOPONTIIDAE
a relationship with the Mesopontia–Arenopontia–
Neoleptastacus lineage, and the morphology of the
anal somite suggests that P. breviarticulata is probably nested within the genus Neoleptastacus. Unfortunately, repeated requests to make the holotype
available for re-examination failed, and its relationships necessarily remain unresolved. Pending the collection of topotype material, we propose to regard
Pararenopontia as a junior subjective synonym of
Neoleptastacus, and A. breviarticulata as a species
incertae sedis in Neoleptastacus. Note that Bodin
(1979) had already listed it under the subgenus
Neoleptastacus.
Pararenopontia trisetosa is clearly closely related to
N. africanus and N. angolensis, the only marked difference being the two-segmented P1 exopod in P. trisetosa. All three species differ from other members of
Neoleptastacus by the lack of the outer spine on P1
exp-1, the presence of only one outer spine on the
distal exopod segment of P1 (= exp-2 in P. trisetosa;
exp-3 in N. africanus and N. angolensis), the onesegmented P2–P3 endopod, and the reduced setal
formula on P2 endopod [110]. Based on these apomorphic character states, P. trisetosa is formally transferred to Neoleptastacus as N. trisetosus comb. nov.
GENUS MESOPONTIA GEN. NOV.
Lang (1965) noted that there is no close affinity
between A. dillonbeachia and A. subterranea, but
believed the former was ‘. . . most nearly related to
the species A. acantha and A. secunda. . .’, both of
which currently belong to the acanthus lineage of
Neoleptastacus. Arenopontia dillonbeachia shares
with Arenopontia (as redefined herein) and Neoleptastacus the presence of an outer spine and an
inner geniculate seta/claw on P1 enp-2, and the
absence of sexual dimorphism on the P3 endopod;
however, the morphology of the P5 indicates an intermediate position between both genera. In Arenopontia, the innermost element is setiform and defined at
the base (as in Psammoleptastacus and Onychopontia
gen. nov.); in A. dillonbeachia it is modified into a
strong articulating spine, whereas in Neoleptastacus
the inner corner of the P5 is modified into a spinous
process. The pinnate nature of this process and the
presence of a posterior surface suture in some species
(e.g. N. spinicaudatus; Fig. 15E) indicate the Neoleptastacus condition originated from the incorporation of a spinous inner element (as expressed in
A. dillonbeachia). The transitionary state of the P5 in
conjunction with the nonprehensile P1 exclude A. dillonbeachia from both Arenopontia and Neoleptastacus, and consequently it is here designated as the
type of a new genus. A unique feature for the genus is
the bicuspidate dorsomedial process on the caudal
ramus.
443
Diagnosis: Arenopontiidae. Urosomites: without conspicuous surface ornamentation. Anal somite: without
paired dorsolateral spinous processes. Anal operculum: not modified. Hyaline frills of abdominal somites
with narrow rectangular lappets. Caudal ramus:
without dorsolateral spur, but with bicuspidate
process near medial margin. P1 exopod: threesegmented, short; exp-1 longest, with outer spine;
exp-3 with two spines and two geniculate setae. P1
endopod: not prehensile, about as long as exopod;
enp-2 with outer spine and inner geniculate seta.
P2–P3 endopods: two-segmented; inner serrate seta of
P2 enp-2 present. P3 endopod: with outer distal
element. P4 endopod: with outer distal element well
developed. Armature formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.121
Endopod
0.120
0.020
0.020
P3 endopod male: not sexually dimorphic, twosegmented. P5: with outer basal seta and four discrete elements in both sexes; innermost element a
strong bipinnate spine. P6 male: with two setae.
Type and only species: Arenopontia dillonbeachia
Lang, 1965 = Mesopontia dillonbeachia (Lang, 1965)
comb. nov.
Etymology: The generic name is derived from the
Greek mesos (mesos meaning the middle), and the
suffix pontia (pontia, meaning the sea), commonly
used in the formation of interstitial copepod names,
and refers to the morphology of the P5, which exhibits
a transitionary state between the Arenopontia condition (innermost element setiform) and the Neoleptastacus condition (inner spinous process).
