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Zootaxa 2579: 45–58 (2010) ISSN 1175-5326 (print edition)

www.mapress.com / zootaxa/ Article ZOOTAXA
Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)
Larval development of the shrimp Hippolyte sapphica d’Udekem d’Acoz, 1993

forma A and B (Decapoda: Caridea: Hippolytidae) reared in the laboratory,
confirmation of the conspecific status of the two forms
ALEXANDROS NTAKIS, CHRYSSA ANASTASIADOU1, ROMAN LIASKO

& IOANNIS D. LEONARDOS2
Laboratory of Zoology, Department of Biological Applications and Technology, University of Ioannina, GR-451 10, Ioannina, Greece.
Corresponding authors. E-mail:2ileonard@uoi.gr; 1chanasta@cc.uoi.gr
Abstract
The complete series of larval staging of Hippolyte sapphica d’Udekem d’Acoz, 1993 forma A and B from Louros
estuary was studied in the laboratory and described in detail for the first time. The reared larvae of H. sapphica passed
through six zoeal stages and one megalopal stage. The larval monitoring completed when the individuals could be
assigned clearly to form A and B via rostra formation. Under the experimental conditions, the average durations of the
larval stages were as follows: three days for 1st and 2nd stages, three to four days for 3rd, 4th, 5th and 6th stages, four days for
the megalopal stage and 19 to 30 days for immature form A and B individuals. Comparison of the larval morphological
characters among the described material and the bibliographic data of closely related species was made and discussed.
The offspring of females of the forma A includes forma A and B and the same can be said of the offspring of the forma
B. This confirms that the formae A and B are indeed conspecific.
Key words: Hippolyte, shrimp, Decapoda, larval development, Greece
Introduction
The Mediterranean shrimp Hippolyte sapphica d’Udekem d’Acoz, 1993 consists of two forms, form A and
form B (d’Udekem d’Acoz, 1996, 2007). The systematic position of the two forms has been considered as
problematic (d’Udekem d’Acoz, 1996) because the only taxonomical difference was detected on the rostrum
morphology. The form A bears a very long dentate rostrum, while form B has a very reduced, toothless
rostrum. d’Udekem d’Acoz (1996) supported that “Considered alone these data would suggest that the two
forms are different species. However it appears – the rostrum expected- that all the morphological structures
of the two forms are perfectly identical and that specimens of the two forms from the same station have also
the same average carapace length”. Moreover, the author presupposes that the two forms belong to the same
species according to their distribution pattern and to their specific extracted ratios in different habitats. No
further publications on this field have been made since then, in order to elucidate the taxonomical status of the
two forms.
H. sapphica forma A is an endemic Mediterranean and Black Sea species which has been reported from
the Adriatic, the Ionian, the Aegean and Black Seas (d’Udekem d’Acoz, 1993, 1996, 1999; Koukouras and
Anastasiadou, 2002). H. sapphica forma B is an endemic species in the central Mediterranean and has been
reported only from the Ionian Sea (Gulf of Amvrakikos) and the northern Adriatic Sea (Venice lagoon)
(d’Udekem d’Acoz, 1996). In the Amvrakikos Gulf seagrasses loci the two forms always have been found
together and demonstrate similar biology (D’ Udekem d’ Acoz, 1996; present data). The species seems to
prefer very sheltered biotopes such as lagoons and closed gulfs with shallow depths ranging from 0.3 m tο 1.5
m. H. sapphica populations have been collected from small seagasses (Zostera marina and Cymodocea
nodosa) and also from Cystoseira (D’ Udekem d’ Acoz, 1996; Koukouras and Anastasiadou, 2002).
Accepted by J. Goy: 13 Jul. 2010; published: 30 Aug. 2010 45

