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Morphological and morphometric comparisons of the first zoea of four species

of purse crabs (Decapoda: Brachyura: Leucosiidae) from the Brazilian South
Atlantic

Article in Zootaxa · January 2012

DOI: 10.11646/zootaxa.3167.1.2

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 Zootaxa 3167: 17–31 (2012) ISSN 1175-5326 (print edition)
www.mapress.com / zootaxa/
Copyright © 2012 · Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)

Morphological and morphometric comparisons of the first zoea of

four species of purse crabs (Decapoda: Brachyura: Leucosiidae)
from the Brazilian South Atlantic

GUSTAVO L. HIROSE1,2, 3, RAFAEL A. GREGATI2, ADILSON FRANSOZO2

& MARIA L. NEGREIROS-FRANSOZO2
1
Laboratório de Carcinologia, Departamento de Biologia, Centro de Ciências Biológicas e da Saúde, Universidade Federal de Ser-
gipe (UFS) Cidade Universitária Prof. José Aloísio de Campos, Av. Marechal Rondon, s/n, Jardim Rosa Elze, 49100-000 São Cris-
tóvão, Sergipe, Brazil
2
NEBECC (Study Group of Crustacean Biology, Ecology and Culture), Departamento de Zoologia, Instituto de Biociências, Universi-
dade Estadual Paulista (UNESP), 18618-000 Botucatu, São Paulo, Brazil
E-mail: [email protected]
3
Corresponging author

Abstract

Morphological descriptions of the first larval stages of Persephona mediterranea, P. lichtensteinii and P. punctata (Leu-
cosiidae: Eballinae) from the South Atlantic are provided, with comparisons among the morphometry of the species. A
detailed description and eight morphometric measurements were made for each larva, and a discriminant function analysis
was used to interpret the data set. The species differed significantly in certain morphological and morphometric features.
These characteristics may help in the identification of species of Leucosioidea found in the South Atlantic, and can be an
important tool in ecological studies, supporting the identification of planktonic morphotypes at the species level.

Key words: Persephona; larval description, larval morphometry

Introduction

Members of the superfamily Leucosioidea Samouelle, 1819, are commonly known as purse crabs. These crabs live
in subtidal marine environments, mainly inhabiting sandy bottoms of coastal regions, although some species can be
found at depths up to 400 m (Melo 1996). The superfamily comprises two families: Iphiculidae Alcock, 1896, and
Leucosiidae Samouelle, 1819, with three subfamilies, Cryptocneminae, Ebaliinae and Leucosiinae. Only the sub-
family Ebaliinae (formed by grouping subfamilies Ebaliinae, Philyrinae, and Iliinae) (Ng et al. 2008) occurs
throughout the Atlantic Ocean. Persephona Leach, 1817, is one of the most common genera of purse crabs in Bra-
zil, with four species: Persephona crinita Rathbun, 1931; P. lichtensteinii Leach, 1817; P. mediterranea (Herbst,
1794); and P. punctata (Linnaeus, 1758) (Melo 1996).
Larvae of several leucosiid genera have been described as follows: three species of Arcania Leach, 1817: A.
septemspinosa (Fabricius, 1793), by Sankolli (1961), A. undecimspinosa De Haan, 1841, by Terada (1979, 1984)
and Quintana (1984, 1986a, 1986b), and A. undecimspinosa elongata Yokoya, 1933, by Terada (1984); six species
of Ebalia Leach, 1817: E. cranchii Leach, 1817, by Lebour (1928, 1982), E. nux A. Milne Edwards, 1883, by Rice
(1980b), E. rotundata A. Milne Edwards, 1880 (described as Lithadia rotundata) by Fransozo & Bertini (2002), E.
tuberosa (Pennant, 1777), by Lebour (1928) and Salman (1982), E. tumefacta (Montagu, 1808), by Salman (1982),
and an unidentified species by Aikawa (1929); one species each of Ilia Leach, 1817and Ixa Leach, 1815: Ilia
nucleus (L., 1758), by Heegaard (1963) and Bartilotti et al. (2009) and I. cylindrus (Fabricius, 1777), by RajaBai
(1960); six species of Leucosia Bell, 1855: L. anatum (Herbst, 1783), by Terada (1984), L. biannulata Tyndale-Bis-
coe & George, 1962, by Ghory & Siddiqui (2008), L. cranicolaris (Herbst, 1783), by Quitana (1984, 1986a,
1986b), L. longifrons De Haan, 1841, by Terada (1979), L. obtusifrons De Haan, 1841, by Terada (1984), and L.