MESOPONTIA DILLONBEACHIA
(LANG, 1965) COMB. NOV.
Arenopontia dillonbeachia Lang, 1965
Arenopontia (Arenopontia) dillonbeachia Lang (1965):
Wells (1967)
Arenopontia spinicaudata (Nicholls, 1945) sensu
Chappuis (1958)
Original description:
figure 231 (씸 only).
Lang
(1965):
pp. 419–422;
Type locality: USA, California, Dillon Beach; about
3 m in depth, fine sand.
Material examined: SMNH: (1) one 씸 (syntype), from
type locality; reg. no. 568 (Typ. Saml. 2210), leg. K.
Lang, 30 August 1960; (2) five 씸씸 (syntypes), Cali-
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S. SAK ET AL.
fornia, Monterey Bay, off Hopkins Marine Laboratory,
fine shell-sand, reg. no. 569 (Typ. Saml. 2211), leg. K.
Lang, 12 September 1960.
Redescription
Female: Total body length from tip of rostrum to
posterior margin of caudal rami: 337 mm. Maximum
width: 41 mm, measured at posterior margin of cephalothorax. Body: slender and cylindrical, without clear
distinction between prosome and urosome (Fig. 18A,
B). Hyaline frills of thoracic somites weakly developed
and crenulated; those of genital double-somite and
free abdominal somites strongly developed, and
consisting of narrow, rectangular, digitate lappets
(Figs 18A–C, 19A). Genital double-somite (Fig. 19A):
1.2 times longer than wide; without chitinous ribs
marking original segmentation; with two middorsal,
two lateral, and two ventral pores. Anal somite
(Figs 18C, D, 19A): with two lateral pores. Anal operculum: with minute spinules along free distal margin
(Fig. 18C). Anus positioned subterminally between
caudal rami.
Caudal rami (Figs 18C, D, 19A): approximately 2.7
times longer than maximum width (measured in
dorsal view), tapering both proximally and distally;
with a pore laterally; outer distal corner produced
into posteriorly directed, recurved, spinous process;
dorsal surface with bicuspidate process near inner
margin (Fig. 18C). Armature consisting of seven
setae: seta I, small; setae II and III, long and naked;
seta IV, short, located between seta V and spinous
process, naked; seta V, long and with fracture plane;
seta VI, small, naked, and located at inner distal
corner; seta VII, spatulate and triarticulate at base.
Rostrum (Fig. 19C): small, broadly subtriangular,
apical part lobate and offset, with two delicate sensillae and a midventral pore near apex.
Antennule (Fig. 19B, C): long, six-segmented. Segment 2: longest, about 2.6 times longer than wide.
Segment 4: with long aesthetasc (37-mm long) fused at
base with seta. Distal segment: with seven naked setae
(two of which are spatulate), and apical acrothek
consisting of short aesthetasc (17-mm long) and two
slender setae. Armature formula: 1-[1], 2-[7 + 1
plumose], 3-[4], 4-[(1 + ae)], 5-[1], 6-[8 + acrothek].
Antenna (Fig. 19D): coxa small, without ornamentation. Allobasis about 2.7 times as long as maximum
width; original segmentation marked by partial
transverse surface suture; with two spinular rows, as
illustrated. Exopod: one-segmented; elongate, with a
naked apical seta (about 3.0 times longer than
exopod). Free endopod with two spinular rows on
anterior surface, and finer spinules at outer distal
corner; lateral armature consisting of two short
spines; apical armature consisting of two spiniform
elements and three geniculate setae, the longest of
which with spinules around geniculation and fused
basally to tiny accessory seta.
