Zootaxa 1977: 21–38 (2009) ISSN 1175-5326 (print edition)

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Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition)

Andeepia ingridae a new genus and species of Pardaliscidae (Crustacea,

Amphipoda) from the Antarctic deep-sea and short redescription of
Nicippe unidentata K.H. Barnard, 1932

TAPATI BISWAS1, CHARLES OLIVER COLEMAN1,3 & ED A. HENDRYCKS2

1
Museum für Naturkunde, Institut für Systematische Zoologie, Invalidenstraße 43,D-10115 Berlin, F.R.G.
E-mail: tapati@zedat.fu-berlin.de
2
Research Services, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario, Canada K1P 6P4.
E-mail: ehendrycks@mus-nature.ca
3
Corresponding author. E-mail: oliver.coleman@museum.hu-berlin.de

Abstract

The new genus and species Andeepia ingridae is described. The main features of this taxon are: urosomite 2 with mid-
dorsal tooth; massive propodi of gnathopods 1–2; carpus of gnathopod 1 short, weakly lobate and half the length of that
of gnathopod 2; dactylus of gnathopod 2 with distinct dentition on the inner margin; dactylus of gnathopod 1 smooth,
lacking dentition, but in the cuticle with faint trace of embedded teeth and reduced setation on pereopods and uropods.
The related southern ocean pardaliscid, Nicippe unidentata K.H. Barnard, 1932 is also re-described and illustrated for the
first time from the type material.

Key words: Taxonomy, Amphipoda, Andeepia gen. n., Andeepia ingridae sp. n., Pardaliscidae, Nicippe unidentata, new
species, deep-sea, Antarctic

Introduction

The ANDEEP (Antarctic benthic DEEP-sea biodiversity) expeditions had the aim to describe the deep-sea
fauna from the eastern Atlantic sector close to the Antarctic continent, in order to examine the deep sea diver-
sity of this area. During the ANDEEP III Expedition in 2005, a remarkable collection of crustaceans was
obtained. Amongst these samples, specimens of the amphipod family Pardaliscidae Boeck, 1871 were espe-
cially numerous. The work on this family started with a species with a striking gnathopod morphology. Ini-
tially, the first observations made on this material suggested that it could be a new species of Nicippe
Bruzelius, 1859. However, upon a more detailed examination it became clear that there were numerous differ-
ences to Nicippe and the other related genera Spelaeonicippe Stock & Vermeulen, 1982 and Antronicippe
Stock & Iliffe, 1990. Therefore, it was necessary to erect a new genus Andeepia to distinguish it.

Material and methods

Material was fixed in 96% ethanol. Pencil drawings were prepared using camera lucida on Leica Wild M8 and
Nikon SMZ-U dissecting microscopes and on Leica DMLB and Olympus BH-2 compound microscopes. Line
drawings were made using the technique described in Coleman (2003). Circles (solid and dotted) on append-
ages indicate insertion points of omitted setae.

Accepted by G. Karaman: 18 Nov. 2008; published: 14 Jan. 2009 21

 Body length measurements were made along the dorsal outline of the animals, beginning at the tip of the
rostrum to the end of the urosome.
Type material of Nicippe unidentata K.H. Barnard, 1932 was borrowed from the collections of The Natu-
ral History Museum (NHM), London. As the type material of this species is in poor condition, a specimen col-
lected during the same expedition in close proximity to the type station has been selected as a paralectotype
and illustrated (refer to Material Examined section).
Type material of the new taxon is deposited in the collections of the Zoological Institute and Zoological
Museum, University of Hamburg (ZIM) and the Canadian Museum of Nature (CMN), Ottawa, Canada. Scale
bars are shown on the figures.

Systematics

Pardaliscidae

Andeepia gen. n.