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The study of H. sapphica larval development arises firstly as a necessity in order to elucidate the
taxonomic status of the two forms. Moreover, the larval staging of the species is given for the first time,
providing both new and useful information on the meroplankton of the Amvrakikos Gulf.
Method
Ovigerous female specimens of H. sapphica were collected in June 2009, by means of a hand net with a frame
of 30 cm X 35 cm and a mesh size of 2 mm. Sampling was carried out in Louros River estuary, next to
Amvrakikos Gulf (39°13.961′N, 20°45.971′E, NW Greece). The habitat was a shallow, muddy, saltwater pond
with rich aquatic vegetation (Cystoseira and Enteromorpha species) and a depth ranging from 0.4 m to 2 m.
Temperature and salinity were measured in situ by the use of portable instruments. The average water
temperature and salinity were 20.5 oC (±0.5 oC) and 16.9 ppt (±0.4 ppt) respectively.
Egg bearing females (25 and 28 individuals from forma A and B, respectively) and habitat water were
transferred to the laboratory in plastic containers. Three types of aquaria were constructed and filled with
aquatic vegetation and water from the habitat, filtered through plankton net, with a mesh size of 125μm. H.
sapphica forma A and B were placed in different aquaria at 25oC and oxygen supplied via air pumps.
After hatching, the females were disposed of and their carapace and total lengths were measured while the
larvae were kept in aquaria in the same conditions. Carapace and total lengths of ovigerous females of forma
A were ranged from 2.68 mm to 3.48 mm and from 10.24 mm to 14.73 mm respectively. Carapace and total
lengths of ovigerous females of forma B were ranged from 2.73 mm to 3.79 mm and from 10.37 mm to 15.11
mm respectively. Through their development, the larvae were fed with natural phytoplankton and rotifers.
After each moult, several larvae were photographed and preserved in 5% formalin solution, n with their
excuviae. All of the specimens are deposited in the Laboratory of Zoology Collection, in the Department of
Biological Applications and Technology, in the University of Ioannina, Greece. The time period between each
moult and the subsequent was recorded for determining the duration of each stage. Larvae mortalities were
very low. The larval anameric development of H. sapphica was investigated up to the stage were the rostrum
discrimination was observed. Larval measurements were accomplished by the means of an optical microscope
(Axioscope 40, Zeiss). The 1st stage is described in detail and only the changes in the preceding stages are
presented below. Total length was measured from the orbit to the end of the telson and carapace length was
calculated from the orbit to the posterolateral margin of the carapace. Descriptive accounts of all appendages
for each larval stage are given according to the proposed methodology after Clark et al. (1998). The current
chromatophore topography and characterization made after the primary system of chromatophore
characterization which was proposed by Keeble and Gamble (1904) (Keeble and Gamble, 1904, Lebour,
1928; Aikawa, 1929; Huus, 1935).
Results
The reared larvae of H. sapphica passed through six zoeal stages and one megalopal stage. These larval stages
are presented in detail, along with the stage where the rostrum morphology allowed the discrimination of the
forma A and B. Under the experimental conditions, the average durations of the larval stages were as follows:
three days for 1st and 2nd stages, three to four days for 3rd, 4th, 5th and 6th stages, four days for the megalopal
stage and 19 to 30 days for immature form A and B individuals. The descriptive accounts and the figures of
the appendages of each larval stage are presented below.
Descriptive accounts
Stage I (Fig. 1) Carapace with supraorbital and pterygostomian pairs of spines. Rostrum short, wide at the
base, sharp at the tip, slightly directed ventrally, slightly overreaching the proximal segment of antennular
46 · Zootaxa 2579 © 2010 Magnolia Press NTAKIS ET AL.