Mandible, maxillule, maxilla, and maxilliped, as in
N. spinicaudatus (see Lang, 1965: Fig. 231d–g).
P1 (Fig. 20A): intercoxal sclerite, long with deeply
concave ventral margin. Praecoxa: minute. Coxa:
without ornamentation. Basis: with spinular row near
base of endopod and at inner distal corner; anterior
surface with small inner seta. Exopod: threesegmented; all segments with spinules around outer
margin; exp-1 longest, with strong, unipinnate outer
spine; exp-2 without outer element; exp-3 with short
naked outer spine, a long curved unipinnate spine,
and two geniculate setae distally. Endopod: twosegmented, not prehensile, about as long as exopod;
enp-1 2.6 times longer than maximum width; with a
serrate inner seta halfway along the segment length,
and several spinules along outer margin; enp-2 about
twice as long as wide, with a short, naked outer spine,
and a long geniculate inner seta.
P2–P4 (Fig. 20B–D): intercoxal sclerites naked,
with deeply concave ventral margin. Praecoxae: small
and naked. Coxae: squarish and without ornamentation. Bases: smaller than coxae, with a spinular row
near base of endopod (P3) or near outer distal corner
(P2, P4); anterior surface with a pore in P3–P4; outer
basal seta absent (P2), plumose (P3), or naked (P4).
Exopods: three-segmented; segments with spinular
ornamentation, as illustrated; inner distal seta of
exp-3 sparsely bipinnate, all other elements unipinnate except for inner seta of P4 exp-3 being apically
serrate; P4 exp-3 with anterior pore. Endopods: twosegmented; P2–P4 enp-1 about 1.3, 1.8, and 3.7 times
longer than their respective distal segments, with few
spinules, as illustrated. P2 enp-2: with a long, apically serrate, backwardly directed seta near proximal
inner corner. P3 enp-2: with a long bipinnate inner
seta, and a short bare outer spine apically. P4 enp-2:
with long, apically serrate inner seta (fused at base),
and long bipinnate seta at outer distal corner. Spine
and seta formula: as for the genus.
Fifth legs (Fig. 19A) closely set together, but not
touching in ventral midline. Baseoendopod and
exopod: fused, forming a subrectangular plate; anterior surface with pore near outer proximal corner;
distal margin with outer naked seta, two medial short
setae, and strong bipinnate spine at inner corner;
outer basal seta long and plumose.
Remarks: Lang (1965) remarked on the apparent
similiarity in antennary exopod morphology between
A. dillonbeachia and Pennak’s (1942a) P. arenaridus;
however, our re-examination and Itô’s (1969) description failed to confirm the very long and slender shape
of the exopod (as well as the completely separated
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CLASSIFICATION OF ARENOPONTIIDAE
445
Figure 18. Mesopontia dillonbeachia (Lang, 1965) comb. nov. (씸). A, habitus, dorsal view. B, habitus ovigerous specimen, lateral view. C, posterior part of penultimate somite, anal somite, and caudal rami, dorsal view. D, anal somite and
right caudal ramus, lateral view.
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446
S. SAK ET AL.
Figure 19. Mesopontia dillonbeachia (Lang, 1965) comb. nov. (씸). A, urosome, ventral view. B, antennule, ventral view.
C, rostrum and antennule, dorsal view. D, antenna.
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CLASSIFICATION OF ARENOPONTIIDAE
447
Figure 20. Mesopontia dillonbeachia (Lang, 1965) comb. nov. (씸). A, P1, anterior view. B, P2, anterior view. C, P3,
anterior view. D, P4, anterior view.
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448
S. SAK ET AL.
basis). Both Lang (1965) and Itô (1969) overlooked the
hyaline frills on the urosomites, and Lang erroneously
illustrated an outer basal seta on P2. Itô provided the
first and only description of the male, and illustrated
the genital field of the female. Our redescription has
shown that the few discrepancies between his specimens and the type material are to the result of small
observational errors made by Lang (1965).
Chappuis’ (1958) specimens of A. spinicaudata from
Puget Sound cannot possibly belong to this species.
His illustrations of the fifth legs, P2 endopod, female
antennule, and caudal rami are in good agreement
with Lang’s description of A. dillonbeachia.
The currently known distribution includes records
from Puget Sound, Washington (Chappuis, 1958),
Dillon Beach and Monterey Bay, California (Lang,
1965), and Samani, Pacific coast of Hokkaido, Japan
(Itô, 1969). The egg sac contains six eggs.
GENUS ONYCHOPONTIA GEN. NOV.
