European Journal of Taxonomy 109: 1-15

http://dx.doi.org/10.5852/ejt.2015.109

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomv. eu

2015 • Cumberlidge N. et al.

Research article

urn:lsid:zoobank.org:pub:373F006E-lA73-4DDF-A351-E31B41F8A8E0

New collections of freshwater crabs from northern Madagascar,

with the description of a new species of Foza Reed &

Cumberlidge, 2006 (Brachyura, Potamonautidae),

and comments on their conservation status

Neil CUMBERLIDGE 15 , Sebastian KLAUS 236 ,

Kirstin S. MEYER 47 & Jennifer C. KOPPIN 18

1 Department of Biology, Northern Michigan University, Marquette, Michigan 49855-5301, U.S.A.

5 Email: [email protected]

2 Dept, of Ecology and Evolution, J.W. Goethe-University, Max-von-Laue-Str. 13,

D-60438 Frankfurt am Main, Germany. Email: [email protected]

3 Chengdu Institute of Biology, Chinese Academy of Sciences, No. 9 Section 4, Renmin

Nan Road, Chengdu, 610041 Sichuan, RR. China

4 Oregon Institute of Marine Biology, 63466 Boat Basin Rd, Charleston,

Oregon 97420 USA. Email: [email protected]

5 urn:lsid:zoobank.org:author:05F6365E-D168-4AE3-B511-80FA7E31ACCl

6 urn:lsid:zoobank.org:author:94EDE055-0B67-4986-8905-EF51B0398239

7 urndsid: zoobank. org: author: AFE96B64-F661-44C4-BE21-11BCFAFF 8192

8 um:lsid:zoobank.org:author:B2F0EEB4-4F33-47BA-876E-27D384F95432

Abstract. We report here on recent collections of freshwater crabs from Antsiranana Province, northern

Madagascar. The specimens belong to three species, one of which is new to science and is described

here. This raises the number of species of freshwater crabs found in Madagascar to 17. All are endemic

to the island and all belong to the Afrotropical family Potamonautidae Bott, 1970. The new species,

Foza manonae sp. nov., is compared to the other species in this genus, and an updated key is provided.

It is distinguished from the other three congeners by characters of the male first gonopod, sternum,

carapace, and cheliped. The conservation status of the Malagasy freshwater crab fauna is summarized and

discussed in light of the new material reported on here belonging to two other species, Madagapotamon

humberti Bott, 1965 and Foza ambohitra Cumberlidge & Meyer, 2009.

Key words. Crustacea, taxonomy, Foza , new species, Madagascar, conservation

Cumberlidge N., Klaus S., Meyer K.S. & Koppin J.C. 2015. New collections of freshwater crabs from northern

Madagascar, with the description of a new species of Foza Reed & Cumberlidge, 2006 (Brachyura, Potamonautidae),

and comments on their conservation status. European Journal of Taxonomy 109: 1-15. http://dx.doi.org/10.5852/

eit.2015.109

European Journal of Taxonomy 109: 1-15 (2015)

Introduction

Freshwater crabs are found throughout Madagascar in forested, savanna, and highland habitats, and are

particularly species rich in the northern parts of the island where several biodiversity hotspots have been

identified (Cumberlidge et al. 2004). These crabs live in lakes, streams, and rivers, as well as in adjacent

terrestrial habitats that include rocky crevices and phytotelmata (Cumberlidge et al. 2002, 2005). All

freshwater crabs found in Madagascar are included in the Afrotropical family Potamonautidae Bott,

1970 (Bott 1960, 1965; Ng & Takeda 1994; Cumberlidge 1999, 2014; Cumberlidge et al. 2002, 2007;

Cumberlidge & Sternberg 2002; Reed & Cumberlidge 2006; Cumberlidge & Meyer 2009; Meyer et al.

