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Downloaded from https://academic.oup.com/zoolinnean/article/152/3/409/2606409 by guest on 22 March 2021Zoological 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,
Downloaded from https://academic.oup.com/zoolinnean/article/152/3/409/2606409 by guest on 22 March 2021
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 Psammoleptasta-
cus 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 409410 S. SAK ET AL.
INTRODUCTION Moura, 1994). Finally, Wells (2007) abandoned the
subgeneric classification altogether, and maintained
In one of the pioneering papers on the interstitial Pararenopontia as a valid genus.
fauna of coastal groundwater (‘Küstengrundwasser’) In this paper we have set out to: (1) redefine the
in north-western Europe, Kunz (1937) proposed the generic boundaries of Arenopontia; (2) provide argu-
genus Arenopontia for a new species Arenopontia ments for the resurrection of Psammoleptastacus as a
subterranea from Schilksee in the Kiel Bay, Germany. valid genus; (3) upgrade Neoleptastacus to its original
Pennak (1942a) erected the genus Psammoleptasta- generic rank; (4) propose two new genera for species
cus for a new species, Psammoleptastacus arenaridus, previously allocated to Arenopontia, and (5) describe
from two sandy beaches in the Woods Hole area, USA. two new species from the Turkish Black Sea coast and
Nicholls (1945) established the genus Neoleptastacus Western Australia, respectively.
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for its type and only species Neoleptastacus spini-
caudatus, from Australia. All three genera were
originally placed in the Canthocamptidae by their MATERIAL AND METHODS
respective authors. Lang (1948) transferred Arenop-
Samples in Turkey were collected using the Karaman–
ontia to the subfamily Leptopontiinae in the
Chappuis method (Delamare Deboutteville, 1953).
Cylindropsyllidae, and Noodt (1955a) suggested
Specimens were cleared in lactic acid and dissected in
Psammoleptastacus should sink as a synonym of this
lactophenol. Dissected parts were mounted on slides in
genus. Chappuis (1955) believed the separate generic
lactophenol mounting medium. Broken glass fibres
status of Neoleptastacus was not warranted, and also
were added to prevent the animal and appendages
relegated the genus to a junior subjective synonym of
from being compressed by the coverslip, and to facili-
Arenopontia. Subsequent authors accepted Chappuis’
tate rotation and manipulation, allowing observation
course of action, with the exception of Krishnaswamy
from all angles. Preparations were sealed with
(1957) who continued using Neoleptastacus as a valid
Entellan® (Merck). All drawings have been prepared
genus, and Wells (1967) who preferred a subgeneric
using a camera lucida on an Olympus BX-50 or Leica
division of Arenopontia into the nominate subgenus
DMR differential interference contrast microscope.
and Neoleptastacus, reflecting the distinct difference
Measurements were made with an ocular micrometer.
in P5 morphology. This subdivision gained wide
Total body length was measured from the anterior
acceptance (e.g. Kunz, 1971; Mielke, 1975; Lindgren,
margin of the rostrum to the posterior margin of the
1976; Itô, 1978; Bodin, 1979, 1988; Bodiou & Colom-
caudal rami. The scale bars in the illustrations are in
ines, 1986; Wells & Rao, 1987; Cottarelli, Bruno &
mm. The descriptive terminology is adopted from Huys
Venanzetti, 1994; Karanovic, 2000), but was not uni-
et al. (1996b). Abbreviations used in the text are as
versally accepted (Masry, 1970; Cottarelli, 1973, 1975;
follows: ae, aesthetasc; enp, endopod; exp, exopod;
Mielke, 1982a, b, 1985, 1987). Bodiou & Colomines
exp-1 (enp-1), exp-2 (enp-2), and exp-3 (enp-3) to
(1986) proposed a new genus Pararenopontia for two
denote the proximal, middle, and distal segment of a
Arenopontia species with reduced leg segmentation:
ramus; P1–P6, for swimming legs 1–6. The type mate-
Arenopontia breviarticulata Mielke, 1975 and Arenop-
rial was deposited in the Natural History Museum,
ontia trisetosa Mielke (1982a).
London (NHM) and Balıkesir University Zoology
Mielke (1982a) questioned the significance attri-
Museum (BUZM), and was borrowed from the
buted to the P5 morphology as a subgeneric discrimi-
National Museum of Natural History, Smithsonian
nant, as some species exhibit a transitionary
Institution, Washington D.C. (NMNH) and the
condition between the Arenopontia and Neoleptasta-
Swedish Museum of Natural History, Stockholm
cus types of P5. Martínez Arbizu & Moura (1994)
(SMNH).
argued that the subgenera of Arenopontia (Arenopon-
tia and Neoleptastacus) are not sustainable on
grounds of potential paraphyly and/or polyphyly, and
RESULTS AND DISCUSSION
that Pararenopontia should be synonymized with
Arenopontia. This amalgamation was disputed by FAMILY ARENOPONTIIDAE MARTÍNEZ ARBIZU &
Huys, Bodiou & Bodin (1996a), who resurrected MOURA, 1994
Pararenopontia, and by Huys et al. (1996b: 35), who Based on the arguments outlined below, a new
maintained the subgeneric classification (although generic classification is proposed for the family,
they did not explicitly list the subgenus Neoleptasta- resulting in the upgrade of the subgenera Arenopon-
cus). Bodin (1997) offered a compromise by adopt- tia and Neoleptastacus, in the resurrection of Psam-
ing Wells’ (1967) original subdivision and adding moleptastacus, in the rejection of Pararenopontia as a
Pararenopontia as a third subgenus (this new rank valid genus, and in the proposal of two new genera,
was erroneously attributed to Martínez Arbizu & Mesopontia and Onychopontia. Table 1 summarizes
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458Table 1. Salient features differentiating arenopontiid genera. Apomorphic character states are set in boldface
Arenopontia Mesopontia gen. nov. Neoleptastacus Psammoleptastacus Onychopontia gen. nov.