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peduncle (Fig. 1A). Eyes sessile (Fig. 1A). Pleopods present as rudiments (Fig. 1A). Abdomen with 6 somites,
sixth somite fused to telson, fifth somite without posterolateral spine (Fig. 1A). Telson, posterior width about
2.7 times anterior width; posterior median notch (Fig. 1B); posterior margin of telson with 7 + 7 spines,
innermost (1st pair) smallest than others: 2nd pair of spines 2.16 times longer than the 1st, 3rd pair 1.16 times
longer than 1st, 4th pair 2.30 times longer than 1st, 5th pair 2.4 times longer than 1st, 6th pair 2.27 times longer
than 1st and 7th pair 1.4 times longer than 1st; 7th pair of spines bears simple setae only in the inner margin while
other pairs bear setae at both sides. Uropods absent. Antennule (Fig. 1C) biarcticulated with long proximal
segment; inner setose flagellum 0.64 times shorter than antennular peduncle; outer flagellum 0.25 times
shorter than peduncle with two long distal simple setae, and one shorter subdistal. Antenna (Fig. 1D):
endopod bearing a triangular strong spine at its distal end; scaphocerite (exopod) 0.88 times shorter than
endopod with 10 long plumose setae along the distal and subdistal margin. Mandible (Fig. 1E) lacking palp,
with molar process well developed and incisor bearing 3 sharp teeth; molar with apical waved surface and
numerous small and large teeth; one well developed, triangular inner posterolateral tooth; tree median teeth
between incisor and molar processes. Maxillula (Fig. 1F): coxal endite with 6 strong simple setae; basial
endite bearing 4 strong triangular teeth; well developed palp (endopod) bearing one subapical plumose setae
and two apical tufts with 4 and 5 simple setae each one; exopodal setae absent. Maxilla (Fig. 1G): coxal endite
bilobed, with proximal and distal lobes bearing 6 and 5 plumose setae, respectively; basial endite divided into
two lobes: proximal lobe with 4 apical plumose setae and distal lobe with 5 plumose setae; endopod with 5
plumose setae, all plumose setae of the basial endite and endopod are spiniform at their distal end;
scaphognathite (exopod) well developed bearing 4 plumose setae, one strong at the basal end and numerous
simple setae on its inner margin. First maxilliped (Fig. 1H): coxa bearing 3 pairs of marginal plumose setae;
basis with 4 pairs of plumose marginal setae and one distal plumose setae; endopod 4-segmented, with setal
formula 3, 1, 2, 4; exopod 3-segmented: proximal segment long, lacking setae, median segment short, 0.2
times shorter than proximal with one long plumose setae and distal segment poor developed, bearing 3 apical
long plumose setae. Second maxilliped (Fig. 1I): endopod 4-segmented, with setal formula 2,2,3,4; basis with
2 plumose setae; exopod 3-segmented with 5 plumose setae at the distal end and one subapical small plumose
setae. Third maxilliped (Fig. 1J): coxa and basis without seta; endopod 4-segmented with setal formula 0, 0, 2,
3; exopod similar to those of previous maxillipeds with 3 apical plumose setae and 2 plumose setae on the
anterior margin of the median segment. One pair of biramous pereiopods and four rudimentary pereiopods
(Fig. 1K); endopod of the first pereiopod 4-segmented; distal segment with a strong claw and two long apical
plumose setae; median segment with two small plumose setae on the inner margin of the articulation and one
smaller on the outer margin; exopod well developed, 0.75 times shorter endopod, bearing 3 long plumose
apical setae and two plumose subapical, bilateral setae.
Red chromatophores (Fig. 1A) distributed over body as follows: 1 subapical on each antennal scale, 1
median on each anntenular penduncle, 4 pairs on each sessile eye and 1 chromatophore on the right: 1 optic
pair, 1 well developed median ocular pair, 1 post ocular pair and 1 corneal pair. 4 pairs on carapace and 1
chromatophore on the right dorsal pancreatic region: 1 supra-cerebral pair, 1 post-cerebral pair, 1 post-cardiac
pair and 1 periopodal pair. 1 abdominal pair on the third segment. 4 pairs on telson: 1 well developed basial
pair, 1 sub-basial pair and 2 lateropaical pairs.
Stage II (Fig. 2): Eyes stalked (Fig. 2A) with ocular penduncle 2.37 times longer than cornea. Pleopods 3-
segmented. Telson (Fig. 2B) posterior median indentation shallower than that of stage I; posterior margin with
8 + 8 plumose spines. Antennular peduncle (Fig. 2C) 3-segmented with setal formula 2,3,0; proximal segment
2.48 times longer than median with a strong marginal spine near the base; median segment 1.3 times longer
than distal; endopod as a long plumose seta; exopod bearing one strong simple setae and tree long plumose
setae. Antenna (Fig. 2D): endopod with 1 small simple setae near the base, 1.33 times longer than exopod;
scaphocerite (exopod) with 13 plumose setae and one strong outer spine; Maxillula (Fig. 2E): coxal endite
with 5 plumose setae and 1 plumose setae on a submarginal lobe; basal endite bearing 8 strong triangular
teeth; endopod unchanged. Maxilla; coxal endite bilobed with the proximal lobe bearing 4 pairs of plumose
setae and 2 plumose setae and the distal lobe with 2 pairs of plumose setae; basial endite bilobed with
proximal lobe with 5 plumose setae, distal lobe with 3 plumose setae and 2 pairs of plumose setae; endopod
LARVAL DEVELOPMENT OF HIPPOLYTE SAPPHICA Zootaxa 2579 © 2010 Magnolia Press · 47