2014). The high degree of endemism shown by the Malagasy freshwater crabs (100% at the genus and

species levels) is a characteristic that they share with many other freshwater organisms from this long-

isolated tropical island.

We report here on recent collections of freshwater crabs from Antsiranana Province, northern

Madagascar. The specimens belong to two genera: Foza Reed & Cumberlidge, 2006 (that currently

includes three species) and Madagapotamon Bott, 1965, a monotypic genus. Foza was established with

the description of F. raimundi Cumberlidge & Reed, 2006, the type species, from Marojejy in northern

Madagascar and expanded in 2009 by the inclusion of two other species, F. ambohitra Cumberlidge &

Meyer, 2009 and F. goudoti (H. Milne Edwards, 1853). The fourth species of this genus, Foza manonae

sp. nov., described here, is recognized by a combination of morphological characters including those of

the mandible, gonopods, carapace, sternum, and cheliped (Table 1).

The new species is described, figured, and compared with other species in this genus, and an updated

key to the genus is provided. In addition, the conservation status of the Malagasy freshwater crab fauna

is discussed for the 14 species known at the time of the last IUCN Red List assessment (Cumberlidge

et al. 2009). Of these, two species ( Boreathelphusa uglowi (Cumberlidge & Sternberg, 2002) and

Madagapotamon humberti Bott, 1965) are currently listed as ‘vulnerable to extinction’, and five other

species are too poorly known to even assess their conservation status (Table 2).

Material and methods

All measurements were made with digital calipers and are given in mm. The terminology is adapted

from Cumberlidge (1999) and Cumberlidge & Sternberg (2002). Line drawings were prepared using a

Leica MZ 16 stereobinocular microscope. The habitus photographs were taken with a Panasonic Lumix

digital camera. Post processing was done in Adobe Photoshop 7.0. Specimens examined are deposited

in the Zoologische Staatssammlung Miinchen (the Bavarian State Collection of Zoology, ZSM).

The following abbreviations are used in this paper:

a7/a6

IUCN

MNHN

pl-p5

abdominal somite

sutures between abdominal somites

carapace height measured at maximum height of cephalothorax (mm)

carapace length measured along median line from anterior to posterior margin ( mm )

carapace width measured at widest point (mm)

thoracic episternite

front width measured along anterior frontal margin between orbits ( mm )

first gonopod

second gonopod

International Union for the Conservation of Nature

Museum national d’Histoire naturelle, Paris

pereiopods 1-5

CUMBERLIDGE N. et al., New species of Malagasy freshwater crab

s = thoracic sternite

s4/s5 = sternal sulci between adjacent thoracic sternites

s4/e4 = episternal sulci between adjacent thoracic sternites and epistemites

SMF = Senckenberg Museum, Frankfurt, Germany

ZSM = Zoologische Staatssammlung Munchen, Germany

Results

Infraorder Brachyura Latreille, 1802

Superfamily Potamoidea Ortmann, 1896

Family Potamonautidae Bott, 1970

Subfamily Deckeniinae Ortmann, 1897

Fozci Reed & Cumberlidge, 2006

Foza manonae sp. nov.

urn:lsid:zoobank.org:act:B73D6977-4FF3-4C7A-A2ED-8CED73466A54

Figs 1-3, Table 1

Diagnosis

Mid-proximal portion of G1 terminal article widened laterally by rounded lobe; s3/s4 incomplete, faint

in middle deep at sides; anterolateral and posterolateral surfaces of carapace with conspicuous carinae;

subhepatic region of carapace sidewall with carinae; anterior pterygostomial region of carapace sidewall

heavily granulated.

Etymology

The new species is named for the first author’s late wife. Dr. Fouise Manon Bourgault, in recognition of

her numerous contributions to freshwater crab biology during her time spent researching communication

in Africa and Madagascar.