P1 exp-3 armature 2 pinnate spines + 1 2 pinnate spines + 2 1–2 pinnate spines + 2 2 pinnate spines + 2 2 naked setae + 2
geniculate seta + 1 geniculate setae geniculate setae geniculate setae geniculate setae
penicillate seta
P1 endopod prehensile not prehensile not prehensile not prehensile not prehensile
longer than exopod as long as exopod as long or longer than shorter than exopod longer than exopod
exopod
P1 enp-2 armature 1 spine + 1 1 spine + 1 1 spine + 1 2 geniculate setae 2 geniculate setae
geniculate claw geniculate seta geniculate seta*
P2–P3 endopods 씸 two-segmented two-segmented one- or two-segmented two-segmented two-segmented
P2 enp-2 inner serrate present present usually present† present absent
seta
P3 endopod 씹 not modified not modified not modified modified modified
segmentation as in 씸 as in 씸 as in 씸 two-segmented one-segmented
P4 exp-3 inner serrate absent present usually present‡ absent absent
seta
P5 with outer basal 3 or 4 discrete 4 discrete elements 1–3 discrete elements + 4 discrete elements 3 or 4 discrete
seta + elements (innermost one inner spinous innermost smaller in elements in 씸
spiniform) process 씹 3 discrete elements in
씹 (with innermost
smaller or fused)
*Except speluncae lineage (N. speluncae and N. phreaticus), which has two geniculate setae.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458
†Except N. ornamentus and N. reductaspina.
‡Except N. australis and N. pacifica; unknown in N. accraensis.
CLASSIFICATION OF ARENOPONTIIDAE
411
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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 NEOLEPTASTACUS NICHOLLS, 1945
A. subterranea Kunz, 1937* N. spinicaudatus Nicholls, 1945*
A. problematica Masry, 1970 N. australis (Chappuis, 1952) comb. nov.
A. nesaie Cottarelli, 1975 N. acanthus (Chappuis, 1954) comb. nov.
A. riedli Lindgren, 1976 N. longiremis (Chappuis, 1955) comb. nov.
N. secundus Krishnaswamy, 1957
PSAMMOLEPTASTACUS PENNAK, 1942 N. africanus (Chappuis & Rouch, 1961) comb. nov.
P. arenaridus Pennak, 1942* N. accraensis (Lang, 1965) comb. nov.
P. stygius (Noodt, 1955) comb. nov. N. indicus (Rao, 1967) comb. nov.
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P. barani sp. nov. N. ishikarianus (Itô, 1968) comb. nov.
N. angolensis (Kunz, 1971) comb. nov.
N. gussoae (Cottarelli, 1973) comb. nov.
ONYCHOPONTIA GEN. NOV. N. trisetosus (Mielke, 1982) comb. nov.
O. orientalis (Krishnaswamy, 1957) comb. nov. N. clasingi (Mielke, 1985) comb. nov.
O. intermedia (Rouch, 1962) comb. nov. N. pacificus (Mielke, 1985) comb. nov.
O. peteraxi (Mielke, 1982) comb. nov. N. spicatus (Mielke, 1985) comb. nov.
O. nichollsi sp. nov.* N. chaufriassei (Bodiou & Colomines, 1986) comb. nov.
N. ornamentus (Mielke, 1987) comb. nov.
N. reductaspina (Mielke, 1987) comb. nov.
MESOPONTIA GEN. NOV. N. phreaticus (Cottarelli et al., 1994) comb. nov.
M. dillonbeachia (Lang, 1965) 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 Arenopontia (Arenopontia), as if Mielke (1985) had
updated generic assignment of the 32 valid species in originally intended such a subgeneric assignment. It
the family is shown in Table 2. 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
GENUS ARENOPONTIA KUNZ, 1937 Wells’ (1967) subdivision bears any phylogenetic sig-
The genus Arenopontia currently contains 32 species nificance. Bodin’s (1988, 1997) error unfortunately
allocated to three subgenera (Bodin, 1997; Karanovic, perpetuated in the literature, as exemplified by Kara-
2000) (or 30 species when the subgenus Pararenop- novic’s (2000) recent key to the subgenus Neoleptasta-
ontia is attributed full generic rank; see Wells, cus, which makes no reference to Mielke’s (1985)
2007). The subgenus Arenopontia encompasses 13 species. Arenopontia clasingi, A. pacifica, and A. spi-
species and possibly one subspecies [Apostolov (1973) cata are here formally transferred to Neoleptastacus,
claimed that Arenopontia pontica Apostolov, 1969 is which will be attributed full generic rank (see below).
a subspecies of A. subterranea], whereas Karanovic The subgenus Arenopontia currently encompasses
(2000) listed 17 valid species in the subgenus Neo- the following species: A. subterranea; Arenopontia
leptastacus [note that Arenopontia sakagamii Itô, arenarida (Pennak, 1942a); Arenopontia stygia Noodt,
1978 was also listed by this author, but according to 1955b; Arenopontia orientalis (Krishnaswamy, 1957);
Wells & Rao (1987) this species is synonymous with Arenopontia intermedia Rouch, 1962; Arenopontia
Arenopontia indica Rao, 1967]. The subgeneric divi- dillonbeachia Lang, 1965; Arenopontia problematica
sion first proposed by Wells (1967) fell into disuse Masry, 1970; Arenopontia nesaie Cottarelli, 1975;
in the 1980s, when Mielke (1982a, b, 1985, 1987) Arenopontia riedli Lindgren, 1976; and Arenopontia
described several new species from Central and South peteraxi Mielke, 1982a. Bodin (1979, 1988, 1997)
America without attributing them to either subgenus. added A. subterranea Kunz? sensu Şerban & Eitel-
Unfortunately, Bodin (1988, 1997) erroneously listed Lang (1957) as a species incertae sedis, but the latter
three of those species, Arenopontia clasingi, Arenop- should be regarded as a nomen nudum. Various
ontia pacifica, and Arenopontia spicata (all described Eastern European authors (e.g. Georgescu, Marcus &
by Mielke, 1985), under the nominate subgenus Şerban, 1962) have repeatedly referred to Şerban &
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 413
Eitel-Lang’s (1957) paper as ‘Notes sur les Copépodes Type species: Arenopontia subterranea Kunz, 1937
de la Mer Noire. Izdanija, Skopje’; however consis- (by monotypy).