The close relationship between A. orientalis (Krishnaswamy, 1957), A. intermedia Rouch, 1962, and
A. peteraxi Mielke, 1982a was first recognized by
Mielke (1982a), who regarded the sexual dimorphism
on the P3 endopod as a synapomorphy linking these
species. The transformation of this ramus is more
derived than the condition displayed in Psammoleptastacus, where the two-segmented state is retained
in the male. Species of the latter genus have a welldeveloped P1 exopod and an inner serrate seta on P2
enp-2. In A. orientalis, A. intermedia, and A. peteraxi,
the P1 exopod is more condensed and the serrate seta
on P2 enp-2 is lost. These three species are here
collectively transferred to a new genus, Onychopontia,
together with a new species from Australia found
among the type material of N. spinicaudatus. Additional autapomorphies for Onychopontia are the modification of the two outer elements on P1 exp-3 into
naked setae and the strong reduction of the outer
geniculate seta; in all other genera the outer elements
are spiniform and pinnate, and the outer geniculate
seta is well developed. Lang’s (1965) claim that P. orientalis belongs to Arenocaris is obviously a slip of the
pen, which was inadvertently adopted by Bodin (1967).
The current known distribution of the genus is highly
disjunct, with species recorded from Brazil, India,
Australia, Galápagos, Panamá, Chile, and Venezuela.
Diagnosis: Arenopontiidae. Urosomites: without conspicuous surface ornamentation. Anal somite: without
paired dorsolateral spinous processes. Anal operculum: not modified. Hyaline frills of abdominal somites
with narrow rectangular lappets. Caudal ramus:
without dorsolateral spur near medial margin. P1
exopod: three-segmented, short; exp-1 longest, with
outer spine; exp-3 with two spines and two geniculate
setae. P1 endopod: not prehensile, longer than
exopod; enp-2 with two geniculate setae. P2–P3 endopods: two-segmented; inner serrate seta of P2 enp-2
absent. P3 endopod: without outer distal element (but
see variability in Onychopontia peteraxi). P4 endopod:
with outer distal element well developed or reduced.
Armature formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.021
Endopod
0.020
0.010
0.020
[or 0.010*]
[or 0.020*†]
P3 endopod male: sexually dimorphic, onesegmented, with distal curved spine. P5: with outer
basal seta and either three or four discrete elements in
female; with outer basal seta and three elements in
male, innermost one distinctly smaller than in female,
and sometimes fused at base. P6 male: with two setae.
Type species: Onychopontia nichollsi sp. nov.
Other species: Psammoleptastacus orientalis Krishnaswamy, 1957 = Onychopontia orientalis (Krishnaswamy, 1957) comb. nov.; A. intermedia Rouch,
1962 = Onychopontia intermedia (Rouch, 1962) comb.
nov.; A. peteraxi Mielke, 1982 = Onychopontia peteraxi
(Mielke, 1982) comb. nov.
Etymology: The generic name is derived from the
Greek onyx (onux; genitive onychos), meaning nail, or
claw, and the suffix pontia (pontia, the sea), commonly used in the formation of interstitial copepod
names, and refers to the morphology of the modified
P3 endopod in the male.
ONYCHOPONTIA ORIENTALIS (KRISHNASWAMY,
1957) COMB. NOV.
Psammoleptastacus orientalis Krishnaswamy, 1957
Arenocaris orientalis (Krishnaswamy, 1957) Lang
(1965) (lapsus calami)
Arenopontia (Arenopontia) orientalis (Krishnaswamy,
1957): Wells (1967)
Original description: Krishnaswamy (1957): pp. 101–
103; text-figure 23.
Type locality: India, Tamil Nadu, Madras; sandy
beach [cf. Krishnaswamy (1957: 144–151) for sand
texture characteristics].