Material examined

Holotype

MADAGASCAR: Adult $, Antsiranana Province, Ankarana Special Reserve, 12.92° S, 49.14° E, near

pitfall trap site, coll. F. Glaw, M. Franzen, J. Kohler & N. d’Cruze, 13 Feb. 2008 (CW 43.7, CF 32.6, CH

18.7, FW 10.5) (ZSM A20145003).

Paratype

MADAGASCAR: Adult $, Antsiranana Province, same locality (CW 42.4, CF 32.2, CH 18.3, FW 10.4)

(ZSM A20145004).

Other material examined

MADAGASCAR: Sub-adult $, Antsiranana Province, Montagne des Frangais Reserve, 12.34° S,

49.35° E, in pitfall trap, coll. N. d’Cruze et al ., 19 Feb. 2008 (CW 36.9, CF 29.0, CH 15.8, FW 9.0)

(ZSM A20145005); juvenile, same locality (CW 12.1, CF 10.4, CH 5.8, FW4.6) (ZSM A20145006).

Description

Based on holotype (adult $, CW 43.7). Carapace outline transversely oval, high (CH/FW 1.75); front

narrow (FW/CW 0.28), deflexed; epibranchial tooth small, pointed, extremely advanced in position;

anterolateral margin evenly curved outward, lined by small granules; postfrontal crest faint, incomplete

postorbital crests, epigastric crests well defined, positioned forward on front; deep wide mid-groove

between epigastric crests; cardiac urogastric grooves deep, cervical grooves deep posteriorly, faint

European Journal of Taxonomy 109: 1-15 (2015)

anteriorly, long, almost reaching postfrontal crest. Anterolateral and posterolateral surfaces of carapace

with conspicuous carinae. Suborbital region of carapace sidewall with small granules, subhepatic region

with carinae, pterygostomial region heavily granulated with setae in inferior part; vertical sulcus on

carapace sidewall curved, granular, running from base of epibranchial tooth to epimeral sulcus. Epistomial

tooth triangular, deflexed, edges smooth. Exopod of third maxilliped long, reaching ischium/merus

junction, flagellum of exopod long, ischium with deep vertical groove. Mandibular palp 2-segmented

Fig. 1 . Foza manonae sp. nov. Holotype, adult CW 43.7 mm, Ankarana Special Reserve, Madagascar

(ZSM A20145003). A. Carapace, frontal view. B. Carapace, dorsal view. C. Third maxilliped, ventral

view. D. Carpus and merus of cheliped, dorsal view. E. Carpus and merus of cheliped, ventral view.

F. Right cheliped, frontal view. G. Left cheliped, frontal view.

CUMBERLIDGE N. et al.. New species of Malagasy freshwater crab

with small, hard, rounded lobe at junction between segments, lobe one-third length of terminal segment.

sl/s2 short, very faint; s2/s3 deep, wide, completely crossing sternum; s3/s4 incomplete, faint in middle

deep at sides. Episternal sulci s4/e4, s5/e5, s6/e6, s7/e7 absent. Male abdomen triangular, tapered, widest

at a3, narrowest at a7 (telson); telson outline forming straight-sided triangle with broad base, rounded

Fig. 2. Foza manonae sp. nov. Holotype, adult CW 43.7 mm, Ankarana Special Reserve, Madagascar

(ZSM A20145003). A. Left and right G1 and G2 shown in situ in ventral view. B. Left Gl, dorsal view.

C. Left Gl, ventral view. D. Sternum and abdomen, ventral view. E. Left mandibular palp, superior

view. F. Left mandibular palp, frontal view. Scale bar: 14 mm (A-B, E), 23.2 mm (C-D).

European Journal of Taxonomy 109: 1-15 (2015)

apex. Sternal grooves s4/s5 meeting telson just beyond abdominal groove between a7/a6; sternal grooves

s5/s6 meeting a6 one half somite length from a6/a5; sternal grooves s6/s7 meeting a5 one third somite

length from a5/a4.