tently, no proper citation of volume number or pagina-
tion has been given. According to Şerban (1959), the Other species: Arenopontia problematica Masry, 1970;
authors were at that time still in the process of Arenopontia nesaie Cottarelli, 1973; Arenopontia
submitting the paper (‘Une description detaillée en riedli Lindgren, 1976.
sera publiée par Şerban & Eitel-Lang’), and it has now
been confirmed (C. Pleşa, pers. comm. to RH, 1 August Species inquirendae: Arenopontia pontica Apostolov,
1996) that the manuscript was never published. The 1969; Arenopontia nesaie Cottarelli, 1975 sensu Mit-
reference nevertheless mistakenly persisted in modern wally & Montagna (2001).
literature (e.g. Apostolov & Marinov, 1988).
Nomen nudum: Arenopontia subterranea Kunz, 1937?
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Morphological comparison revealed a core group of
closely related species within the (sub)genus Arenop- sensu Şerban & Eitel-Lang (1957).
ontia, encompassing the type species A. subterranea,
A. pontica, A. problematica, A. nesaie, and A. riedli. ARENOPONTIA SUBTERRANEA KUNZ, 1937
These five species differ from other members of the Arenopontia (Arenopontia) subterranea Kunz, 1937:
family in their unique P1 morphology, including: (1) Wells (1967)
the prehensile endopod with enp-1 being distinctly
elongate, and with enp-2 bearing an outer spine and Original description: Kunz (1937): pp. 107–110; Abb.
an inner geniculate claw, and (2) the modification of 8 (figs 38–42), 9 (figs 43–47), 10 (figs 48–51).
the inner distal element of exp-3 into a penicillate
seta. Based on these autapomorphies, we here restrict Type locality: Germany, Kieler Förde, Schilksee; ‘Küs-
the generic concept of Arenopontia to this subterranea tengrundwasser’ (intertidal coastal groundwater).
group. Our unpublished studies based on sandy beach Arenopontia subterranea has been reported from a
samples from all over Europe revealed that many new wide range of localities throughout Europe, from the
species await description (e.g. Sak, Karaytuǧ & Huys, Baltic to the Black Sea basin. With additional records
in press a), and that the five currently known species from Madeira (Delamare Deboutteville, 1960b), India
only represent the tip of the iceberg. The genus is (Rao, 1967, 1968, 1970, 1980, 1991; Rao & Ganapati,
primarily restricted to the Northern Hemisphere, the 1968, 1969; Rao & Misra, 1983), Mozambique
only exception being Wells’ (1967) doubtful outlier of (Wells, 1967), and North Carolina (Lindgren, 1976) it
A. subterranea in Mozambique. is not surprising that this species has been regarded as
potentially cosmopolitan (Wells, 1967, 1986; Lindgren,
Diagnosis: Arenopontiidae. Urosomites without con- 1976). Lindgren (1976) claimed its range might be
spicuous surface ornamentation. Anal somite without extended with more investigation of sandy beaches in
paired dorsolateral spinous processes. Anal opercu- the Pacific. Unfortunately, the great majority of these
lum not modified. Hyaline frills of abdominal somites records are not accompanied by illustrations, and
with rectangular digitate lappets. Caudal ramus with consequently their authenticity cannot be verified. The
dorsolateral spur or raised spinular row near medial discovery of a closely related species from the Isle of
margin. P1 exopod: three-segmented; exp-1 with outer Sylt (Sak, 2004) casts further doubt on the validity of
spine; exp-3 with two spines, one outer distal genicu- most north-western European, and even some
late seta, and one inner distal penicillate seta. P1 German, records. Arlt’s (1983) illustrations show that
endopod: prehensile, longer than exopod; enp-2 with his Baltic specimen does not belong to A. subterranea
one outer distal spine and one inner distal geniculate either, raising the suspicion that not all records from
claw. P2–P3 endopods: two-segmented. P3 enp-2 with east of the Skagerrak necessarily pertain to the type
outer distal element defined at base or absent. P4 species. There is no doubt that many authors have
enp-2 with well developed outer distal element. Arma- attributed their material to A. subterranea on the sole
ture formula as follows: basis that this species shows extensive intraspecific
variability. The true range of the species is as yet
Exopod Endopod
unknown, and the only reliable records appear to be
P2 0.0.021 0.110 or 0.120
restricted to German waters: (1) North Sea coast – Isle
P3 0.0.021 0.010 or 0.020
of Sylt (Noodt, 1952, 1956, 1957; Mielke, 1975, 1976),
P4 0.0.021 0.020
Amrum (Noodt, 1956, 1957), Sankt Peter-Ording
P3 endopod male: not sexually dimorphic, two- (Noodt, 1956), and Helgoland (Martínez Arbizu &
segmented. P5 with outer basal seta and three or four Moura, 1994); (2) Kieler Bucht – Schilksee (Kunz,
discrete elements: innermost one distinctly smaller in 1937; Noodt, 1956), Bottsand, Gelting Birk, Weißen-
males. P6 male with one or two seta(e). haus, and Heiligenhafen (Noodt, 1956, 1957).
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458414 S. SAK ET AL.