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with 2 pairs of plumose setae; scaphognathite (exopod) bearing 6 plumose setae. First maxilliped with 3 apical
long plumose setae on the distal segment of the exopod. Second maxilliped: endopod 5-segmented, with setal
formula 3, 2, 1, 4, 4; exopod with 2 plumose setae on the distal part of median segment and 4 plumose setae
on the terminal end of the distal segment. Third maxilliped (Fig. 2F): endopod 4-segmented with setal formula
2,1,3,4; exopod similar to those of previous maxillipeds with 4 apical plumose setae and 2 plumose setae on
the distal margin of the median segment. One pair of biramous pereiopods (Fig. 2G) and four rudimentary
pereiopods. Endopod of the first pereiopod 4-segmented; distal segment modified to chela; exopod 2-
segmented, with 3 plumose setae on the terminal end and 2 plumose setae near the base of the distal segment.
Stage III (Fig. 2): Posterior margin of telson (Fig. 2H) with 7 + 7 plumose spines. Uropods free with
exopod 0.68 times shorter than telson. Pleopods (Fig. 2I) 3-segmented with 2 apical simple small setae.
Antennal endopod (Fig. 2J) 9-segmented with setal formula 1,0,0,0,0,0,0,0,2; 0.93 times shorter than
scaphocerite. Maxilla (Fig. 2K): coxal endite bilobed, proximal and distal lobes with 6 and 2 plumose setae,
respectively; basial endite bilobed, proximal lobe with 4 pairs of plumose setae, distal lobe with 3 pairs of
plumose setae and 2 plumose setae; endopod with 4 plumose setae; scaphognathite (exopod) bearing 7
plumose setae. Second maxilliped (Fig. 2L): basis with 3 marginal pairs of plumose setae and 1 marginal
posterior plumose setae; endopod 4-segmented, with setal formula 3,1,2,4; exopod with 4 terminal and 2
subterminal plumose setae;. First pereiopod biramous (Fig. 2M); endopod 4-segmented, chela more
developed with 3 simple setae; exopod nude and unsegmented. Second pereiopod 4-segmented (Fig. 2N);
distal segment modified to chela, bearing 4 simple setae. Third, fourth and fifth pereiopods unsegmented with
2 strong spines on the tip (Fig. 2N). Other segments unchanged.
Stage IV (Fig. 3): Eyes with ocular penduncle 2.54 times longer than cornea. Pleopods (Fig. 3A) 3-
segmented with 4 apical, long plumose setae and 1 subapical plumose setae on the distal segment. Telson (Fig.
3B) bearing 1 pair of dorsalolateral spines. Uropods biramous, endopod nacked, 0.57 times shorter than
exopod and 0.55 times shorter than telson; exopod bearing 9 plumose setae, 0.97 times shorter than telson.
Antennular peduncle 3-segmented with setal formula 2,2,0; proximal segment 3.72 times longer than median
with a strong marginal spine near the base; median segment 1.14 times longer than distal; outer flagellum
bearing 2 long and 1 small simple setae. Antennal endopod (Fig. 3C) 10-segmented, each segment bearing
one pair of aesthetasks, distal segment bearing 1 aesthetask and 2 simple setae; 3.5 times longer than
scaphocerite. Maxillula (Fig. 3D): basial endite bearing 12 strong triangular teeth and 5 plumose setae; coxal
endite with 7 plumose setae; palp (endopod) bearing one apical simple setae. Maxilla (Fig. 3E): coxal endite
bearing 5 plumose setae; basial endite bilobed with proximal lobe bearing 3 pairs of plumose setae and 3
plumose setae on a submarginal small lobe, distal lobe with 3 pairs of plumose setae and one plumose setae;
well developed palp (endopod) bearing 1 apical plumose setae; scaphognathite (exopod) bearing 6 long and
one short plumose setae; First maxilliped (Fig. 3F): protopodite bearing 10 pairs of marginal plumose setae;
endopod 2-segmented, with setal formula 5,1; exopod 3-segmented with setal formula 1,0,4; median segment
long, 1.2 times longer than proximal and 2.7 times longer than distal. Second maxilliped (Fig. 3G): base
bearing 4 marginal plumose setae; endopod 4-segmented, with setal formula 0, 0, 1, 5; exopod 3-segmented
with setal formula 0, 1, 4; median segment long, 22.25 times longer than proximal and 4 times longer than
basial. Third maxilliped (Fig. 3H): endopod 4-segmented with setal formula 10, 2, 4, 0; distal segment bearing
4 spines; exopod unsegmented with 4 apical plumose setae and 1 simple setae on the outer margin. First
pereiopod (Fig. 3I): 4-segmented with setal formula 0, 0, 2, 11; chela strong and robust. Second pereiopod
(Fig. 3J): 4-segmented with setal formula 0, 0, 0, 9; chela well developed. Third pereiopod (Fig. 3K) 4-
segmented with setal formula 1, 3, 10, 2; basis and coxa well developed bearing 0 and 1 simple setae
respectively; propodus with one pair of ventral spines; dactylus with 1 ventral and 2 strong apical spines.
Fourth and fifth pereiopods same as the third.
Stage V (Fig. 4): Telson (Fig. 4A) margins almost parallel; posterior width of telson slightly wider than
anterior; posterior margin with 5 + 5 spines and two pairs of dorsolateral spines. Uropod’s (Fig. 4B) exopod
bearing 11 plumose setae and 1 distolateral spine; endopod with 7 plumose setae, 0.85 times shorter than
exopod. Antennular peduncle 3-segmented with setal formula 2,4,4; proximal segment 4.1 times longer than
median with a strong marginal spine; median segment 1.3 times longer than distal; outer flagellum bearing 1