G1 terminal article short (ratio of length of terminal article to subterminal segment 0.27), cone-shaped,

straight, tapering to broad tip, apical opening narrow, directed slightly outward, smooth; mid-proximal

portion of G1 terminal article widened laterally by rounded lobe; lateral, medial folds basally separated,

meeting midway along ventral face, forming longitudinal groove that continues almost to tip of article;

groove not visible on dorsal face. Subterminal segment of G1 with raised rounded shoulder on lateral

margin near junction with terminal article. Terminal article/subterminal segment junction of G1 marked

by faint diagonal line on ventral side, dorsal side marked by broad, subtriangular dorsal membrane;

superior margin of dorsal membrane formed by horizontal basal margin of terminal article, inferior

margin of membrane formed by U-shaped distal edge of subterminal segment; lateral, medial margins

of dorsal membrane narrow. Terminal article of G2 flagellum-like, long, reaching anterior margin of

sternoabdominal cavity; flagellae of G2 viewed together in situ forming heart shape; distal parts of

terminal articles of both G2s touching, sometimes protruding from under closed abdominal telson.

Movable finger (dactylus) of major (right) cheliped slender, upper margin smooth. Fixed finger long,

l A height of palm; lower margin of palm slightly indented; proximal region of cutting edge of fixed

finger with four large, fused molars followed distally by series of small teeth. Carpus with two teeth on

inner margin, first tooth large, pointed, second tooth smaller, pointed, followed by series of very small

teeth. Medial, lateral margins of inferior face of cheliped merus distinctly toothed, inferior face with

pointed, granulated, distal tooth; superior margin and superior face of merus covered with granules

and short carinae; granules on medial margin of merus continuous with granules on medial margin of

cheliped ischium, inferior margin of ischium rounded, smooth. Walking legs (p2-p5) of normal length,

not strikingly elongated, inner margins of propodi of p2 to p5 smooth.

Size

The largest known specimen is the male holotype, CW 43.7. Adults judged by size at pubertal molt are

between CW 37 and CW 42.

Type locality

Ankarana Special Reserve, 12.92° S, 49.14° E, Antsiranana Province, northern Madagascar.

Distribution

Foza manonae sp. nov. is known only from two localities in Antsiranana Province in northern Mada¬

gascar: Ankarana Special Reserve and Montagne des Frangais Reserve (Fig. 4).

Ecology

At the Ankarana Special Reserve the new species is found in sympatry with two other species of

freshwater crabs: Foza ambohitra (ZSM A20145001, ZSM A20145002) and Madagapotamon humberti

(ZSM A20145007, ZSM A20145008, ZSM A20145009, ZSM A20145010). Foza manonae sp. nov. is

also found in sympatry with M. humberti (ZSM A20145012) at Montagne des Frangais Reserve, where

the two species were caught together in pitfall traps. Both F. ambohitra and M. humberti are known to

be semi-terrestrial air-breathing crabs (Cumberlidge & Meyer 2009), so it is likely that F. manonae sp.

nov. also has similar habits and abilities, and also has a similar degree of independence from permanent

water sources. Furthermore, the branchial chambers of A manonae sp. nov. (like those of A ambohitra)

each house two different sets of respiratory organs: a dorsal pseudolung for aerial respiration and ventral

gills for aquatic respiration (Sternberg & Cumberlidge 2001).

CUMBERLIDGE N. el al.. New species of Malagasy freshwater crab

Fig. 3. Foza manonae sp. nov. Holotype, adult S, CW 43.7 mm, Ankarana Special Reserve, Madagascar

(ZSMA20145003). Entire animal. A. Dorsal view. B. Frontal view. C. Sternal view. Scale bar: 14.6 mm

European Journal of Taxonomy 109: 1-15 (2015)

Remarks

The differences between Foza and the other Malagasy freshwater crab genera are discussed by Reed &