Diagnosis. Mouthparts not cone-shaped, buccal mass shallow. Coxal plates short, shallow, weakly to not
overlapping. Urosomite 1 tooth lacking; urosomite 2 with a conspicuous tooth projecting over posterior mar-
gin. Gnathopod 1 with short carpus and short ventral lobe, propodus massive with straight palm margin and
strongly curved, long dactylus, inner margin of dactylus smooth, but inside the cuticle faint embedded teeth
can just be discerned (only visible under high magnification).
Gnathopod 2 with carpus twice as long as that of gnathopod 1, propodus less robust, shorter and tapering
distally, palm convex. Dactylus of gnathopod 2 shorter than that of gnathopod 1 and with distinct toothed den-
tition on the inner margin. Pereopods 3–7 weakly spinose/setose (especially on pereopods 5–7). Uropods 1–3
rami lanceolate, rami of each uropod subequal in length, spines short and sparse. Uropod 1 peduncle lacking
distolateral tooth, but with short tooth distomedially. Uropod 2 peduncle with short distomedial tooth. Uropod
3, outer ramus 2-articulate, distal article minute. Telson cleft about 2/3 total length apices rounded, not
notched.
Type species: Andeepia ingridae sp. n. by monotypy.
Etymology: The genus name is derived from the Andeep Expedition with the research vessel Polarstern
during which the material was collected.
Remarks. The new genus appears to be most similar to Nicippe, and the related genera Spelaeonicippe
Stock & Vermeulen, 1982 and Antronicippe Stock & Iliffe, 1990 but due to some remarkable differences (see
table 1), it required a new genus to accommodate it.
From Nicippe and all other pardaliscids known, Andeepia differs most obviously in the much more robust
and different form of the gnathopod propodi, especially obvious in gnathopod 1. In comparison to Nicippe, the
enlarged propodus, short carpus and straight palmar margin of gnathopod 1 are strikingly different. Numerous
other characters of difference from Nicippe are shown in table 1.
Andeepia differs from Spelaeonicippe and Antronicippe in many characters. Spelaeonicippe provo Stock
& Vermeulen, 1982 has an elongate antenna 2 (vs shorter in the new taxon); pereopods 6–7 basis with poster-
oventral lobe (vs non-lobate, straight); the outer ramus of uropods 1–2 is much shorter than the inner (vs sub-
equal) and the telson has strong marginal spines (vs lacking spines).
Antronicippe serrata Stock & Iliffe, 1990 has very slender gnathopods (vs robust) and their dactyli have
fine serrated teeth on the inner margin; pereopod 7 basis is strongly expanded posteroventrally and lobate (vs
rectangular and non-lobate); the outer ramus of uropods 1–2 is much shorter than the inner (vs subequal) and
the telson is very shallowly (10%) notched (vs cleft 68%).

22 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

TABLE 1. Differences between the genera Nicippe and Andeepia gen. n.

Nicippe Andeepia gen. n.

Right mandible incisor
Maxilla 2 plates
Maxilliped palp
Antenna 1 peduncular article 3
Pereopod coxae
Gnathopod carpus lobes
Carpus length of gnathopod 1 com-
pared to gnathopod 2
Propodus of gnathopods 1 and 2
Gnathopod 1 palm margin
Merus/carpus ratio of pereopods 3–4
Merus to propodus of pereopods 5–7
Middorsal posterior margin of uro-
somite 1
Middorsal posterior margin of uro-
somite 2
Setation/spination of uropods 1–3
Uropod 1
Uropods 1–2 apical third of rami
Uropod 3
Telson
-strongly toothed- 4 large teeth, very -very weakly toothed, not much differ-
asymmetric from left ent from left
- narrow, subequal in length - much wider, outer plate somewhat
longer
- inner plates long and tapering - inner plates wide and short
- palp long and relatively wide - palp is much shorter
- toothed inner margin of dactylus - smooth dactylus
- short, length 0.4 x or less than article -longer, length 0.57–0.6 x article 2
2
- coxa 1 broadest and rounded, coxae - coxa 1 short and narrow, coxae 2–4
2–4 progressively smaller wider and subequal in length
- strongly produced - weakly produced
- subequal - half the length
- rather narrow - robust, massive
- strongly convex - straight
- nearly the same length (0.88–0.98 x) - much shorter than carpus (0.55–0.58
x)
- strongly setose, especially the ante- - with few and short setae
rior margins
- uni- or bidentate - not dentate
- not dentate - unidentate
- strong - weak
- peduncle with strong distolateral - peduncle lacking distolateral tooth;
tooth; peduncle subequal in length to peduncle much longer, 1.4–1.5 x rami
rami length
- setose - not setose (and pointed)
- long and strongly surpassing uropods - short and equal to uropods 1–2 in lat-
1–2 in lateral view eral view
- narrowly lanceolate - lanceolate
- outer ramus clearly longer than inner, - rami subequal
1-articulate - outer ramus 2-articulate
- inner margins of rami with dense and - inner margins with sparse setation
long setation
- long (length 2–2.5 x width) - short and wide (length 1.6 x width)
- lobes with terminal notch - lobes without terminal notch,
- deeply cleft rounded
- shallow cleft

The genera Spelaeonicippe and Antronicippe both occur in anchialine caves. Spelaeonicippe contains two
species, one found in Lanzarote (Canary Islands), the other in the Turks and Caicos Island (West Indies).
Antronicippe was collected in a cave on Santa Cruz Island (Galapagos Islands). Thus, both genera occur in
totally different regions and habitats than Andeepia, a Southern Ocean cold-water and deep-sea species.

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Key to the genera of Pardaliscidae (modified after Karaman, 1974)