*Variability noted in some populations of Onychopontia
peteraxi; †there is some confusion about the correct number
of elements in Onychopontia orientalis and Onychopontia
intermedia.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
CLASSIFICATION OF ARENOPONTIIDAE
Remarks. The type material no longer exists. Krishnaswamy (1957) observed sexual dimorphism on the
P3 endopod, but the relevant illustration was mislabelled as ‘end P씹?’. His claims of five elements on P1
exp-3 and an outer spine on exp-2 are undoubtedly
wrong, and the outer basal seta on 씹 P5 was probably overlooked. Also, Krishnaswamy (1957) did not
illustrate the P3, but stated that it ‘. . . resembles the
second one, except for the increase in the number of
spines and the presence of longer setae in the
endopod’, implying that there are probably two setae
on enp-2 (as shown in his setal formula).
Onychopontia orientalis appears to prefer the drier
zones on the beach, and reaches maximum densities
around the high-tide level; known only from the type
locality.
ONYCHOPONTIA
INTERMEDIA
(ROUCH, 1962)
COMB. NOV.
Arenopontia intermedia Rouch, 1962
Arenopontia (Arenopontia) intermedia Rouch, 1962:
Bodin (1979)
Original description: Rouch (1962): pp. 277–278;
figures 158–166 (씹 unknown).
Type locality: Brazil, Pernambuco State, Recife,
Olinda; sandy beach.
Remarks: Mielke (1985, 1987) commented on the
similarity in body sculpturing between O. intermedia
and some species of Neoleptastacus (N. clasingi,
N. ornamentus, and N. reductaspina); however, this
was based on an erroneous interpretation of Rouch’s
(1962) text and figures. In the Neoleptastacus species
the abdominal somites display rectangular integumental plates both ventrally and dorsally. Rouch’s
statement that each somite boundary has ‘. . . une
série de cannelures longitudinales en forme de pallissade’ clearly refers to the deeply incised hyaline frill
(as suggested by his figs 162, 163, 166). Mielke
(1982a) remarked on the discrepancy in the number
of spines on P3 enp-2 between Rouch’s (1962) text and
his illustration (fig. 160). Onychopontia intermedia is
known only from the type locality. The type material
is no longer available for examination.
ONYCHOPONTIA
PETERAXI
(Mielke, 1982a)
COMB. NOV.
Arenopontia peteraxi Mielke, 1982a
Arenopontia (Arenopontia) peteraxi Mielke (1982a):
Bodin (1988)
449
Original description: Mielke (1982a): pp. 36–42; Abb.
22–24, 28 (distribution map).
Type locality: Ecuador, Galápagos, Jervis, north coast;
sandy beach.
Remarks: The species has also been reported from
three other islands of the Galápagos archipelago
(Isabela, James, and Marchena) (Mielke, 1982a),
three localities along the Pacific seaboard of
Panamá (Isla Taboga, Playa Nueva Gorgona, and
Playa Lagomar) (Mielke, 1982b), Arica in northern
Chile (Mielke, 1987), and Isla Margarita in Venezuela (Martínez Arbizu & Moura, 1994). Variability in
the number of elements (either one or two) on P2
enp-2 and P3 enp-2 was recorded in the Panamá
and Galápagos populations, respectively. Additional
variability can be noted in the length of the caudal
ramus spinous process, and the robustness of the
spinules on the anal operculum. Mielke (1982b) also
reported one individual with an outer spine on P1
exp-2 (atavism!); some Galápagos specimens were
found in copula. Mielke’s (1982a) claim that there
is a short outer basal seta on P1 is probably
false.
ONYCHOPONTIA
NICHOLLSI SP. NOV.
Type locality: Australia, Western Australia, ‘Back
Beach’ at Dongarra; washed from sand taken from
below 15 cm of the surface.
Material examined: Type material found among
syntype series of N. spinicaudatus (see above).
Holotype 씸: dissected on six slides (NHM
reg. no. 1994.4807). Paratypes: eight 씸씸 and one 씹
in alcohol (NHM reg. nos 1994.4808–4816; antennule
of 씹 paratype mounted on separate slide). Collected
by A.G. Nicholls on 25–26 March 1940.