Cumberlidge (2006), Cumberlidge & Meyer (2009), and Meyer et al. (2014). Foza manonae sp. nov. is

assigned to the genus Foza on the basis of characters that it shares with F. raimundi Reed & Cumberlidge,

2006, the type species of the genus (Reed & Cumberlidge 2006). These characters include a bilobed

mandibular palp, a faint postfrontal crest, sternal grooves s6/s7 that meet the margin of abdominal

segment a5 in the middle of the segment, and curved, elongated G2s that together form a distinctive

heart shape (Fig. 2A). A preliminary DNA comparison of 16S rRNA sequences of F. manonae sp. nov.

with those of other Malagasy freshwater crab species available from GenBank was made by the second

author (SK, unpublished data). The results positioned the new species in a separate clade, although the

taxonomic sampling and exact identity of the species to which the other sequences belong is currently

the focus of a larger molecular phylogenetic study of the Malagasy freshwater crab fauna using a wider

range of molecular markers (S.R. Daniels, pers. comm.).

The four species of Foza are compared in Table 1 and can be identified as follows: Foza manonae sp.

nov. can be distinguished from F. raimundi Reed & Cumberlidge, 2006 by the texture of the anterolateral

surfaces of the carapace (which have heavy carinae in F. manonae sp. nov. but are smooth, or with only a

few light carinae in F. raimundi ), by the texture of the suborbital and subhepatic regions of the carapace

sidewall (which have granules and/or carinae in F. manonae sp. nov. but are smooth in F. raimundi ), by

the terminal article of G1 (which is cone-shaped in F. manonae sp. nov. and tube-shaped in F. raimundi ),

Fig. 4. Distribution map of northern Madagascar showing the seven localities (black circles) where

freshwater crabs were collected in the present study: Foza manonae sp. nov. (white circles), F. ambohitra

(triangle), and Madagapotamon humberti (squares).

CUMBERLIDGE N. et al.. New species of Malagasy freshwater crab

Table 1. Comparison of the morphological characters of the species in the genus Foza.

Character

F. raimundi

F. goudoti

F. ambohitra

F. manonae sp. nov.

Postfrontal crest

faint

distinct

faint

Anterolateral surface

of carapace

smooth,

light carapace

smooth

smooth,

light carapace

heavy carinae

Postlateral surface

of carapace

light carinae

smooth

heavy carinae

Cervical grooves

short

long

Suborbital region

smooth

large granules

smooth

small granules

Subhepatic region

smooth

carinae

Anterior pterygo-

dense setae

inferior region

inferior region with

stomial region

all over

with setae

setae

Anterior pterygo-

smooth

superior region

heavily granulated

stomial region

with granules

all over

G1 terminal article

tube-shaped,

cone-shaped,

cone-shaped, laterally

not widened

widened by raised lobe

G2 terminal

segment

left one shaped

like question mark,

together form heart

shape

straight, long,

no distal curve

left one shaped

like question mark,

together form heart

shape

left one shaped like

question mark, toge¬

ther fonn heart shape

Sternoabdominal

cavity

dense setae

no/few setae

s3/s4

complete,

shallow in middle,

u-shaped

v-shaped

u-shaped

deep at sides

s6/s7 meets a5

at middle margins

of somite

at a5/a6 junction

at middle margins

of somite

Major cheliped

proximal

propodus

one molar

3-4 molars

by the pterygostomial region and the sternoabdominal cavity (which largely lack setae in F. manonae sp.

nov. but have dense fields of setae in F. raimundi), and by the sternal sulcus s3/s4 (which is incomplete

and only visible at the sides in F. manonae sp. nov., but complete and crossing the entire thoracic

sternum in F. raimundi).