1. Pereopods 3–4 similar to gnathopods 1–2 in form, propodi broadened and strongly subchelate; antenna 1
peduncular article 1 with strong, acute distal process. ............. OCTOMANA Hendrycks and Conlan, 2003
- Pereopods 3–4 not similar to gnathopods 1–2, propodi not gnathopod like; antenna 1 peduncular article 1
lacking strong distal process.........................................................................................................................2
2. Mouthparts forming a cone-like bundle........................................................................................................3
- Mouthparts not forming a cone-like bundle, more or less quadratiform in lateral view ..............................4
3. Palp of maxilla 1 biarticulate; mandibular palp 3-articulate; maxilliped palp exceeding 1/2 of outer lobe;
maxilla 2 composed of 2 lobes .................................................................. HALICELLA Schellenberg, 1926
- Palp of maxilla 1 uniarticulate; mandibular palp vestigial, represented by one very short tubercle with 2
setae; maxilliped palp not exceeding outer lobe; maxilla 2 composed of one lobe. ......................................
............................................................................................................ RHYNOHALICELLA Karaman, 1974
4. Pereopods 3–4 gracile, ischium extremely elongated, almost as long as basis; pereopod 7 especially long
(1.5 x) compared to pereopods 5–6; uropods 1–2 very long, greater than 2X urosome length, uropod
peduncles 3X rami length and strongly spinose ..............MACROARTHRUS Hendrycks and Conlan, 2003
- Pereopods 3–4 not gracile, ischium much shorter than basis; pereopod 7 similar in length to pereopods
5–6; uropods 1–2 less than 2X urosome length, uropod peduncles much less than 3X rami length ...........5
5. Telson entire..................................................................................................................................................6
- Telson cleft....................................................................................................................................................7
6. Carpus of gnathopods 1–2 very short, about 1/7 X length of propodus, propodus elongated........................
........................................................................................................................ PARPANO J.L. Barnard, 1964
- Carpus of gnathopods 1–2 broad and much longer than propodus, propodus not elongated .........................
..............................................................................................................................EPEROPEUS Mills, 1967
7. Gnathopods 1–2 different in form, gnathopod 1 simple and gnathopod 2 subchelate; mandibular palp arti-
cle 3 minute...................................................................................................ARCULFIA J.L. Barnard, 1961
- Gnathopods 1–2 simple or subchelate; mandibular palp article 3 not minute..............................................8
8. Pereopods 3–7 prehensile ..................................................... PARAHALICE Birstein and Vinogradov, 1962
- Pereopods 3–7 simple ...................................................................................................................................9
9. Coxa 5–6 broad and deep, extending to depth of basis of pereopods 5–6; maxilla 2 vestigal, composed of
two tiny lobes provided with one seta each ............................................... NECOCHEA J.L. Barnard, 1962
- Coxa 5–6 shallow, not extending to depth of basis of pereopods 5–6; maxilla 2 well developed, composed
of two separated setose lobes......................................................................................................................10
10. Antenna 1 lacking (or scale-like?) accessory flagellum .................................. HALICOIDES Walker, 1896
- Antenna 1 with well developed accessory flagellum, composed of several articles ..................................11
11. Peduncular article 2 of antenna 1 longer than article 1..........................PARDALISCOIDES Stebbing, 1888
- Peduncular article 2 of antenna 1 shorter than article 1..............................................................................12
12. Lateral cephalic lobes slightly produced anteroventrally................................. TOSILUS J.L. Barnard, 1966
Lateral cephalic lobes not produced anteroventrally ..................................................................................13
13. Lobes of maxilla 2 short, dilated ..........................................................CALEIDOSCOPSIS Karaman, 1974
- Lobes of maxilla 2 longer and slender........................................................................................................14
14. Palp of maxilliped very long, more than 2 times as long as inner edge of outer lobe ...................................
............................................................................................................................ PRINCAXELIA Dahl, 1959
- Palp of maxilliped shorter, nearly as long as inner edge of outer lobe.......................................................15
15. Gnathopods 1–2, carpus much longer than slender propodus, dactylus short...PARDALISCA Krøyer, 1842
- Gnathopods 1–2, carpus subequal or shorter than propodus, dactylus regular...........................................16
16. Palp of maxilla 1 narrow, not dilated distally ............................................................. HALICE Boeck, 1871

24 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

- Palp of maxilla 1 dilated distally ................................................................................................................17
17. Urosome 1–2 each with a dorsal tooth; dactyls of gnathopods 1–2 smooth on inner margin ........................
.............................................................................................................. PARDALISCOPSIS Chevreux, 1911
- Urosome smooth or with only a dorsal process (sometimes double-toothed) on 1 or 2; dactyls of gnatho-
pods 1–2 with one or more teeth or spines on inner margin.......................................................................18
18. Gnathopod 1 propodus robust, palm straight, dactylus strongly curved, carpus short with weak lobe; basis
of pereopods 5–7 similar ................................................................................................ ANDEEPIA gen. n.
- Gnathopod 1 propodus smaller, palm not straight, dactylus straight to weakly curved, carpus long, with
prominent lobe; basis of pereopod 7 wider than basis of pereopod 6. .......................................................19
19. Urosome 1 with pointed and produced dorsal process (sometimes double-toothed); uropod 1 peduncle
with strong distolateral tooth ................................................................................ NICIPPE Bruzelius, 1859
- Urosome 1 smooth; uropod 1 peduncle lacking strong distolateral tooth ..................................................20
20. Peduncle of antenna 1 more than 1/2 as long as peduncle of antenna 2; gnathopods 1–2 simple, not
strongly developed.......................................................................................... PARDALISCELLA Sars, 1895
- Peduncle of antenna 1 less than 1/2 as long as peduncle of antenna 2; gnathopods 1–2 subchelate, strongly
developed....................................................................................................................................................21
21. Gnathopods 1–2 slender; basis of pereopod 6 without posteroventral lobe, pereopod 7 with a broad,
rounded posteroventral lobe; telson shallowly notched ....................ANTRONICIPPE Stock & Iliffe, 1990
- Gnathopods 1–2 broad; basis of pereopods 6–7 with a narrow posteroventral lobe; telson deeply cleft …
............................................................................................. SPELAEONICIPPE Stock & Vermeulen, 1982

Andeepia ingridae sp. n.