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aesthetask and 1 simple setae. Scaphocerite (exopod) with 15 plumose setae and 1 distolateral spine; endopod
11-segmented, 4.1 times longer than scaphocerite. Maxillula: coxal endite with 6 apical plumose setae and 1
plumose setae on the outer margin; basial endite bearing 12 strong triangular teeth and 4 plumose setae; palp
(endopod) unchanged. First maxilliped with protopodite bearing 19 marginal plumose setae; endopod
unsegmented, with one terminal plumose setae; exopod unsegmented, 2.1 times longer than endopod, with 4
subbasal plumose setae and 4 terminal plumose setae. Second maxilliped with protopodite bearing 5 plumose
setae and 2 simple short setae; endopod 4-segmented, with setal formula 0, 0, 3, 5; exopod unsegmented, 1.9
times longer than endopod, with 3 apical and 1 subapical plumose setae. Third maxilliped (Fig. 4C): endopod
3-segmented and setal formula 2, 3, 6; distal segment bearing 5 spines; exopod unsegmented with 4 apical
plumose setae and 1 simple setae on the outer margin. First (Fig. 4D) and second pereiopods (Fig. 4E) 4-
segmented with setal formulas 0, 0, 0, 19 and 0, 0, 0, 14, respectively.
Stage VI (Fig. 4): Telson with 3 dorsolateral spines (Fig. 4F). Endopod and exopod of uropods with 9 and
14 plumose setae, respectively; endopod 0.85 times shorter than exopod. First maxilliped (Fig. 4G):
protopodite bearing 13 marginal and 3 submarginal plumose setae; endopod unsegmented, with 1 distal
plumose setae, 1 terminal simple setae and 1 proximal plumose setae; exopod unsegmented with 4 proximal
and 3 distal plumose setae; exopod 2.1 times longer than endopod. Second maxilliped (Fig. 4H): protopodite
bearing 7 plumose setae; endopod 4-segmented, with setal formula 1, 0, 3, 6; exopod unsegmented with 4
apical plumose setae and 1 subapical simple setae; exopod 1.9 times longer than endopod.
Stage VII (Megalopa) (Fig. 4): Pleopods biramous (Fig. 4I); endopod bearing 4 apical and 3 subapical
plumose setae; exopod with 2 apical plumose setae. Telson (Fig. 4J) margins parallel, with 3 pairs of well
developed dorsolateral spines. Uropod’s exopod with 21 plumose setae and 1 distolateral spine; endopod with
17 plumose setae, almost equal to exopod. Antennula (Fig. 4K) 3-segmented with setal formula 3, 4, 0;
proximal segment 2.66 times longer than median with a strong marginal spine near the base and a well
developed stylocerite; median segment 1.3 times longer than distal; inner flagellum with 2 apical simple setae;
outer flagellum 2-segmented with setal folmula 1, 1; 3 terminal aesthetasks. Antenna: scaphocerite (exopod)
with 17 plumose setae; endopod 15-segmented. Second maxilliped: protopodite bearing 7 plumose setae;
endopod 4-segmented, with setal formula 1, 0, 4, 7; exopod unsegmented with 3 apical and 1 subapical
plumose setae; exopod 1.9 times longer than endopod.
Stage Immature (Figs. 5 and 6): Form A: rostrum long, narrow, inclined slightly downwards with one
median dorsal tooth (Fig. 5A and 5B). Form B: rostrum short, reduced to a toothless spine (Fig. 5C). Sex
discrimination: Males’ first pleopod with endopod bearing 2 apical plumose setae and 1 simple setae (Fig.
5D); exopod with 7 plumose setae. Males’s second pleopod with endopod bearing 5 plumose setae and at its
inner margin a short appendix masculina with 1 plumose setae and an outer longer appendix interna (Fig. 5E);
exopod with 9 plumose setae. Females’ first pleopod (Fig. 5F) with endopod bearing 4 plumose setae; exopod
with 7 plumose setae. Females’ second pleopod with endopod bearing 4 plumose seate and an appendix
interna on its inner margin; exopod with 9 plumose setae (Fig. 5G). Anterior width of telson (Fig. 5H) slightly
wider than posterior with lateral sides almost parallel. Uropods’ endopod and exopod (Fig. 5I) with 16 and 19
plumose setae, respectively; endopod 0.95 times shorter than exopod. Antennula (Fig. 5J) 3-segmented with
setal formula 5, 6, 3; proximal segment 2.73 times longer than median with a strong well developed
stylocerite; median segment 1.15 times longer than distal; inner flagellum with 4 apical and 2 subapical
simple setae; outer flagellum 3-segmented with setal folmula 0, 2, 4; distal and median segments with 2
aesthetasks for each one. Antennal endopod 26-segmented with a pair of aesthetasks on each one;
scaphocerite (exopod) bearing 28 plumose setae. Maxilla (Fig. 6A): coxal endite with 4 plumose setae; basial
endite bilobed with the proximal lobe bearing 9 plumose setae and the distal lobe bearing with 11 plumose
setae; palp (endopod) well developed with 1 simple setae; scaphognathite (exopod) bearing 24 plumose setae.
First maxilliped (Fig. 6B) with protopodite bearing 26 marginal and 14 submarginal plumose setae; endopod
unsegmented, with 4 plumose setae; exopod unsegmented with 3 distal and 7 proximal plumose setae; exopod
2 times longer than endopod. Second maxilliped (Fig. 6C): protopodite bearing 7 marginal plumose setae and
2 submarginal simple setae; endopod 4-segmented, with setal formula 3, 0, 5, 9; exopod unsegmented with 5
apical and 2 posterior plumose setae; exopod 2.24 times longer than endopod. Third maxilliped (Fig. 6D):