Description
Female: Total body length from tip of rostrum to
posterior margin of caudal rami: 210–245 mm
(mean = 225 mm, N = 9). Body: slender and cylindrical, without clear distinction between prosome and
urosome (Fig. 21A, B). Hyaline frills of thoracic
somites weakly developed and crenulated; those of
genital double-somite and free abdominal somites
strongly developed, and consisting of narrow, rectangular digitate lappets (Fig. 21A–D). Genital doublesomite (Fig. 21A–C): as long as wide; without
chitinous ribs marking original segmentation; with
one middorsal, two lateral, and two ventral pores.
Anal somite (Fig. 21A, B, D): with two dorsal and two
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450
S. SAK ET AL.
Figure 21. Onychopontia nichollsi gen. nov. sp. nov. (씸). A, habitus, dorsal view. B, habitus, lateral view. C,
P5-bearing somite and genital double-somite, ventral view. D, posterior half of penultimate somite, anal somite, and left
caudal ramus, ventral view. E, left caudal ramus, lateral view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
CLASSIFICATION OF ARENOPONTIIDAE
lateral pores. Anal operculum: naked. Anus positioned
subterminally between caudal rami.
Caudal rami (Figs 21D, E, 22F): approximately 2.3
times longer than wide (measured in dorsal view),
tapering posteriorly; with a pore dorsally (Fig. 22F)
and two pores laterally (Fig. 21E); outer distal corner
produced into posteriorly directed recurved spinous
process, accompanied by outer spinular row at base
(Fig. 21E); dorsal surface without process or spinular
row near inner margin. Armature consisting of seven
setae: seta I, small; setae II and III, long and naked;
seta IV, short, sparsely pinnate, located between
seta V and spinous process, and fused basally to
seta V; seta V, long, sparsely plumose, and with fracture plane; seta VI, small, naked, and located at inner
distal corner; seta VII, foliaceous and triarticulate at
base.
Rostrum (Fig. 21A): small, broadly subtriangular,
tapering distally, with two delicate sensillae.
Antennule (Fig. 22A): long, six-segmented. Segment 1: with a tiny seta near the anterodistal margin.
Segment 2: longest, about 2.8 times longer than wide.
Segment 4: with long aesthetasc (22-mm long) fused at
base with seta. Distal segment: with seven naked
setae (one of which spatulate), and apical acrothek
consisting of short aesthetasc (13-mm long) and two
slender setae. Armature formula: 1-[1], 2-[6 + 1
plumose], 3-[4], 4-[(1 + ae)], 5-[1], 6-[7 + acrothek].
Antenna (Fig. 22B): coxa small, without ornamentation. Allobasis: about 2.4 times as long as maximum
width; original segmentation marked by partial
transverse surface suture; without spinular ornamentation. Exopod: one-segmented, elongate, with a
naked apical seta (slightly longer than exopod). Free
endopod with two spinular rows on anterior surface;
lateral armature consisting of two short spines; apical
armature consisting of two spines and three geniculate setae, strongest of which with spinules around
geniculation and fused basally to tiny accessory seta.
Mandibular palp: two-segmented (Fig. 22C, D);
basis elongate with one lateral seta; endopod with one
inner, one outer, and three apical setae (two fused at
base); all armature elements naked. Gnathobase:
with few sharp teeth distally and one naked seta at
dorsal corner.
Maxillule and maxilla: as in O. peteraxi.
Maxilliped (Fig. 22E): syncoxa, small and unarmed.
Basis: elongate and unarmed. Endopod: with small
accessory seta and slightly curved claw bearing subterminal spinule.
P1 (Fig. 23A): intercoxal sclerite, long and narrow.
Praecoxa: triangular and naked. Coxa: without ornamentation. Basis: with spinular row near base of
endopod; anterior surface with a small inner seta.
Exopod: three-segmented, condensed; exp-1 and exp-2
with spinules around outer margin; exp-1 longest,
451
with strong, naked outer spine; exp-2 without outer
element; exp-3 with two naked setae, one short, and
one long geniculate seta. Endopod: two-segmented,
longer than exopod, not prehensile; enp-1 2.2 times
longer than wide, with a short serrate inner seta at
about halfway along the segment length, and with
few spinules along outer margin; enp-2 longer than
wide, with a short outer and a long inner geniculate
seta.