Accepted by P. Castro: 22 Nov. 2011; published: 20 Jan. 2012 17

pubescens Miers, 1877, by Tufail & Hashimi (1964);. two species of Myra Leach, 1817: M. coalita Hilgendorf,
1878, by Quitana (1984, 1986a, 1986b), and M. fugax (Fabricius, 1798), by Terada (1979, 1984); one species each
of Nucia Dana, 1852 and Persephona Leach, 1817: N. laminata (Doflein, 1904), by Quitana (1984, 1986a, 1986b),
and P. mediterranea by Negreiros-Fransozo et al. (1989); seven species of Philyra Leach, 1817: P. corallicola
Alcock, 1896, by Sankolli (1961) and Hashmi (1970), P. globosa (Fabricius, 1888), by Chhapgar (1955) and Krish-
nan & Kannupandi (1990), P. pisum De Haan, 1841, by Aikawa (1929) and Terada (1979, 1984), P. platycheira De
Haan, 1841, by Quitana (1984, 1986a, 1986b), Ko (2000) and Ghory & Siddiqui (2008), P. scabriuscula (Fabricius,
1798), by Menon (1937) and RajaBai (1960), P. syndactyla Ortmann, 1892, by Terada (1979, 1984) and Ghory &
Siddiqui (2008), and P. kanekoi Sakai 1934, by Ko (2001). Several unidentified larvae were also described by
Aikawa (1933), which he called “Leucozoeas”.
This study redescribes the first larval stage of P. mediterranea, and provides the first description of the first lar-
val stages of P. lichtensteinii and P. punctata in order to compare the species of the subfamily Ebaliinae from the
South Atlantic. Morphological and morphometric data are also provided to aid in the separation and identification
of species, including those from plankton samples.

Material and methods

Ovigerous females of Persephona mediterranea, P. punctata, and P. lichtensteinii were collected in Ubatuba Bay,
São Paulo, Brazil (23° 26'S and 45° 05'W) during October 1984, May 1989, and May 1991 respectively, using a
double-rig trawl on the sandy bottom at depths of 10 and 15 m. The crabs were transported to the laboratory in
coolers containing water from the collection site and kept alive in tanks with approximately 10 l of filtered, U.V-
irradiated seawater and under constant aeration. The temperature was maintained between 22 and 26° C, and salin-
ity similar to that at the collection site (35‰).
The crabs were inspected daily to verify the release of larvae. After hatching, the larvae were carefully
removed from the tanks with the use of a light source and a pipette and fixed in a solution of ethanol (70%) and
glycerin in a 1:1 ratio. The samples were deposited in the collection of NEBECC (Study Group of Crustacean Biol-
ogy, Ecology and Culture) under numbers 00020, 00033 and 00067 for P. mediterranea, P. punctata, and P. lichten-
steinii respectively.
Morphological descriptions. Drawings and descriptions for the zoea I of each species were made from fixed
larvae, with the aid of a Zeiss microscope equipped with a drawing tube. Approximately 10 randomly selected
specimens were used for each larval stage. The illustrations were prepared in accordance with Clark et al. (1998).
Morphometric comparison
Fifteen larvae of each species (including E. rotundata, which was also deposited in the collection of NEBECC
under the number 00052) were measured for the morphometric analysis, using a Zeiss microscope equipped with
an imaging and measurement tool. Eight different morphometric dimensions of each larva were obtained as fol-
lows: rostral spine length (RS), lateral spine length (LS); dorsal spine length (DS), distance from the lateral to the
dorsal spine (DLD), distance from the rostral to the lateral spine (DRL), carapace length (CL), carapace height
(CH), and carapace width (CW) (Fig. 1).
The data were submitted to a discriminant function analysis to determine which morphometric variables were
important in separating the groups (species). The discriminatory power of each variable was observed by Wilks'
lambda, which can range from 0 to 1, with 0 = total discrimination and 1 = absence of discrimination. Subse-
quently, a canonical discriminant analysis was performed to graphically show the separation of groups. The canon-
ical variables (roots) were tested for significance by a Chi-square test.
An analysis of variance (one-way ANOVA) with an a posteriori Tukey test was performed for each morpho-
metric variable. Tests of normality (Shapiro-Wilks) and homoscedasticity (Levene) were performed to determine if
the assumptions of the tests were satisfied. The data were transformed (log10) when necessary.

18 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

FIGURE 1. Morphometric measurements adopted for each larva analyzed.