(Figures 1–7)

Type material. Four specimens. Holotype: Female with non-setose oostegites, 12.5 mm (catalogue number
ZIM K-41506). Locus typicus: RV Polarstern: ANT XXII/3 EBS 121-11-E: 63°38.27S–63°37.31S;
50°37.16W–50°38.04W; 14 March 2005; 2659 m; leg. (collector) Angelika Brandt.
1 paratype (a), female with non-setose oostegites, 14 mm (catalogue number ZIM K-51507); 1 paratype
(c), juvenile 5.9 mm (catalogue number ZIM K-41509) both from type locality.
1 paratype (b), female with non-setose oostegites, 15.2 mm (catalogue number CMNC 2008-0109): RV
Polarstern: ANT XXII/3 EBS 80-9-S: 70°38.45S–70°39.18S; 14°42.86W–14°43.43W; 23 February 2005;
3100 m; leg. Angelika Brandt.
Etymology. This species is named for Ingrid Biswas, the mother of the senior author.
Description of holotype. Body (Fig. 1a) slender, appearing elongate. Head without eyes, longer than
wide. Rostrum short and broad, distally pointed. Lateral cephalic lobe broadly triangular, rounded. Ventral
margin of head shallowly concave. Pereonite 1 slightly shorter than head. Pereonites 1–4 subequal in length.
Pereonites 5–7 subequal but longer than 1–4. Pleonites 1–3 elongated, longer than preceding tergites and suc-
cessively longer; epimeron 1–3 posteroventral corners rounded. Urosomite 1 longest; urosomite 2 with a dis-
tinct tooth projecting over posterior margin; urosomite 3 longer than 2.
Antenna 1 (Fig. 1c) longer than 2. Antenna 1 peduncular article 1 about 2 x as wide as articles 2–3, disto-
laterally strongly angled at insertion of article 2; article 2 0.5 x the length of article 1; length of article 3 0.60 x
the length of article 2; accessory flagellum 4–articulate; primary flagellum 33-articulate, not strongly setose.
Antenna 2 (Fig. 1b) peduncular article 1 ovoid, article 2 with rounded blunt gland cone; article 3 wider than
4–5, distolaterally strongly angled at insertion of article 4, relative lengths of articles 3–5 = 1 : 1.27 : 1; flagel-
lum 16-articulate, not setose. Labrum and lower lip damaged in holotype (see paratype a, b below).

NEW DEEP-SEA PARDALISCID AMPHIPOD Zootaxa 1977 © 2009 Magnolia Press · 25

FIGURE 1a–c,e. Andeepia ingridae sp. n., holotype female 12.5 mm (ZIM K-41506). a) habitus; b) antenna 2; c)
antenna 1; e) left mandible. d. paratype a, female with non-setose oostegites, 14 mm (ZIM K-51507). d) labrum.

26 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

FIGURE 2d–f. Andeepia ingridae sp. n., holotype female 12.5 mm (ZIM K-41506). d) maxilla 2; e) maxilla 1; f) maxil-
liped; a–c. paratype a, female with non-setose oostegites, 14 mm (ZIM K-51507). a) epistome; b) left mandible; c) right
mandible.

Mouthparts not cone-like, shallow in lateral view (Fig. 1a and in paratype 6b). Left mandible (Fig. 1e,
compare details of paratype a and b in Figs. 2b, and 7e, j,) incisor wide with some rounded teeth; left lacinia
mobilis wide, multidentate; setal row consisting of 2 raker spines and fine setae; pars molaris not present;
mandibular palp 3-articulate, length ratio of articles = 1 : 2.5 : 2.25 ; article 2 widest with 3 marginal setae;
article 3 slender, with 2 marginal setae and 2 apical setae. Maxilla 1 (Fig. 2e) inner plate damaged; outer plate
apically oblique with 8 spine-teeth; palp 2-articulate, article 1 long, length 0.77 x article 2, palp surpassing
outer plate, article 2 expanded distally with distal spines and marginal setae. Maxilla 2 (Fig. 2d) both plates
relatively wide, inner plate wider than outer plate, somewhat shorter, bordered with 9 setulated setae (only 7
shown) along the inner margin; outer plate with 3 terminal setae. Maxilliped (Fig. 2f) inner plates with 2 api-
cal setae; outer plates shorter than distal margin of 1st palp article, with 4 stout setae on oblique apices; article
3 = 1.21 x length of palp article 1; palp rather long, 4-articulate, very slender, sparsely setose; article 4 falcate,
relatively stout, inner margin not serrate, except for small subterminal notch with seta.
Coxae of all pereopods (Fig. 1a) slightly separate from each other or weakly overlapping, wider than long;
coxae 1–4 subequal in length, posteroventral margin with a fine seta. Gnathopod 1 (Figs. 3a, b) coxa short,
narrow, anteroventrally pointed; basis stout, curved, expanded distally; ischium short, subrectangular; merus