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FIGURE 1. Stage I. A. Whole specimen; B. Telson; C. Antennula; D. Antenna; E. Mandible; F. Maxillula; G. Maxilla;
H. 1st Maxilliped; I. 2nd Maxilliped; J. 3rd Maxilliped; K. 1st Pereiopod. Scales: A. 0.0464mm; B. 0.046mm; C. 0.0057mm;
D. 0.033mm; E. 0.02mm; F. 0.0057mm; G. 0.0215mm; H. 0.0023mm; I. 0.0044mm; J. 0.0083mm; K. 0.0215mm.

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FIGURE 2. A – G Stage II and H – N Stage III. A. Eye; B. Telson; C. Antennula; D. Antenna; E. Maxillula; F. 3 rd
Maxilliped; G. 1st Pereiopod; H. Telson; I. Pleopod; J. Antenna; K. Maxilla; L. 2nd Maxilliped; M.1st Pereiopod; N.
Pereiopods. Scales: A. 0.0731mm; B. 0.0731mm; C. 0.033mm; D. 0.044mm; E. 0.002mm; F. 0.0044mm; G. 0.0066mm;
H. 0.078mm; I. 0.018mm; J. 0.034mm; K. 0.002mm; L. 0.028mm; M. 0.0012mm; N. 0.045mm.