The type material of A. subterranea, as well as the Anal somite (Fig. 3A, B): with two dorsal and two
remainder of Kunz’ earlier collections, were destroyed lateral pores. Anal operculum: with minute pinnules
during World War II when the Institut für along free distal margin (Fig. 3A). Anus: positioned
Meereskunde was heavily bombed in 1944 (Schriever, subterminally between caudal rami. Rostrum
1984). We have been unable to obtain topotype or (Fig. 1C): small, broadly subtriangular, tapering dis-
other material that could be attributed with confi- tally, with two delicate sensillae.
dence to A. subterranea, and instead we have selected Caudal rami: approximately twice longer than wide
A. nesaie for the model description. Illustrations and (measured in dorsal view), tapering posteriorly; with
text are based on material collected from the Turkish a proximal pore dorsally (Fig. 3A), one pore near the
west coast (Marmara Sea), which represents a con- ventral proximal margin (Fig. 2A), and one pore lat-
siderable extension of the range for the species. erally near the insertion site of seta III (Fig. 3B);
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outer distal corner produced into posteriorly directed
ARENOPONTIA NESAIE COTTARELLI, 1975 recurved spinous process, accompanied by outer
spinular row at base (Fig. 3A, B); dorsal surface with
Arenopontia (Arenopontia) nesaie Cottarelli, 1975 flagellate spur-like process near inner margin, accom-
Arenopontia nesiae Cottarelli, 1975: panied by a few tiny spinules near base (Figs 1D, 3B).
Martínez Arbizu & Moura (1994: 57) (lapsus Armature consisting of seven setae: seta I, small;
calami) setae II and III, long and naked; seta IV, short,
sparsely pinnate, located between seta V and spinous
Arenopontia nessiae Cottarelli, 1975: process; seta V, long and with fracture plane; seta VI,
Martínez Arbizu & Moura (1994: 63) (lapsus small, naked, and located at inner distal corner;
calami) seta VII, foliaceous and triarticulate at base.
Arenopontia ciplaki Sak, 2004 (nomen nudum) Antennule (Fig. 3C): long, six-segmented. Seg-
ment 1 with a tiny seta near the anterodistal margin.
Original description: Cottarelli (1975): pp. 65–70; Segment 2 longest, about 3.5 times longer than wide.
figures 1–11, 13–16, 18–19, 21–23. Segment 4 with long aesthetasc (32-mm long) fused at
base with seta. Distal segment: with seven naked setae
Type locality: Italy, Sardinia, near Cagliari, Bay of (two of which are spatulate) and apical acrothek,
Quartu S. Elena, Poetto beach. consisting of short aesthetasc (20-mm long) and two
slender setae. Armature formula: 1-[1], 2-[7 + 1
Material examined: (1) one 씸 dissected on eight slides plumose], 3-[4], 4-[(1 + ae)], 5-[1], 6-[7 + acrothek].
(NHM reg. no. 2006. 1953), one 씹 mounted in toto on Antenna (Fig. 3D, E): coxa small, without ornamen-
slide (NHM reg. no. 2006. 1954), one 씹 dissected on tation. Allobasis: about 2.7 times as long as maximum
eight slides (NHM reg. no. 2006. 1955), 22 씸씸 and 22 width; original segmentation marked by partial trans-
씹씹 preserved in alcohol (NHM reg. no. 2006. 1956– verse surface suture; with two spinular rows, as illus-
1965); (2) > 50 씸씸 and > 50 씹씹 preserved in alcohol trated. Exopod one-segmented, elongate, with a naked
(deposited in BUZM). All material was collected apical seta (about 3.3 times longer than exopod). Free
from Dutlimanı Beach (Marmara Sea), 40°22.479′N, endopod with two spinular rows on anterior surface,
28°03.080′E, Balıkesir Province, Turkey; leg. S. and with finer spinules at outer distal corner; lateral
Karaytuğ and S. Sak, 18 September 2001. armature consisting of two short spines; apical arma-
ture consisting of two spines and three geniculate
Redescription setae, the longest of which with spinules around
Female: Total body length from tip of rostrum to geniculation, and fused basally to tiny accessory
posterior margin of caudal rami: 341–396 mm seta.
(mean = 366 mm, n = 25). Maximum width: 38 mm Mandible: with two-segmented palp (Fig. 2D); basis
(mean of 20 individuals = 41 mm), measured at poste- elongate with one lateral seta; endopod with one
rior margin of cephalothorax. Body: slender and cylin- inner, one outer, and three apical setae; all armature
drical, without clear distinction between prosome and elements naked. Gnathobase: with coarse teeth dis-
urosome (Fig. 1A, B). Hyaline frills of thoracic somites tally, and with one naked seta at dorsal corner.
weakly developed and crenulated; those of genital Maxillule (Fig. 1E): with praecoxal arthrite bearing
double-somite and free abdominal somites strongly two setae and five spines around distal margin. Coxal
developed, and consisting of rectangular digitate endite: with two long naked setae. Basis with rami
lappets (Figs 1A, B, 2A, B). Genital double-somite entirely incorporated; palp represented by nine naked
(Figs 1A, B, 2A): slightly longer than wide; without setae.
chitinous ribs marking original segmentation; with Maxilla (Fig. 2E): syncoxa with two cylindrical
two mid-dorsal, two lateral, and two ventral pores. endites; proximal endite with three setae; distal
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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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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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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.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458418 S. SAK ET AL.
endite with two setae. Allobasis: drawn out into long spinous processes on either side; copulatory pore
claw; with one accessory setae. Endopod one- located midventrally, close to genital slit; seminal
segmented , and with three setae. All elements naked. receptacles difficult to discern.
Maxilliped (Fig. 2F): syncoxa small and unarmed.
Basis: elongate and unarmed. Endopod with small Male: Total body length from tip of rostrum to
accessory seta, and with slightly curved claw bearing posterior margin of caudal rami: 320–374 mm
subterminal spinule. (mean = 346 mm; N = 25). Maximum width: 40 mm
P1 (Fig. 4A): intercoxal sclerite long and rectan- (mean = 38, N = 20), measured at cephalothorax.
gular. Praecoxa: triangular and naked. Coxa: Body ornamentation (Fig. 5A): essentially as in
without ornamentation. Basis: with spinular row female. Sexual dimorphism: in antennule, genital seg-
near bases of endopod and exopod; anterior surface mentation, and P5 and P6. Spermatophore length:
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with a proximal pore and a small inner seta. approximately 35 mm.