P2–P4 (Fig. 23B–D): intercoxal sclerites naked.
Praecoxae: small and naked. Coxae: without ornamentation. Bases: generally smaller than coxae,
with a spinular row near the base of the endopod
and with few spinules along the outer margin; anterior surface with a pore in P4; outer basal seta
absent (P2), plumose (P3), or naked (P4). Exopods:
three-segmented; segments with spinular ornamentation, as illustrated; inner distal seta of exp-3
sparsely bipinnate, all other elements unipinnate;
P3–P4 exp-3 with anterior pore. Endopods: twosegmented, enp-2 very small; P2–P4 enp-1 about
3.5, 3.4, and 3.8 times longer than their respective
distal segments, with few spinules along outer
margin, as illustrated. P2 enp-2: without inner seta.
P2–P3 enp-2: with a sparsely bipinnate apical seta.
P4 enp-2: with large, apically serrate seta, fused at
base, and minute seta at outer distal corner. Spine
and seta formula as follows:
P2
P3
P4
Exopod
0.0.021
0.0.021
0.0.021
Endopod
0.010
0.010
0.020
Fifth legs (Figs 21C, 23E): closely set together, but
not touching in ventral midline. Baseoendopod and
exopod: fused, forming a squarish plate; anterior
surface with two pores; distal margin with two
pinnate setae (inner one twice the length of the outer
one) and long, naked outer seta; outer basal seta, long
and plumose.
Genital field positioned near anterior margin of
genital double-somite (Fig. 21C). Genital apertures
fused, forming median common slit; closed off by
fused P6 forming operculum, with two minute spinous
processes on either side; copulatory pore located midventrally, close to genital slit, and flanked by paired
secretory pores; seminal receptacles difficult to
discern.
Male: Total body length from tip of rostrum to posterior margin of caudal rami: 295 mm. Body ornamentation (Fig. 24A, B): essentially as in female. Sexual
dimorphism: in antennule, genital segmentation, P3
endopod, P5, and P6. Spermatophore length: approximately 18 mm.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
452
S. SAK ET AL.
Figure 22. Onychopontia nichollsi gen. nov. sp. nov. (씸). A, antennule, dorsal view. B, antenna. C, mandible. D,
mandibular palp. E, maxilliped. F, right caudal ramus, dorsal view.
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CLASSIFICATION OF ARENOPONTIIDAE
453
Figure 23. Onychopontia nichollsi gen. nov. sp. nov. A, P1 씸, anterior view. B, P2 씸, anterior view. C, P3 씸,
posterior view. D, P4 씸, anterior view. E, P5 씸, anterior view. F, P3 endopod 씹, anterior view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
454
S. SAK ET AL.
Figure 24. Onychopontia nichollsi gen. nov. sp. nov. (씹). A, urosome, ventral view. B, urosome, lateral view. C,
antennule, dorsal view. D, antennulary segments 3–7, anterior view. E, P5, anterior view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
CLASSIFICATION OF ARENOPONTIIDAE
KEY
455
TO SPECIES
Although some variability has been noted for O. peteraxi, we consider 0.020 and 0.010 as the normal formulae for
P2–P3 endopods, respectively (as found in the Chilean population; Mielke, 1987).
1. P4 enp-2: with two well-developed setae ............................................................................................... 2.
P4 enp-2: with vestigial outer seta.......................................................................................................3.
2. P2 enp-2: with inner spine over twice the length of the outer spine; P5 female with five setae ...... O. intermedia.
P2 enp-2: with both elements equally long; P5 female with four setae.........................................O. orientalis.
3. P5 male: with innermost element setiform and defined at base...................................................O. peteraxi.*
P5 male: with innermost element spiniform and fused to segment ................................. O. nichollsi sp. nov.*
*Females of these species can be separated by the relative length of P2–P4 exp-1, being much shorter in O. nichollsi.
KEY
TO
GENERA
1. P1 endopod: prehensile; innermost element of P1 exp-3 penicillate .............................................. Arenopontia.