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and carpus subequal in length; carpus weakly lobate, ventrally with 5–6 setae, with 3–4 spines, about half the
length of gnathopod 2 carpus; propodus very robust, anterior margin strongly convex with only few setae,
palm occupying nearly complete straight ventral margin, bordered with setulated setae of varying lengths;
dactylus very long and strongly curved, reaching nearly the complete length of the ventral margin of propo-
dus, tip of dactylus moves to medial face of propodus when adducted; inner margin of dactylus smooth; how-
ever, under high magnification there are hints of 9 (?) teeth embedded in the cuticle (Fig. 3b).
Gnathopod 2 (Figs. 3c, d) of a different shape, propodus less robust, slightly shorter (0.87 x) and more
slender compared to preceding appendage; coxa ovoid, bluntly rounded anteriorly and posteriorly; compared
to gnathopod 1: basis longer, straighter; ischium longer than wide and tapering distally (vs wider than long
and subrectangular); merus parallelogram-shaped (vs triangular); carpus large, longer than wide, somewhat
lobate, strongly setose ventrally (vs trapezoid with fewer short setae); propodus less robust, shorter and taper-
ing distally, palm shorter and convex, with shorter setae; dactylus shorter and not as strongly curved, tip of
dactylus fits to medial pit on propodus when adducted; inner margin with 10 strongly pointed flattened teeth
directed posteriorly towards the tip of the dactylus (vs smooth margin).
Pereopods 3–4 (Figs. 3e, 4a) alike (note - pereopod 4 slightly longer than 3 and pereopod 4 carpus less
setose); coxa subrectangular, posteroventral margin with a fine seta; basis long and slightly expanded distally;
ischium longer than wide; merus anterior margin expanded into a subacute apex, posterior margin straight,
length much shorter than carpus (0.55–0.58 x); carpus and propodus subequal in length, propodus narrower,
both articles with posteromarginal setation; dactylus long and slender not strongly curved. Pereopods 5–7
(Figs. 4b–d) merus to propodus successively longer on each pereopod; coxae of pereopods 5–6 much wider
than long, that of pereopod 7 slightly indented with narrowly rounded posterior lobe; basis of 5–7 slender,
rectangular without posteroventral lobes; merus to propodus 5–7 slender with few and short setae; dactyli all
long and slender, slightly curved.
Gills (Fig. 1a) cylindrical and short, length much less than basis, found on pereopods 2–6. Oostegites
present on pereopods 2–5, not fully developed (lacking setae), somewhat narrowing distally. Pleopods strong,
both peduncles and rami long with many long, plumose setae.
Spination of uropods weak. Uropod 1 (Fig. 5a) lanceolate, peduncle long, 1.4 x rami, subrectangular, mar-
gins with short spines, with a distomedial broad tooth; rami subequal in length with few short spines, apical
third of both rami lacking setae. Uropod 2 (Fig. 5b) lanceolate, peduncle length 0.79 x length of uropod 1
peduncle, with a short distomedial tooth; rami subequal in length, apical third of both rami lacking setae. Uro-
pod 3 (Fig. 5c) short, with stout peduncle, length 0.64 x outer ramus, equipped with distoventral short flange,
and relatively short subequal flattened rami (insertion points of few setae present, setae missing, but most
likely long plumose), outer ramus with small distal segment with 2 terminal setae (see detail of Fig. 5c).
Fringes of microtrichs on the margins of peduncle and rami of all uropods. Telson (Fig. 5d) relatively short
(length 1.6 x width), cleft (68%) with narrowly rounded unnotched apices.
Details from paratype a. Epistome (Fig. 2a), triangular, without keel. Labrum (Fig. 1d) wider than long,
with shallow incision, slightly asymmetrical; both apical lobes short and wide, of about equal length. Mandi-
ble (Figs. 2b, c) incisors wide and irregularly toothed; left lacinia mobilis (Fig. 2b) wide, 4-dentate; spine row
on right mandible (Fig. 2c) with only 1 raker spine. Lower lip with wide hypopharyngeal gap; lobes of lower
lip blunt (damaged?). Maxilla 1 inner plate short, triangular, with 1 long terminal seta. Maxilla 2, inner margin
of inner plate with 10 setae.
Details from paratype b. Mandible (Figs. 7d, e, h, j, k) incisors very slightly asymmetric, margins wide
with 4 irregular teeth, anteroventral corners with a large pointed tooth; left mandible (Figs. 7e, j) lacinia mobi-
lis very wide, triangular, irregularly 7–8 dentate; spine row with 2 curved raker spines and proximal tuft of
setae; palp article 3 with 2 marginal setae and 3 apical setae; right mandible (Figs. 7d, h, k) with apparent lac-
inia as a cluster of 2 broad spines; spine row with 1 raker spine and proximal tuft of straight setae, palp article
2 with 4 marginal setae. Lower lip (Fig 7c), outer lobes rounded. Maxilla 1 (Figs. 7l, m) palp article 1,

28 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

FIGURE 3a–e. Andeepia ingridae sp. n., holotype female 12.5 mm (ZIM K-41506). a) gnathopod 1; b) dactylus of gna-
thopod 1; c) gnathopod 2; d) dactylus of gnathopod 2; e) pereopod 3.