Exopod: three-segmented; exp-1 and exp-2 with Antennule (Fig. 5B, C): nine-segmented, haplocer;
spinules around outer margin; exp-1 longest, with geniculation between segments 7 and 8. Segment 2
long unipinnate outer spine; exp-2 without outer longest, and about 2.7 times longer than wide;
element; exp-3 with short unipinnate outer spine, a segment 4 an incomplete sclerite with one modified
long curved unipinnate spine, and one geniculate (fused at base) and one tiny element; segment 5 with
seta distally, and one inner, apically penicillate seta three setae plus long aesthetasc (42-mm long) fused
subdistally. Endopod: two-segmented, prehensile; basally to a small slender seta; segment 6 with a
enp-1 9.3 times longer than wide, and about twice spinulose spine and long distal seta; segment 7 with
longer than exopod; with a serrate inner seta in three modified spines and a seta; segment 8 with a
proximal third, and a subdistal spinule along outer modified spine; distal segment with seven naked
margin; enp-2 slightly longer than wide, with a setae (two of which spatulate) and apical acrothek.
short unipinnate spine, a geniculate claw, and a Setal formula: 1-[1], 2-[7 + 1 plumose], 3-[4 + 2
small inner spinule. spines], 4-[1 + 1 modified], 5-[3 + (1 + ae)], 6-[1 + 1
P2–P4 (Fig. 4B–D): intercoxal sclerites naked, modified], 7-[1 + 3 modified], 8-[1 modified],
wider in P2, but more deeply concave in P3–P4. 9-[7 + acrothek]. Acrothek consisting of short aes-
Praecoxae: small and naked. Coxae: squarish and thetasc (16-mm long) fused basally to two slender
without ornamentation. Bases: smaller than coxae, setae.
with a spinular row near base of endopod (P3–P4); P5 (Fig. 2B): with armature as in female, but with
anterior surface with a pore; outer basal seta absent middle and inner elements comparatively shorter.
(P2), plumose (P3), or naked (P4). Exopods: three- Sixth legs (Fig. 2B): asymmetrical, with smallest P6
segmented; segments with spinular ornamentation, closing off functional gonopore; each with a long
as illustrated; inner distal seta of exp-3 sparsely plumose seta.
bipinnate, all other elements unipinnate; P3–P4 exp-3
with anterior pore. Endopods: two-segmented; P2–P4 Remarks: Arenopontia nesaie was originally des-
enp-1 about 1.5, 2.2, and 3.0 times longer than their cribed from Poetto Beach in the Bay of Quartu S.
respective distal segments, with few spinules, as illus- Elena near Cagliari, Sardinia (Italy). Our material
trated. P2: enp-1 with a long, apically serrate, back- differs from Cottarelli’s (1975) description in some
wardly directed seta near proximal inner corner. aspects, but these are most likely attributable to
P2–P3: enp-2 with a long, bipinnate, apical seta. P4: deficiencies in the original figures. In the type mate-
enp-2 with apically serrate seta, fused at base, and rial the marginal spines on the P5 of both sexes
long unipinnate seta at outer distal corner. Armature appear shorter; however, the flagellate distal parts of
formula as follows: P2, exopod, 0.0.021, endopod, these elements are usually difficult to discern, and it
0.110; P3, exopod, 0.0.021, endopod, 0.010; P4, seems conceivable that they were not illustrated cor-
exopod, 0.0.021, endopod, 0.020. rectly in the original description. Similarly, Cottarelli
Fifth legs (Fig. 2A) closely set together, but not (1975) did not illustrate the spinules at the base of
touching in ventral midline. Baseoendopod and the spur and around the terminal process of the
exopod: fused, forming a rectangular plate; distal caudal ramus, but such morphological minutiae were
margin with three pinnate setae, middle one mark- generally overlooked prior to the advent of differential
edly shorter than the others, but not vestigial; outer interference contrast microscopy. The female anten-
basal seta, long and plumose. nule has fewer setae on the proximal segments than
Genital field: positioned near anterior margin of in the Turkish material, but this can be attributed to
genital double-somite (Fig. 2A). Genital apertures the fact that Cottarelli viewed the appendage in
(Fig. 2C): fused forming median common slit; closed dorsal aspect and hence overlooked various setae
off by fused P6 forming operculum with three minute arising from the ventral surface. The three-
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 419
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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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Figure 5. Arenopontia nesaie Cottarelli, 1975 (씹). A, habitus, dorsal view. B, antennule, ventral view. C, antennule,
anterior view.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 421
segmented mandibular palp, which was considered ARENOPONTIA PONTICA APOSTOLOV, 1969
diagnostic for A. nesaie, was not observed in our Original description: Apostolov (1969): pp. 125–127;
material, and requires confirmation. The extra Abb. 36–45.
segment boundary indicates a two-segmented
endopod, which has thus far not been reported for any
other oligoarthran harpacticoid (but see Mitwally & Type locality: Bulgaria, south of Lozenetz, Düni
Montagna, 2001; cf. below). It is also noteworthy that Beach; 5 m from low-tide mark.