Results

Morphological descriptions

Persephona mediterranea (Herbst, 1794)

First zoea. Carapace (Fig. 2A): Globose, small plumose setae present near base of dorsal spine. Dorsal, rostral and
pair of lateral spines present. Eyes sessile.
Antennule (Fig. 4A): Uniramous, unsegmented, conical, 3 aesthetascs + 1 simple seta terminally.
Antenna (Fig. 4A): Present as small bud, about one-third as large as antennule.
Maxillule (Figure 6A): Coxal endite, 5 plumose setae. Basial endite, 3 plumose setae + 1 plumodenticulate seta
and 1 simple seta may also be present. Endopod unsegmented, 4 simple setae terminally.
Maxilla (Figure 7A): Coxal endite bilobed, 2+3 plumose setae and microtrichia. Basial endite bilobed, 8 plu-
mose setae (4+4) and microtrichia. Endopod, 4 plumose setae (2+2) and microtrichia. Scaphognathite, 4 marginal
plumose setae, posterior part with strong apical process bearing many microtrichia.
First maxilliped (Fig. 8A): Coxa, without setae. Basis, 8 medial plumose setae (2+2+2+2) on the ventral side.
Endopodite 5-segmented, 2+2+0+2+ 5 (one subterminal and four terminal) setae. Exopod 2-segmented, 4 long plu-
mose natatory setae terminally.
Second maxilliped (Figure 9A): Coxa without setae. Basis, 4 medial setae (1+1+1+1) on inner side. Endopod
unsegmented, 3 setae. Exopod 2-segmented, 4 long plumose natatory setae terminally.
Abdomen (Fig. 3A): 5-segmented, pair of lateral knobs on posterior margin of second abdominal segment, and
second pair in middle portion of third segment. Telson wider than long, convex lateral margins with small terminal
spine. Posterior margin slightly concave, 3 pairs of plumose setae in central region.

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 19
FIGURE 2. Lateral view of the first larval stage (zoea I): A, Persephona mediterranea, B, P. lichtensteinii, and C, P. punctata.

Persephona lichtensteinii Leach, 1817

First zoea. Carapace (Fig. 2B): Dorsal, rostral and pair of curved lateral (about half size of rostral) spines present.
Eyes sessile.
Antennule (Fig. 4B): Uniramous, unsegmented, conical, 3 aesthetascs + 1 simple seta terminally. Aesthetascs
variable in size, approximately 2.5, 2 and 1.5 times size of antennules.
Antenna (Fig. 4B): Present as small bud, about half size of antennules, 1 small simple seta.
Maxillule (Fig. 6B): Coxal endite, 4 plumodenticulate setae, 3 plumose setae. Basial endite, 3 plumodenticu-
late setae + 1 simple seta; 1 small simple seta may also be present. Endopod unsegmented, 4 plumose setae termi-
nally.
Maxilla (Fig. 7B): Coxal endite bilobed, 4+4 plumodenticulate setae and microtrichia. Basial endite bilobed, 9
plumodenticulate setae (5+4) and microtrichia. Endopod, 4 plumose setae (2+2) and microtrichia. Scaphognathite,
4 marginal plumose setae, posterior part with strong apical process bearing many microtrichia.
First maxilliped (Fig. 8B): Coxa, without setae. Basis, 8 medial simple setae (2+2+2+2) on ventral side. Endo-
podite 5-segmented, 2+2+0+2+ 5 (one subterminal and four terminal) setae. Exopod 2-segmented, 4 long plumose
natatory setae terminally.
Second maxilliped (Fig. 9B): Coxa without setae. Basis, 4 medial setae (1+1+1+1) on the inner side. Endopod
unsegmented, 4 setae. Exopod 2-segmented, 4 long plumose natatory setae terminally.
Abdomen (Fig. 3B): 5-segmented, pair of lateral knobs on posterior margin of second abdominal segment, and
second pair in middle portion of third segment. Telson wider than long, lateral margins convex, with small terminal
spine. Posterior margin slightly concave, 3 pairs of plumose setae positioned in central region.

Persephona punctata (Linnaeus, 1758)

First zoea. Carapace (Fig. 2C): Evident posterior projection. Dorsal, rostral and pair of curved lateral spines pres-
ent. Eyes sessile.
Antennule (Fig. 4C): Uniramous, unsegmented, conical, 2 aesthetascs + 1 simple seta terminally. Aesthetascs
similar in size, approximately 1.5 times size of antennule.
Antenna (Fig. 4C): Present as small bud, about 1/4 size of antennule.
Maxillule (Fig. 6C): Coxal endite, 7 plumodenticulate setae. Basial endite, 4 plumodenticulate setae; 1 small
simple seta may also be present. Endopod unsegmented, 4 plumose setae terminally.