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FIGURE 3. Stage IV. A. Pleopod; B. Telson; C. Antenna; D. Maxillula; E. Maxilla; F. 1st Maxilliped; G. 2nd Maxilliped;
H. 3rd Maxilliped; I. 1st Pereiopod; J. 2nd Pereiopod; K. 3rd Pereiopod. Scales: A. 0.029mm; B. 0.039mm; C. 0.016mm; D.
0.021mm; E. 0.0013mm; F. 0.0044mm; G. 0.0025mm; H. 0.044mm; I. 0.017mm; J. 0.017mm; K. 0.028mm.

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FIGURE 4. A–E Stage V, F–H Stage VI and I– K Stage VII. A. Telson; B. Uropod; C. 3rd Maxilliped; D. 1st Pereiopod;
E. 2nd Pereiopod; F. Telson; G. 1st Maxilliped; H. 2nd Maxilliped; I. 1st and 2nd Pleopods; J. Telson; K. Antennula. Scales:
A. 0.015mm; B. 0.040mm; C. 0.003mm; D. 0.0011mm; E. 0.028mm; F. 0.038mm; G. 0.066mm; H. 0.028mm; I.
0.019mm; J. 0.039mm; K. 0.046mm.

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FIGURE 5. Stage Immature. A and B. Forma A Rostrum; C. Forma B Rostrum; D. 1st Pleopod (male); E. 2nd Pleopod
(male); F. 1st Pleopod (female); G. 2 nd Pleopod (female); H. Telson; I. Uropod; J. Antennula. Scales: A. 0.019mm; B.
0.019mm; C. 0.2475mm; D. 0.044mm; E. 0.044mm; F. 0.039mm; G. 0.038mm; H. 0.090mm; I. 0.090mm ; J. 0.026mm.

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FIGURE 6. Stage Immature. A. Maxilla; B. 1st Maxilliped; C. 2nd Maxilliped; D. 3rd Maxilliped; E. 1st Pereiopod; F. 2nd
Pereiopod; G. 3rd Pereiopod. Scales: A. 0.036mm; B. 0.046mm; C. 0.011mm; D. 0.038mm; E. 0.043mm; F. 0.046mm; G.
0.023mm.

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endopod 3-segmented with setal formula 12, 6, 16; distal segment bearing 5 ventral and 2 apical strong spines;
exopod unsegmented with 9 plumose setae. First pereiopod (Fig. 6E) 4-segmented with setal formula 0, 3, 3,
33. Second pereiopod (Fig. 6F): 5-segmented with setal formula 0, 0, 0, 2, 28 with basis and coxa well
developed; carpus three-arculated. Third pereiopod (Fig. 6G): 5-segmented with setal formula 2, 3, 3, 13, 2;
dactylus with 3 ventral and 2 apical spines; propodus with 5 ventral pairs of spines. Propodus of fourth and
fifth pereiopods with 3 pairs of ventral spines.
TABLE 1. Means, standard deviation (SD), minimum (min) and maximum (max) values for the measured total length
(TL) and carapace length (CL). NI: number of individuals.
Stage NI TL (mm) CL (mm)
Mean Min Max SD Mean Min Max SD
I 20 1,05 0,95 1,15 0,59 0,30 0,27 0,33 0,36
II 20 1,57 1,52 1,62 0,51 0,36 0,31 0,4 0,42
III 18 1,62 1,51 1,72 0,49 0,42 0,38 0,48 0,41
IV 15 1,68 1,59 1,79 0,56 0,47 0,41 0,54 0,37
V 15 1,75 1,42 1,93 1,35 0,53 0,45 0,59 0,49
VI 12 2,44 1,96 2,75 2,24 0,69 0,56 0,79 0,72
VII 20 2,55 2,21 3,31 2,47 0,74 0,63 0,94 0,71
Immature 18 4,52 3,05 5,85 8,81 1,01 0,72 1,25 1,66
Discussion
Some Hippolyte species like H. leptocerus (Heller, 1863) have a highly variable rostrum morphology, with a
continuum of short-rostred to long-rostred forms (d'Udekem d'Acoz, 1996). H. obliquimanus also
demonstrates four forms or morphotypes (Terrosi and Mantelatto, 2010). However, among the thirty nine
species of the genus Hippolyte (d’Udekem d’Acoz, 1996; 2007), H. sapphica is the only one which consists of
two discontinuous forms, without intermediates: form A (elongated rostrum) and form B (concave, short
rostrum). Although the rostrum morphology suggests that the two forms could be different species, the study
of the larval development revealed the opposite. Laboratory rearing showed that larvae with short and
elongate rostra are hatched in different percentages by ovigerous females of the two forms. Our present data
confirm the hypothesis of d’Udekem d’Acoz (1996) that the two forms are not different species.
Information on the complete larval development or on certain larval stages is available only for seven
Hippolyte species (H. coerulescens, H. inermis, H. pleuracanthus, H. prideauxiana, H. proteus, H. varians
and H. zostericola) till now (G.O. Sars, 1911; Webb, 1921; Gurney, 1927, 1936; Lebour, 1931; Chace, 1972;
Shield, 1978; Zupo and Buttino, 2001; González–Gordillo et al., 2001). Especially for the Mediterranean
representatives of the genus Hippolyte, the descriptive accounts for larval development focused on Hippolyte
inermis (Lebour, 1931; Williamson 1957; Bourdillon-Casanova, 1960; Le Roux, 1963; Pessani and Robotti,
1992; Zupo and Buttino, 2001). Although H. inermis and H. sapphica have been found together inhabiting
shallow, sheltered bays of the sub-littoral zone at the northeastern coast of Ionian Sea (present data), H.
sapphica prefers more closed habitats with low rate of water recycling and rich aquatic vegetation.
The comparison of H. inermis and H. sapphica larval development along with the main five generic
morphological characters of Hippolyte larvae listed by Gurney (1923) and Lebour (1931) is given in Table 2.
First of all, H. sapphica demonstrates seven larval stages (six zoeal and one megalopal stage) instead of eight
for H. inermis, given by Lebour (1931). Furthermore, the duration of each stage usually ranges from 3 to 4
days, while in H. inermis has been recorded to be lower (2 to 3 days). Moreover, the acceleration of the
formation of certain appendages such as pleopods and pereiopods is obvious in H. sapphica and is
accomplished in former stages in comparison with H. inermis. Established populations of H. sapphica forma