Cottarelli (1975) had accidentally rotated exp-3 in his
drawing of the P2. His drawing of the female genital Remarks: Apostolov’s (1969) description of A. pontica,
field superimposes external and internal structures; from the Bulgarian Black Sea coast, is a taxonomic
for a more accurate interpretation see Fig. 3C and nightmare because of several internal inconsistencies
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Martínez Arbizu & Moura (1994: fig. 2c, as A. nes- between the text and illustrations. Apostolov (1969:
siae). The Sardinian specimens are somewhat smaller 111) claimed to have found two females (although on
[334 mm (씸), 285–300 mm (씹)] than the Marmara p. 125 he stated that three females were recorded),
population [341–396 mm (씸), 320–374 mm (씹)], but it but for some inexplicable reason provided a brief
is questionable whether this size discrepancy has diagnosis of the male. He referred to Figure 46 in his
any significance beyond the range of intraspecific description of the male P5, but this figure is not
variability. printed. His illustrations of the female show several
Arenopontia nesaie appears to be widely distributed extraordinary features not found in any other
in the Mediterranean, with confirmed intertidal member of the Arenopontiidae: (1) the antennary
records from El Saler (Valencia, Spain) by Martínez exopod is bisetose – in all species of Arenopontia this
Arbizu & Moura (1994, as A. nesiae), Sardinia ramus displays only one apical seta; (2) P1 exp-2
(Cottarelli, 1975), the mouth of the Trigno River on the bears an outer spine – the absence of this spine is a
Adriatic coast (Molise, Italy) by Bruno, Cottarelli & high-level diagnostic, being a synapomorphy linking
Berrera (1998), and Dutlimanı beach (Sea of the Parastenocarididae, Leptopontiidae, and Arenop-
Marmara, Turkey) by Sak (2004, as A. ciplaki; present ontiidae (Martínez Arbizu & Moura, 1994); (3) P1
account). Mitwally & Montagna (2001) provided a enp-1 lacks an inner seta – this seta is present in all
redescription of A. nesaie based on specimens collected species, except for the inadequately described A. prob-
from three beaches (Bir Masoud, El Mamoura, and El lematica and Arenopontia accraensis Lang, 1965 – we
Shatby) near Alexandria, Egypt, but the many defi- have been able to confirm its presence in the types of
ciencies in their illustrations make it difficult to vali- A. problematica; (4) P2–P3 exp-3 with four elements,
date their identification. According to Wells (2007), i.e. with two outer spines and two terminal setae – all
Mitwally & Montagna (2001) make statements about Arenopontiidae have only one outer spine and share a
the setation of P1–P4 that, if true, mean that their [021] setal formula on the distal exopod segment –
material cannot belong to Arenopontia. It is obvious note that Apostolov (1969) contradicts himself in the
that their atypical setal formula results from a failure setal formula table on p. 125 (three elements), his
to distinguish between ornamentation elements (such Figure 42 (four elements), and the comparative table
as long spinules) and genuine setae/spines. Their on p. 127 (four elements); (5) P2–P3 exp-2 bears a
reports of an outer seta on P1 enp-1 and P3–P4 enp-1, long inner seta – the latter seta is absent in all other
as well as their claim of four elements on P2 exp-3, are arenopontiids, except for Arenopontia angolensis
false and do not reflect deficiencies in Cottarelli’s Kunz, 1971, which according to Kunz’ (1971) setal
(1975) original description, as claimed by the authors. formula possesses a seta on P2 exp-2. However, as
The elements on the female P5 are distinctly longer Kunz neither illustrated the P2 nor mentioned this
than in A. nesaie (but are similar to our specimens), character in the text or the table comparing Arenop-
and the caudal ramus appears shorter. The variability ontia africana f. africana and A. africana f. angolensis
illustrated for the male P5 suggests that Mitwally & (he does state that the P2 is as in the nominate
Montagna (2001) had an amalgam of Arenopontia subspecies, apart from the ornamentation of the inner
species in their samples. No information was given on seta on enp-2), we strongly suspect that his report is
the number of setae on the male P6. The distal based on a slip of the pen in his table, rather than on
segment of the P4 exopod appears rotated in their an observational error.
Fig. 11G. Finally, the mandibular palp is erroneously Apostolov (1969) recognized a close relationship
illustrated as three-segmented (see above). Pending with A. subterranea, A. indica and A. sp. sensu Griga
re-examination of more material of the Egyptian (1964) [the latter was later identified as conspecific
populations, A. nesaie Cottarelli (1975) sensu Mit- with Stenocaropsis valkanovi (Marinov, 1974), family
wally & Montagna (2001) is considered species Cylindropsyllidae]. In our opinion it is impossible to
inquirenda in Arenopontia. make any positive statement on the identity and
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458422 S. SAK ET AL.
Table 3. Diagnostic characters of Arenopontia species
P1 enp-1:exp P2 enp P3 enp P5씸씹 P6씹 CR An Op
A. riedli 1.4 0.120 0.020 5 2 Spur Smooth
A. nesaie 2.0 0.110 0.010 4 1 Spur Pinnate
A. subterranea 1.5 0.110 0.010 4 2?* Spinules Smooth
A. problematica 1.5† 0.110‡ 0.010‡ 4 2?* Spinules 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.
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‡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 GENUS PSAMMOLEPTASTACUS PENNAK, 1942A
this species can be assigned to the genus Arenopontia Pennak (1942a) proposed this genus for a new species,
as diagnosed herein. Pending redescription, A. pon- P. arenaridus, collected from two sandy beaches near
tica is considered here as species inquirenda. This Woods Hole, and placed it without any further
course of action is in contrast to Marinov’s (1971) comment in the Canthocamptidae. He remarked on the
suggestion to relegate A. pontica to a junior subjective superficial resemblance with other interstitial genera
synonym of A. subterranea. Marinov rightly pointed (Leptastacus, Paraleptastacus, and Arenopontia), but
out some of the weaknesses in Apostolov’s (1969) considered the differences in the antenna, maxillipeds,
description, but it remains a mystery how he recon- P5, and caudal rami sufficient for generic distinction.
ciled the many differences between the latter and his Pennak’s (1942a) paper remained largely unnoticed
own illustrations of A. subterranea from the Bulgar- until Noodt (1955b) synonymized Psammoleptastacus
ian coast. with Arenopontia, a course of action that was endorsed
Inspired by the variability reported for French by Lang (1965) but was overlooked by Krishnaswamy
mediterranean (Chappuis, 1954a) and Romanian (1957), who added a second species, Psammoleptasta-
populations (Şerban, 1959) of A. subterranea, but cus orientalis Krishnaswamy, 1957 from the Madras
apparently unaware of Marinov’s (1971) paper, coast. As noted by Wells (1967), Lang’s (1965) state-
Apostolov (1973) claimed that A. pontica may well ment that P. orientalis belongs to Arenocaris (Leptast-
be a synonym of the latter. He further proposed that acidae) is obviously a slip of the pen.