20 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

FIGURE 3. Dorsal view of abdomen of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; C, P.
punctata; and C’, P. punctata (telson enlarged, showing the detail of the simple setae present near the distal margin).

Maxilla (Fig. 7C): Coxal endite bilobed, 3+4 plumodenticulate setae and microtrichia. Basial endite bilobed, 8
plumodenticulate setae (4+4) and microtrichia. Endopod, 4 plumose setae (2+2) and microtrichia. Scaphognathite,
4 marginal plumose setae, posterior part with strong apical process bearing many microtrichia.
First maxilliped (Fig. 8C): Coxa, without setae. Basis, 8 medial simple setae (2+2+2+2) on the ventral side.
Endopodite 5-segmented, 2+2+0+2+ 4 setae. Exopod unsegmented, 4 long plumose natatory setae terminally.
Second maxilliped (Fig. 9C): Coxa without setae. Basis, 4 medial setae (1+1+1+1) on the inner side. Endopod
unsegmented, 4 setae. Exopod 2-segmented, 4 long plumose natatory setae terminally.

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 21
FIGURE 4. First larval stage (zoea I): antennule (A, Persephona mediterranea; B, P. lichtensteinii; and C, P. punctata) and
antenna (A’, P. mediterranea; B’, P. lichtensteinii; and C’, P. punctata).

FIGURE 5. Mandible of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; and C, P. punctata.

22 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

FIGURE 6. Maxillule of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; and C, P. punctata.

FIGURE 7. Maxilla of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; and C, P. punctata.

Abdomen (Fig. 3C): 5-segmented, pair of lateral knobs on posterior margin of second abdominal segment, and
second pair in middle portion of third segment. Telson wider than long, lateral margins convex, with small terminal
spine. Posterior margin slightly concave, 3 pairs of plumose setae positioned in central region; 1 small simple seta
may also be present on distal margin of telson.

Morphometric comparison

The morphometric analysis showed significant differences among species. These differences are represented by
different discriminant functions (Table 1), which showed high prediction accuracy (100%) for all species. Exami-
nation of the eigenvalues obtained by canonical discriminant analysis showed the first 3 variables (roots) as signif-
icant (p <0.001) (Table 2), and estimated the cumulative contributions of variability as 86.9%, 95.5%, and 100%
from the first to third eigenvalue, respectively. The first two canonical variables derived from the canonical dis-
criminant analysis are illustrated for each group (Fig. 10).

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 23
FIGURE 8. First maxilliped of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; and C, P. punc-
tata.

FIGURE 9. Second maxilliped of the first larval stage (zoea I): A, Persephona mediterranea; B, P. lichtensteinii; and C, P.
punctata.

24 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

TABLE 1. Classification functions, derived from the discriminant analysis for the species utilized in this study.
Variable E. rotundata P. mediterranea P. lichtensteinii P. punctata
Log RS 2.09 1.08 1.08 1.03
Log LS -0.65 -0.51 -1.19 -1.06
Log DS 2.13 1.14 0.82 1.14
Log DLD 0.03 0.14 0.05 0.04
Log DRL 1.09 0.77 0.58 0.65
Log CL 1.01 0.95 1.02 0.89
Log CH 2.98 2.96 2.87 2.67
Log CW -0.41 0.26 0.43 0.27
Constant -2097.53 -1389.76 -1163.83 -1093.04

TABLE 2. Analysis of the significance of the canonical variables (Roots) by the chi-square (χ2) test.
“Roots” Removed Eigenvalue Canonical-R Wilks’ lambda χ2 d.f. p-level
0 123.571 0.995 0.000 441.717 24 <0.001
1 9.370 0.950 0.021 195.649 14 <0.001
2 3.469 0.881 0.223 76.363 6 <0.001

d.f. = degrees of freedom

FIGURE 10. Relationship between the first two variables (canonical discriminants) for morphometric data of the species stud-
ied. The circle represents their respective confidence interval of 95%.

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 25
FIGURE 11. Morphometric variables utilized in the separation of species by discriminant analysis (µm). Letters in common,
indicate no statistically significant difference (Tukey, α = 0.05).