NEW DEEP-SEA PARDALISCID AMPHIPOD Zootaxa 1977 © 2009 Magnolia Press · 29

FIGURE 4a–d. Andeepia ingridae sp. n., holotype female 12.5 mm (ZIM K-41506). a) pereopod 4; b) pereopod 5; c)
pereopod 6; d) pereopod 7.

30 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

FIGURE 5a–d. Andeepia ingridae sp. n., holotype female 12.5 mm (ZIM K-41506). a) uropod 1, detail shows distome-
dial process; b) uropod 2; c) uropod 3, detail shows apices of rami, outer ramus with minute article; d) telson; e–h. para-
type b, female with non-setose oostegites, 15.2 mm (CMNC 2008-0109). e) uropod 1, detail of distomedial peduncular
tooth; f) uropod 2, detail of distomedial peduncular tooth; g) telson; h) uropod 3.

NEW DEEP-SEA PARDALISCID AMPHIPOD Zootaxa 1977 © 2009 Magnolia Press · 31

FIGURE 6a–g. Andeepia ingridae sp. n., paratype b, female with non-setose oostegites, 15.2 mm (CMNC 2008-0109).
a) right aspect of anterior part of body with head and gnathopod 1; b) left aspect of head with buccal mass; c) antenna 1,
detail shows distal articles of accessory flagellum; d) left aspect of pleon and urosome (pleopod peduncles shown with-
out rami), detail shows urosomite 2 tooth; e) propodus of gnathopod 2, detail shows inner margin of dactylus (8 of 10–11
teeth shown); f) right aspect of coxae 3–6, with coxa-merus of pereopods 3–4; g) overview of gnathopod 2.

32 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

FIGURE 7a–m. Andeepia ingridae sp. n. paratype b, female with non-setose oostegites, 15.2 mm (CMNC 2008-0109).
a) labrum; b) apex of outer plate of maxilliped; c) lower lip; d) lateral view of right mandible, incisor and lacinia; e) left
mandible, incisor and lacinia mobilis; f) right side of maxilliped; g) rostrum; h) right mandible, incisor and lacinia (?); i)
maxilla 2; j) left mandible; k) right mandible, 3rd article of palp not shown; l, m) maxilla 1 (l is missing inner plate).

NEW DEEP-SEA PARDALISCID AMPHIPOD Zootaxa 1977 © 2009 Magnolia Press · 33

length 0.56 x article 2. Maxilla 2 (Fig. 7i) inner plate narrowing distally and truncate, with 3 distal setae and
7–8 inner marginal setae. Maxilliped (Figs. 7b, f) outer plate just reaching distal inner margin of 1st palp arti-
cle, with 3 stout setae and 1 long inner marginal seta on oblique apices; article 3 = 1.44 x length of palp article
1.
Antenna 1 (Figs. 6a, c) peduncular article 2 0.57 x length of article 1; length of article 3 0.56 x length of
article 2; accessory flagellum 5-articulate; primary flagellum 25-articulate. Ventral margin of head (Figs. 6a,
b) at antenna 2 insertion shallowly concave. Antenna 2 (Fig. 6b) article 2 with narrowly rounded gland cone;
relative lengths of peduncular articles 3–5 = 1 : 1.27 : 1.05; flagellum 19-articulate. Gnathopod 1 carpal lobe
ventrally with 4 short spines and 1 large spine. Oostegites (Fig. 6g) lacking setae, not fully developed, fairly
narrow, tapering distally. Urosomite 2 (Fig. 6d) dorsal tooth with a very fine seta underneath it. Uropod 3 (Fig.
5h) peduncle with distoventral flange, flange margin microserrate. Telson (Fig. 5g) length 1.68 x width, cleft
64%.
Remarks. The new taxon superficially resembles species from the genus Nicippe, such as N. tumida and
N. unidentata but due to many remarkable differences (see table 1), the new genus Andeepia was introduced
to accept it.
The other two genera in the taxonomic complex, Spelaeonicippe and Antronicippe differ from the new
species to a greater extent. Spelaeonicippe provo Stock & Vermeulen, 1982 has an elongate antenna 2 (vs
shorter in the new taxon); pereopods 6–7 with strong posteroventral lobes of basis (vs lacking lobes) and the
rami of uropods 1–2 are of unequal length (vs subequal).
Antronicippe serrata Stock & Iliffe, 1990 has a shallowly cleft telson with long marginal spines; the gna-
thopod propodi are very slender and their dactyli have serrated teeth on the inner margin; pereopod 7 basis is
rounded and strongly lobate and the rami of uropods 1–2 are strongly unequal in length.