the Black Sea specimens represent a new subspecies Noodt (1955b) considered A. stygia to be most closely
of A. subterranea, but refrained from formally related to A. arenarida, recognizing some subtle differ-
naming it. Apostolov stated that considerable vari- ences in the caudal rami and P2–P4, whereas Lindgren
ability was found in the caudal rami, the P1 exopod, (1976) suggested A. stygia is potentially ‘. . . an
and the P5, but it is conceivable that this is at least intraspecific variation of A. arenarida’. The genus
partly attributable to his failure to discriminate Psammoleptastacus is reinstated herein for the latter
between two or more coexisting species. His draw- two species and a new species, P. barani sp. nov., from
ings of the female P5 clearly refer to two different the Turkish Black Sea coast, which had previously
species: his Figure 5 shows a fifth leg of the subter- been misidentified as A. stygia by Marinov (1971).
ranea type, whereas Figure 6 was almost certainly Arenopontia subterranea Kunz, 1937 sensu Rao &
based on the species previously identified by Ganapati (1969) is regarded as a species inquirenda in
Marinov (1971) as A. stygia (and described below as Psammoleptastacus, and P. orientalis is transferred to
Psammoleptastacus barani sp. nov.). In accordance Onychopontia gen. nov. Psammoleptastacus differs
with Şerban’s (1959) observations, Apostolov (1973) from Arenopontia and other arenopontiid genera in the
maintained that his material did not display the small size of the P1 endopod, which is shorter than the
foliaceous seta VII, or the penicillate seta on P1 exopod. It is most closely related to Onychopontia, with
exp-3. which it shares the sexual dimorphism on the P3
endopod (apomorphic) and the presence of two genicu-
late setae on P1 enp-2.
Key to species: A simple dichotomous identification
key is difficult to construct; however, species can be
reliably identified by considering the salient diagnos- Diagnosis: Arenopontiidae. Urosomites: without con-
tic characters summarized in Table 3. spicuous surface ornamentation. Anal somite: without
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 423
paired dorsolateral spinous processes. Anal opercu- Material examined: NMNH: one 씹 syntype mounted
lum: not modified. Hyaline frills of abdominal somites in toto on slide, and partly remounted by one of us
with rectangular digitate lappets. Caudal ramus: (RH); erroneously labelled ‘Paraleptastacus arenari-
with dorsolateral spur near medial margin. P1 dus n.g. n. sp.’; Cat. no. 81982; leg. R.W. Pennak, Sep-
exopod: three-segmented; exp-1 with outer spine; tember 1939.
exp-3 with two spines and two geniculate setae. P1
endopod: not prehensile, shorter than exopod; enp-2 Partial redescription
with two geniculate setae. P2–P3 endopods: two- Male: Total body length from tip of rostrum to pos-
segmented. P3 endopod: with outer distal element terior margin of caudal rami: 325 mm. Body: slender
defined at base. P4 endopod: with well-developed and cylindrical, without clear distinction between
outer distal element. Armature formula as follows: prosome and urosome. Hyaline frills of thoracic
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somites weakly developed and crenulated (Fig. 6A,
Exopod Endopod B); those of abdominal somites strongly developed
P2 0.0.021 0.120 and consisting of rectangular digitate lappets
P3 0.0.021 0.020 (Fig. 6A).
P4 0.0.021 0.020 Caudal rami (Fig. 6A, C): approximately 2.8 times
P3 endopod male: sexually dimorphic, two- longer than basal width, tapering posteriorly; with
segmented; enp-1 unarmed; enp-2 minute, with one pore dorsally, one pore near ventral proximal
strong spinule on outer margin, curved spine distally margin, and two pores laterally near outer spinules;
(sometimes fused at base), and fine seta on inner outer distal corner produced into posteriorly directed
margin. P5: with outer basal seta and four discrete recurved spinous process, accompanied by ventral
elements; innermost one distinctly smaller in male. spinular row at base; dorsomedial surface with pos-
P6 male: with two setae. teriorly directed spinous process. Armature consisting
of seven setae: seta I, small; setae II and III, long and
Type species: Psammoleptastacus arenaridus Pennak, naked; seta IV, short, sparsely pinnate, located
1942a (by monotypy). 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 folia-
Other species: Arenopontia stygia Noodt, 1955b = P.
ceous and triarticulate at base.
stygius (Noodt, 1955b) comb. nov.; P. barani sp. nov.
Rostrum (Fig. 6D): small, broadly subtriangular,
tapering distally, with two delicate sensillae and sub-
Species inquirenda: Arenopontia subterranea Kunz,
apical pore.
1937 sensu Rao & Ganapati (1969)
Antennule (Fig. 7A): nine-segmented, haplocer;
geniculation between segments 7 and 8. Segment 2
PSAMMOLEPTASTACUS ARENARIDUS PENNAK, 1942A longest; segment 4 an incomplete sclerite with one
Psammoleptastacus arenardius Pennak, 1942a: Coull modified (fused at base) and one tiny element; seg-
(1977) (lapsus calami) ment 5 with long aesthetasc fused basally to seta;
segments 6–8 with one seta and one basally fused
Arenopontia arenarida (Pennak, 1942a) Noodt spiniform element. Setal formula: 1-[1], 2-[7 + 1
(1955a) plumose], 3-[4 + 1 pinnate spine], 4-[1 + 1 modified],
Arenopontia (Arenopontia) arenarida (Pennak, 5-[2 + (1 + ae)], 6-[1 + 1 modified], 7-[1 + 1 modified],
1942a): Wells (1967) 8-[1 + 1 modified], 9-[7 + acrothek]. Acrothek consist-
Arenopontia arenardia (Pennak, 1942a): Coull (1971, ing of short aesthetasc fused basally to two slender
1977) (lapsus calami) setae.