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A and B have been recorded in sheltered, closed habitats, where the temperature is significantly higher (2 °C)
than in less sheltered open bays of the Ionian coast. It is also well known that at temperate climates, caridean
shrimps demonstrate abbreviated development during hot summers and retarded gametogenesis and larval
development through winter due to low temperatures (Williams, 1977; Dudgeon, 1985; Hancock, 1998). This
could possibly explain why the larval development of H. inermis is elongated (8 stages) in comparison with
H. sapphica.
TABLE 2. Comparison of the morphometric characters of Hippolyte sapphica D’ Udekem d’ Acoz, 1993 to Hippolyte
inermis Leach, 1815 and to main generic characters of Hippolyte larvae according to Lebour (1931) description.
Morphological Characters Main Generic characters of Hippolyte inermis Hippolyte sapphica
Hippolyte larvae (Gurney, (Lebour, 1931; Le Roux, (present data)
1923; Lebour, 1931) 1963)
Number of larval stages 6 8 7
Duration (days) no data available 2-3, 2-3, 4,4,4,4,4,4 3,3,3-4, 3-4, 3-4, 3-4,4
Maxillula With an unjointed seta on no data available With an unjointed palp
the base, and an unjointed
palp
Maxilla With 4 lobes no data available With 4 lobes and a palp
First Maxilliped With a small epipodite no data available Not observed
1st and 2nd pair of pereiopods Rudimentary buds Rudimentary buds 1st pair biramous, 2nd pair
in first stage rudimentary
1st and 2nd pair of pereiopods no data available 1st pereiopod chelated. 1st pereiopod chelated.
in third stage Second pereiopod Second pereiopod
rudimentary segmented
Segmentation of pleopods no data available At 4th stage At 2nd stage
Acknowledgements
The authors owe special thanks to Mr. Neil Walker for the text corrections and to Mr. Kypriotis Spyros and
Chipis Art Studio for the digital elaboration of the figures of the present manuscript.
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Zootaxa 2579: 45–58 (2010) ISSN 1175-5326 (print edition)

Larval development of the shrimp Hippolyte sapphica d’Udekem d’Acoz, 1993

ALEXANDROS NTAKIS, CHRYSSA ANASTASIADOU1, ROMAN LIASKO

Key words: Hippolyte, shrimp, Decapoda, larval development, Greece

Accepted by J. Goy: 13 Jul. 2010; published: 30 Aug. 2010 45

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