Nicippe unidentata K.H. Barnard, 1932

(Figure 8)

Nicippe unidentata K.H. Barnard, 1932: p. 133; ? Enequist, 1949: 325. (probably Nicippe unidentata = Nicippe tumida
Bruzelius, 1859, according to J.L. Barnard, 1959); J.L. Barnard, 1958: 110; ? J.L. Barnard, 1959: 38. (probably
Nicippe unidentata = Nicippe tumida); Karaman, 1974: 23; Lowry & Bullock, 1976: 124.
? Voss, 1988: 54. (Nicippe ?unidentata); Barnard & Karaman, 1991: 578; ? Klages, 1991: 51; De Broyer & Jazdzewski,
1993: 82; De Broyer et al., 2007: 179.

Material examined: 1 male, 10 mm, from the type series designated as lectotype herein; paralectotypes: 1
ovigerous female, 11 mm; 1 female, 11 mm; 1 damaged female, approx. 12 mm; 2 males (?), 10 mm; 1 dam-
aged male, approx. 12 mm; all specimens from locus typicus: Schollaert Channel, Palmer Archipelago,
278–500 m, 14 March 1927, The Natural History Museum London, 1936.11.2 1136-1143, Discovery Expedi-
tion St. 182.
1 paralectotype, not exactly from type locality, female with setose oostegites, 14 mm (designated herein,
illustrated specimen); Schollaert Channel, Palmer Archipelago, 160–335 m, 12 March 1927, The Natural His-
tory Museum London, 1936.11.2 1144, Discovery Expedition St. 181, designated as cotype on the museum
label.
Description based on male lectotype and illustrated female paralectotype. Head without any trace of eyes
or ommatidea; with short rostrum; lateral cephalic lobe acute, with a keel starting at this sharp angle and
extending more than half the head length posteriorly; head quadrately lobate posteroventrally. Pereon seg-
ments 1–4 subequal in length, segments 5–6 longer, segment 7 longest. Pleon segments 1–2 subequal in
length, segment 3 longest. Epimeral plates 1–3 with posteroventral sharp cusps. Urosomite 1 as long as 2–3
combined, with a strong pointed tooth extending beyond the posterior segmental margin; small pointed pro-
cess ventrally at insertion of uropod 1. Urosomites 2–3 smooth, without dorsal teeth.

34 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

FIGURE 8. Nicippe unidentata K.H. Barnard, 1932. Habitus of female paralectotype, 14 mm, Schollaert Channel,
Palmer Archipelago, 12 March 1927, Discovery Expedition St. 181, 160–335 m, 1936.11.2 1144.

Antenna 1 longer than antenna 2; peduncular articles 1–3 relative lengths = 1 : 0.6 : 0.3; accessory flagel-
lum 3-articulate in male lectotype, with elongate and widened article 1 and 4-articulate in female; flagellum
article 1 stouter and longer in male. Antenna 2 peduncular article 3 twice as wide as article 4; articles 4–5 sub-
equal in length. Epistome weakly produced, rounded. Labrum wider than long, medially excavate with right
lobe longer than left one. Both mandibles with a sharp incisor with projecting tooth posteriorly; left lacinia
mobilis wide, with dentate cutting edge, on the right mandible spine-like; mandibular palp 3-articulate, article
2 longest, about 1.75 x length of article 3, article 2–3 with long setae on ventral margin. Maxilla 1 outer plate
with 7 smooth apical spine-teeth, the lateral one strongest and longest; palp 2-articulate, second article
expanded distally, with apical row of short spine-like setae; inner plate small with 1 apical seta. Maxilla 2
small, inner lobe slightly shorter than outer, with row of setae on medial margin; both lobes with long apical
setation. Maxilliped inner lobes narrow and pointed with long apical setae; outer plate slender, extending up
the distal margin of palp article 1, with setae along medial margin; palp long, robust, article 2 longest and wid-
est, densely covered with setae posteromarginally; article 4 slender with serrate inner margin.
Coxal plates all wider than long, not much overlapping. Gnathopod 1 coxa with rounded anterior project-
ing lobe; basis expanded distally, anterior margin widened and flat, triangular in cross section, posterior mar-
gin with groups of long setae; ischium triangular, subequal in length to merus; carpus with wide, rounded
ventrally projecting lobe, with long setae posteromarginally and medially; propodus about as wide as carpus,
tapering distally, straight palm with long setae; dactylus slender, slightly curved, with one proximal tooth on
inner margin. Gnathopod 2 coxa subrectangular; compared to that of gnathopod 1; basis longer, more slender,
less expanded; ischium shorter than merus; carpus more slender, but with similar groups of long setae; propo-
dus and dactylus of similar shape but slightly shorter. Pereopods 3–4 similar in shape and length, both with
long setation, but pereopod 3 with slightly wider basis to propodus, propodus slightly shortened; coxa 3–4
subequal to coxa 2; basis slender, anterior margin straight, posterior margin weakly convex; ischium small
with anteromarginal notch; merus expanded distally, anterodistal angle acutely drawn out; carpus as wide as
merus, anterior margin of carpus produced; propodus slender, less than 0.5 x width of carpus; dactylus long,