P1 (Fig. 7B): coxa without ornamentation. Basis:
Arenopontia stygia Noodt (1955b) sensu Coull (1971) with spinular row near bases of endopod and exopod;
and Lindgren (1976) anterior surface with inner naked seta. Exopod:
three-segmented; about 1.3 times the length of
Original description: Pennak (1942a): pp. 275–278; endopod; all segments with spinules along outer
plate I, figures 1–11. margin; exp-1 longest, with long unipinnate outer
spine; exp-2 without outer element; exp-3 with two
Type locality: USA, Massachusetts, Woods Hole. unipinnate spines and two geniculate setae of differ-
Pennak (1942a) collected material from both Nobska ent lengths. Endopod: two-segmented, not prehensile;
and north Cape Cod beaches, but did not specify the enp-1 slightly longer than exp-1, with a serrate seta
type locality; sand washings in vicinity of high tide at about halfway along the length of the inner
mark. margin, and with two subdistal spinules along outer
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458424 S. SAK ET AL.
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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.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 425
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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.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458426 S. SAK ET AL.
margin; enp-2 about half the size of enp-1, with two least North Inlet, South Carolina (Coull & Dudley,
geniculate setae and a few spinules. 1985) in the south. Pennak (1942b) provides data on
P2 endopod (Fig. 7C): two-segmented; enp-1 with the horizontal distribution and relative abundance.
few spinules along outer margin; enp-2 short, with a
long, apically serrate, backwardly directed seta near
PSAMMOLEPTASTACUS STYGIUS (NOODT, 1955B)
proximal inner corner, and a long bipinnate inner seta
COMB. NOV.
and a short bare outer spine around distal margin.
P3 endopod (Fig. 7D): two-segmented; enp-1 with Arenopontia stygia Noodt (1955b)
few strong spinules along outer margin; enp-2 Arenopontia (Arenopontia) stygia Noodt (1955b):
minute, with strong spinule at outer distal corner, Wells (1967)
short thin seta arising from inner distal corner
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(homologous with long inner distal seta of female), Original description: Noodt (1955b): pp. 101–102;
and naked curved apical spine, fused at base Tafel 35 (figs 75–82) (씸 only).
(homologous with outer distal spine of female).
P4 endopod (Fig. 7E): two-segmented; enp-1 with Type locality: France, Landes, Mimizan-Plage;
five strong spinules along outer margin; distal margin medium coarse sand.
of enp-2 with long, basally fused, serrate seta and
long, unipinnate outer seta. Remarks: The type material of A. stygia (a single
P2–P4: spine and seta formula as for the genus. female) is no longer extant. Noodt’s (1955b) dorsal
P5 (Fig. 6B): forming subrectangular plate; outer view of the caudal ramus shows three setae inserting
basal seta sparsely plumose. Free distal margin: with at about the same level; the short inner one (adjacent
three short bipinnate spines and one long bipinnate to seta VII) is not a setal element but the dorsolateral
outer seta; inner spine longer than the other two. spur. In addition to the type locality (Delamare
Sixth legs (Fig. 6B): asymmetrical, with smallest P6 Deboutteville, Gerlach & Siewing, 1955; Noodt,
closing off functional gonopore; each with a short 1955b, c; Delamare Deboutteville, 1960a), P. stygius
inner and a long plumose outer seta. has been recorded from the Bassin d’Arcachon,
Gironde (Renaud-Debyser, 1963a, b) and the Portu-
Remarks: Pennak’s (1942a) illustrations of the male guese coast (Francelos, south of Porto) (Galhano,
are restricted to the antennule, and no reference was 1970). Marinov’s (1971) record from Bulgaria is
made to the sexual dimorphism on the P3 endopod. attributable to P. barani sp. nov. (see below).
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 PSAMMOLEPTASTACUS BARANI SP. NOV.
a source of confusion for subsequent identifications. Arenopontia stygia Noodt, 1955b sensu Marinov
Re-examination of a male paratype proved: (1) caudal (1971)
ramus seta V to be normally developed, and the ter-
minal spinous process to be much longer and more Type locality: Turkey, Black Sea coast, Istanbul,
sharply pointed; and (2) P1 endopod to be markedly Sahilköy (east of Bosporus); sandy beach.
shorter than the exopod. Both aspects contradict Pen-
nak’s (1942a) description, but agree with Lindgren’s Material examined: Holotype 씸 (dissected on eight
(1976) observations based on North Carolina speci- slides) (BUZM). Paratypes are one 씸 and one 씹 in
mens. Lindgren also found that the proximal third of alcohol (NHM reg. nos. 2006. 1966–1967), and two
seta V was modified in approximately 60% of the 씹씹 dissected on two and seven slides, respectively
population; individuals with unmodified setae were (NHM reg. nos. 2006. 1968–1969); all collected at type
provisionally identified as A. stygia. Given this locality; leg. S. Karaytuğ and S. Sak, 01 May 2001.
intraspecific variability, in conjunction with both
‘populations’ having the same distribution within the Description
beach, we attribute all North Carolina records of Female: Total body length from tip of rostrum to
A. stygia to P. arenaridus. These include Lindgren’s posterior margin of caudal rami: 330–380 mm
(1976) intertidal record from west of the Iron Steamer (mean = 361 mm; N = 7). Maximum width measured
Pier near Morehead City, and, although provisionally, at P5-bearing somite. Body: slender and cylindrical,
Coull’s (1971) subtidal record north of Cape Hatteras without clear distinction between prosome and
(at a depth of 100 m!). Psammoleptastacus arenaridus urosome (Fig. 8A, B). Hyaline frills of thoracic somites
appears to be restricted to the north-eastern Atlantic weakly developed and crenulated; those of genital
seaboard of the USA, from the Woods Hole area in the double-somite and free abdominal somites strongly
north (Pennak, 1942a, b, 1952; Lindgren, 1976) to at developed, and consisting of rectangular digitate
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 409–458CLASSIFICATION OF ARENOPONTIIDAE 427
Downloaded from https://academic.oup.com/zoolinnean/article/152/3/409/2606409 by guest on 22 March 2021
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.
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