26 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

 Among the morphological structures utilized, based on the values of Wilks' lambda, the measurements related
to the size of the rostral, lateral, and dorsal spines were the most important in discriminating the species (Table 3).
These differences were also shown by the analysis of variance (one-way ANOVA) (Table 4 and Fig. 11). The mean
values for each morphological structure used are shown in Table 5.
The spines/ carapace width and spines/ spines ratios also revealed significant differences among species. A
clear difference was detected for E. rotundata in the ratios RS/CW, LS/CW, DS/CW, with higher values, and DS/
RS with lowest values. Higher values were found for P. mediterranea in the DS/RS and LS/RS ratios, while for P.
lichtensteinii the lowest values were found in the LS/CW, DS/CW and LS/RS ratios, whereas higher values were
found for P. punctata in the DS/RS and DS/LS ratios (Table 6).

TABLE 3. Discriminatory power of each morphometric variable and its respective significance value.
Variable Wilks’ lambda F p-value
RS 0.163 80.190 0.000
LS 0.467 17.822 0.000
DS 0.380 25.514 0.000
DLD 0.880 2.122 0.110
DRL 0.767 4.733 0.005
CL 0.843 2.905 0.044
CH 0.752 5.156 0.003
CW 0.785 4.282 0.009

TABLE 4. Results of the analysis of variance (ANOVA α = 0.05) and the a posteriori Tukey test for each variable used in the
morphometric study. Var = variable; Rel= Relationship; E = Ebalia rotundata; Pm = Persephona mediterranea; Pp= Perse-
phona punctata; Pl = Persephona lichtensteinii. *=<0.05; **=<0.01; Sign. = Significance.
Var. ANOVA Tukey
Rel. E vs Pm E vs Pp E vs Pl Pm vs Pp Pm vs Pl Pp vs Pl
RS p 0.00 P 0.000 0.000 0.000 0.000 0.000 0.994
F 743.2 Sign. s (**) s (**) s(**) s(**) s(**) ns
LS p 0.00 P 0.000 0.000 0.000 0.000 0.000 0.000
F 608.2 Sign. s (**) s (**) s(**) s(**) s(**) s(**)
DS p 0.00 P 0.000 0.000 0.000 0.055 0.000 0.000
F 516.7 Sign. s (**) s (**) s(**) ns s(**) s(**)
DLD p 0.00 P 0.000 0.000 0.000 0.000 0.297 0.040
F 53.2 Sign. s (**) s (**) s(**) s(**) ns s(*)
DRL p 0.00 P 0.080 0.000 0.000 0.000 0.000 0.984
F 36.9 Sign. ns s (**) s(**) s(**) s(**) ns
CL p 0.00 P 0.003 0.000 0.999 0.000 0.000 0.000
F 28 Sign. s (**) s(**) ns s (**) s(**) s(**)
CH p 0.00 P 0.272 0.000 0.000 0.000 0.000 0.000
F 77 Sign. ns s (**) s(**) s(**) s(**) s(**)
CW p 0.00 P 0.705 0.000 0.000 0.000 0.003 0.977
F 13.1 Sign. ns s (**) s(**) s(**) s(**) ns

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 27
TABLE 5. Descriptive statistics for each morphometric variable for the species studied (µm).
sp. Var. E. rotundata P. mediterranea P. lichtensteinii P. punctata
mean sd mean sd mean sd mean sd
RS 558.00 24.04 262.18 21.51 229.78 13.62 228.49 9.29
LS 245.56 4.35 173.18 19.24 89.63 5.08 100.97 7.66
DS 555.57 13.28 361.76 24.30 279.76 9.21 345.16 19.18
DLD 243.96 14.64 352.63 39.20 332.95 22.18 306.71 24.50
DRL 351.61 8.88 323.29 38.07 259.20 31.41 255.15 22.79
CL 448.65 12.12 472.91 24.75 447.94 19.67 414.50 12.64
CH 439.58 14.13 449.33 14.59 413.91 12.03 383.51 12.97
CW 362.57 14.06 355.79 28.92 330.75 13.38 328.12 11.02

sp = species; Var. = Variable; sd = standard deviation.