NEW DEEP-SEA PARDALISCID AMPHIPOD Zootaxa 1977 © 2009 Magnolia Press · 35

slender and weakly curved. Pereopod 5 much shorter than 6 but of similar shape, pereopod 7 longest, slightly
surpassing ends of all uropods. Pereopod 5 coxa bilobate, anterior lobe stronger and slightly longer; basis pos-
terior margin straight and flat, posteroventral corner with short rounded lobe; ischium shortest with antero-
marginal notch, merus to propodus relative lengths = 1 : 0.7 : 0.83; dactylus slender and straight, longer than
on pereopods 3–4. Pereopod 6 coxa bilobate, both lobes alike; basis longer compared to pereopod 5; merus to
propodus relative lengths = 1 : 0.76 : 1.22; dactylus as for pereopod 5. Pereopod 7 coxa short, tapering poste-
riorly; basis anteroproximally widened, posterior margin straight and flat with long setae, posteroventral cor-
ner roundly lobate; merus to propodus relative lengths = 1 : 1.2 : 1.2.

FIGURE 9. a–f. Main diagnostic differences between Nicippe unidentata (a, c, e) and Nicippe tumida (b, d, f, redrawn
from Karaman, 1993). a, b) urosome, left aspect: development of dorsal process on urosomite 1 and position of distal
process of uropod 2 peduncle; c, d) length of distal article of mandibular palp; e, f) distal margin of uropod 3 peduncle.

Uropod 1 peduncle slightly shorter than rami, distolateral peduncular tooth very strong; outer ramus
somewhat longer than inner ramus, both rami slightly surpassing the uropod 2 apex; rami margins bordered
with relatively long spine-like setae, with stout spine at tip of both rami. Uropod 2 of similar shape as uropod
1, but both peduncle and rami shorter than on preceding appendage; distolateral peduncular tooth about half
the length of that on uropod 1, situated rather dorsally; rami subequal with long spine-like setae. Uropod 3
peduncle with 3 pointed processes dorsally; outer ramus slightly longer than inner; medial margins of both
rami with long setulated setae. Telson deeply cleft (almost down to the base) with straight margins of incision,
with a long stout seta proximodorsally and a few laterally on both sides; apex of each telson lobe incised, lat-
eral part of apex slighter longer than medial part.

36 · Zootaxa 1977 © 2009 Magnolia Press BISWAS ET AL.

 Remarks. The status of Nicippe unidentata K.H. Barnard, 1932, has never been clear (see synonymy).
Much of the taxonomic confusion was due to the fact that K.H. Barnard (1932) gave a very brief description
of only 6 lines of text, stating the overall similarity to Nicippe tumida Bruzelius, 1859, but unfortunately, pro-
viding no illustrations for comparison. Later descriptions of N. tumida (see synonymy) addressed the possible
synonymy, but never resolved the problem. Enequist (1949) considered the south Atlantic N. unidentata to be
synonymous with the considered cosmopolitan N. tumida, as he found two forms of this species: one with the
typical dorsal bidentate urosomite 1 and one with only a single dorsal tooth. In the meantime collections made
during polar ecological surveys (Voss 1988; Klages 1991) were found to contain this pardaliscid, but the spec-
imens could not be identified with certainty. As N. unidentata has never been studied in detail since Barnard’s
description, the need for a decision on the validity of the species is long overdue.
After our examination of the type series, we are convinced that N. unidentata is a valid and distinct spe-
cies. However, the situation of N. tumida seems to be more complicated and this cosmopolitan taxon may yet
be a complex of several local (North Atlantic, Mediterranean, north east Pacific, South Africa) species.
Nicippe unidentata differs from this complex by several subtle characters (compare Fig. 9) which are
especially well visible in the female paralectotype, the largest specimen collected by K.H. Barnard: the char-
acteristic shape of the mid-dorsal process of urosomite 1, which is large and drawn out into a thin acute pro-
cess (vs a rather angular process); mandibular palp article 3 is shortened, article 2 is 1.75 x as long as this
article (vs 1.18 x); maxilliped article 2 appears to be expanded (vs more slender); pereopod 5–6 coxae are
clearly bilobed and excavated ventrally (vs almost straight); pereopod 7 coxa is tapering posteriorly (vs sub-
rectangular shape); the peduncle of uropod 2 has a straight, distolateral tooth (vs a curved, distolateral tooth);
uropod 3 peduncle with 3 dorsoapical teeth (vs none) and the telson cleft has straight inner margins of incision
(vs sinuous inner margins).

Acknowledgements

We would like to express our thanks to Prof. Angelika Brandt and Dr. Hans-Georg Andres (Zoological Insti-
tute and Museum of the University of Hamburg) for providing the material for this study. Special thanks to
Miranda Lowe (The Natural History Museum, London) for allowing us to study the type material of Nicippe
unidentata K.H. Barnard, 1932. Judith Price (Canadian Museum of Nature, Ottawa) provided the catalogue
number for the paratype deposited at the CMN.

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