P1 endopod: not prehensile; innermost element of P1 exp-3 geniculate ....................................................... 2.
2. P5 of both sexes: with innermost element forming distinct spinous process...............................Neoleptastacus.
P5 of both sexes: with innermost element not modified (occasionally fused at base)......................................3.
3. P1 enp-2: with one geniculate seta and one spine; P3 endopod not modified in male.........Mesopontia gen. nov.
P1 enp-2: with two geniculate setae; P3 endopod sexually dimorphic..........................................................4.
4. P2 enp-2: with inner serrate seta; P3 endopod male two-segmented...................................Psammoleptastacus.
P2 enp-2: without inner serrate seta; P3 endopod male one-segmented ....................... Onychopontia gen. nov.
Antennule (Fig. 24C, D): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 2
longest, and about 2.2 times longer than wide;
segment 4 an incomplete sclerite with two short
spiniform elements; segment 5 with three setae
plus long aesthetasc (29-mm long), fused basally to a
small seta; segment 6 with a fused spine and long
distal seta; segment 7 with two modified, fused spines
and a seta; segment 8 with a modified fused spine;
distal segment with seven naked setae (one of which
spatulate) and apical acrothek. Setal formula: 1-[1],
2-[7 + 1 plumose], 3-[4 + 1 spine], 4-[2 modified],
5-[3 + (1 + ae)], 6-[1 + 1 modified], 7-[1 + 2 modified],
8-[1 modified], 9-[7 + acrothek]. Acrothek consisting of
short aesthetasc (10-mm long) fused basally to two
slender setae.
P3 endopod (Fig. 23F): a small segment with two
spinules on anterior surface, and with a curved clawlike spine apically.
P5 (Fig. 24E): longer than wide; with same number
of armature elements as in female, but with middle
and inner elements of distal margin fused to segment;
inner one being replaced by a short bipinnate spine,
and middle one naked and comparatively shorter
than in female.
Sixth legs (Fig. 24A): slightly asymmetrical,
with smallest P6 closing off functional gonopore; each
with a short inner spine and a long, naked outer
seta.
Remarks: Onychopontia nichollsi sp. nov. is closely
related to O. peteraxi, with which it shares the vestigial outer seta on P4 enp-2. It differs in the antennule
(longer, in particular in segment 2), P1 (outer geniculate seta shorter), and male P5 (plate more rectangular; middle and inner elements fused at base). The
species is known only from the type locality where it
coexists with N. spinicaudatus.
ADDITIONAL
REMARKS
There has been some controversy over the taxonomic
significance of the foliaceous seta VII, and the
absence/presence of this character has often led to
confusion as to the true specific identity of certain
populations (e.g. Şerban, 1959; Apostolov, 1973;
Lindgren, 1976). In some species, including A. subterranea, the transparent leaf-like flanges of this seta
are very well developed, and are easy to observe
under low magnification; however, it is clear that
some authors have overlooked them in their descriptions (e.g. Masry, 1970; A. problematica). Recent highquality descriptions (Mielke, 1982a, b, 1985, 1987)
have demonstrated that this character is widespread
in the family. Our observations showed that it is
expressed (but often only weakly) in all genera, indicating that it is a potential autapomorphy for the
family.
Arenopontiids appear to have a propensity for
developing modified setation elements. Our study
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
456
S. SAK ET AL.
revealed that at least some members of every genus
exhibit one or (usually) two spatulate setae on the
posterior margin of the distal antennulary segment in
both sexes. This character, which is probably another
arenopontiid diagnostic, has thus far remained unnoticed in descriptions, except for Mielke’s (1982a)
excellent illustrations of N. trisetosus (as A. trisetosa).
ACKNOWLEDGEMENTS
Results included in this paper partly originated from
a PhD thesis by the senior author. Funding by
TÜBITAK [project no TBAG-1962(100T120)] and the
Balıkesir University Research Fund (project no BAP
2003/21) is gratefully acknowledged by SS and SK.
Thanks are also due to Dr Karin Sindemark (Swedish
Museum of Natural History, Stockholm) for arranging
the loan of Mesopontia dillonbeachia.
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