TABLE 6. Ratios among different morphometric variables for the species studied (µm).
Species Ratios
RS/CW LS/CW DS/CW DS/RS LS/RS DS/LS
E. rotundata 1.54 0.67 1.52 0.98 0.43 2.25
P. mediterranea 0.73 0.48 1.01 1.37 0.66 2.08
P. lichtensteinii 0.69 0.27 0.84 1.21 0.39 3.12
P. punctata 0.69 0.30 1.05 1.51 0.44 3.41

Discussion

The three species studied show the characteristic diagnostic morphology of Leucoisoidea. Rice (1980a) distin-
guished the larvae of Leucosioidea from others by the: (1) reduction in the number of processes added to the telson
in succeeding zoeal stages, (2) reduction or loss of spines on the outer portion of the telson, (3) reduction or loss of
the antenna, (4) reduction in the armature of the antennal exopod and the size of the spinous process, (5) loss of evi-
dent segmentation in the endopod of the maxilla, (6) reduction of the setation of the basis and endopod of the first
maxilliped, (7) reduction of the setation of the basis and endopod of the second maxilliped, and (8) fusion of the
endopodal segments of the second maxilliped.
The zoea 1 of Persephona mediterranea, previously described by Negreiros-Fransozo et al. (1989), is rede-
scribed here. Some differences, mainly regarding the number of aesthetascs on the antennules and the number and
type of setae on the maxillule (endopod with 4 simple setae), maxilla, and second maxilliped (see Table 7) were
found. These differences can be attributed to possible individual variation, or even to a deficiency of optical instru-
ments used in the original study.
Some differences are noted, which facilitate the separation of the four leucosiid species, i.e., the shape of the
carapace and the size and curvature of lateral spines. Whereas the format of the carapace is very similar between P.
mediterranea and P. lichtensteinii, P. punctata has a projection on the posterior region of the carapace, which dis-
tinguishes this species from the others. In P. lichtensteinii, the lateral spines are shorter and curved, differing from
P. mediterranea and from E. rotundata, which has the largest spines among the four species.
Based on a more detailed analysis of the number and types of setae in each structure, the main distinguishing
characteristics of the four species are the number and types of setae of the coxal endite and endopod of the maxil-
lule and maxilla, and the endopod of the first and second maxilliped (see Table 6).
No great differences in morphology appeared when comparing the species studied with the description of Eba-
lia rotundata (as Lithadia rotundata) by Fransozo & Bertini (2002). This morphological similarity suggests an
affinity between the groups. The presence of well-developed spines on the carapace is a plesiomorphic character,
and indicates that these species are less derived among leucosiids.

28 · Zootaxa 3167 © 2012 Magnolia Press HIROSE ET AL.

MORPHOLOGICAL & MORPHOMETRIC COMPARISONS OF PURSE CRABS Zootaxa 3167 © 2012 Magnolia Press · 29
 As mentioned by Rice (1980b), Salman (1982), and Wear & Fielder (1985), Ebalia nux, E. tuberosa, E.
cranchii, and E. laevis show no rostral, lateral, or dorsal spines on the carapace; only E. longipedata has lateral
spines (Aikawa 1937). These features place Ebalia as one of the most derived genera in this group (Ko 2000). This
information reinforces the need to reexamine the taxonomic status of E. rotundata as well as the entire family Leu-
cosiidae, as previously suggested by Ng et al. (2008).
There were significant morphometric differences among the species studied. The size of the dorsal, rostral, and
lateral spines were the most important in separating the species. Ebalia rotundata proved to be quite different from
the other species, and together with Persephona mediterranea formed a distinct group. P. punctata and P. lichten-
steinii are more similar but still have significant morphometric differences. The ratios among morphometric mea-
surements can also contribute to the identification of these purse crabs, and provide an easier and more rapid tool
for this purpose than those previously used.
On the other hand, larval mophometric data should be used with caution, especially when making comparisons
between geographically distant areas, or areas with disparate environmental conditions. Size and relative lengths of
extremities and appendages commonly vary along latitudinal gradients, and variation due to non-genetic alterations
of phenotype may be a result of environmental factors (Shirley et al. 1987). This morphological variability in deca-
pod larvae has been frequently discussed as an ecological strategy and adaptation to changing environmental con-
ditions (Shirley et al. 1987; Anger 2001; Weiss et al. 2010).
The morphological characteristics, combined with the morphometric relationships studied here, may help in
the identification of the planktonic larvae of Leucosioidea from the western South Atlantic, which may be an
important tool in ecological studies. Because of the inadequate knowledge of these larval forms, further studies are
needed to better clarify their taxonomic relationships, which are of fundamental importance for understanding the
phylogeny of this group.

Acknowledgements

We thank the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) (94/4878-4; 98/3136-4; 04/
15194-6), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (133740/2008-0), Coordena-
ção de Aperfeiçoamento de Pessoal de Ensino Superior (CAPES) for financial support. We are grateful to anony-
mous reviewers for their valuable suggestions. The crabs in this study were collected according to Brazilian laws
concerning sampling wild animals.

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