Foza manonae sp. nov. can be distinguished from F. ambohitra by the texture of the posterolateral

corners of the carapace (which have heavy carinae in F. manonae sp. nov., but are smooth with a few

light carinae in F. ambohitra ), by the texture of the suborbital and subhepatic regions of the carapace

sidewall (which have granules and/or carinae in F. manonae sp. nov., but are smooth in F. ambohitra ),

by the suborbital and subhepatic regions of the carapace sidewall (which are granulated in F. manonae

sp. nov., but smooth in F. ambohitra :), and by the terminal article of G1 (which is widened in F. manonae

sp. nov., but slim and evenly tapered in F. ambohitra).

Foza manonae sp. nov. can be distinguished from F. goudoti by the postfrontal crest (which is faint and

incomplete in F. manonae sp. nov., but distinct and completely crosses the carapace in F. goudoti ), by

the anterolateral surfaces of the carapace (which have heavy carinae in F. manonae sp. nov., but are

European Journal of Taxonomy 109: 1-15 (2015)

smooth or with only a few light carinae in F. goudoti), by the subhepatic region of the carapace sidewall

(which has carinae in F. manonae sp. nov., but is smooth in F. goudoti ), and by the terminal articles of

G2 (which are curved distally, forming a distinctive heart shape in F. manonae sp. nov., but are slim,

long, and straight in F. goudoti ).

Foza ambohitra Cumberlidge & Meyer, 2009

Material examined

MADAGASCAR: 2 adult SS, Antsiranana Province, Ankarana Special Reserve, 12.92° S, 49.14° E,

coll. F. Glaw, M. Franzen, J. Kohler & N. d’Cruze, 13 Feb. 2008 (CW 50, Cl 31.6; CW 43, CF 30) (ZSM

A20145001, ZSM A20145002).

Distribution

This species was described by Cumberlidge & Meyer (2009) from specimens collected in Antsiranana

Province, northern Madagascar, at Ambohitra (fonnerly Joffreville) in the Diana Region, and at two

localities in the Analamerana Special Reserve. The present report adds a new locality for this species:

the Ankarana Special Reserve (Fig. 4). In light of the new material and preliminary DNA analysis, it is

possible that the specimen from Toamasina Province, Montagne dAkirindro (NMU PN 17-21.3.2003),

included in F. ambohitra by Cumberlidge & Meyer (2009), may not prove to belong to this species (S.R.

Daniels, pers. comm.).

Remarks

Foza ambohitra is a medium-sized species that lives in the mixed dry deciduous and humid forests of

northern Madagascar. The species can be recognized by its anterolateral margins (granular), its carapace

sidewalls (completely smooth except for a small field of granules at the junction of the longitudinal and

vertical sutures), and by its sternal sulcus s3/s4 (which is complete, U-shaped, and does not meet the

sternoabdominal cavity).

The material reported on here also includes two small juvenile crabs (CW 9.5, CF 7.6 and CW 9.0, CF

7.2) (ZSM A20145014) from the Ankarana Special Reserve that are difficult to identify because their

morphology includes a number of characters that have yet to develop to the adult form. Nevertheless,

these specimens clearly belong to the genus Foza and were collected at the same locality as the specimens

of F. ambohitra (ZSM A20145001, ZSM A20145002). However, we hesitate to assign these juvenile

specimens to F. ambohitra because they possess dense fields of setae on the anterior pterygostomial

region of the carapace sidewall, and because they have very elongated walking legs, both of which are

characters that would place them close to F. raimundi.

Key to the species of Foza

1. G1 terminal article widened by raised lobe; s3/s4 incomplete, faint in middle, deep at sides;

subhepatic region of carapace sidewall with carinae. F. manonae sp. nov.

G1 terminal article slim, not widened; s3/s4 deep, completely crossing thoracic sternum; subhepatic

region of carapace sidewall smooth.2

2. Dense setae in pterygostomial region and in sternoabdominal cavity; cervical grooves short;

proximal region of fixed finger of major cheliped with single large molar; s6/s7 meeting margins

of a5 in middle of somite. F. raimundi Reed & Cumberlidge, 2006

Dense setae lacking in pterygostomial region and in sternoabdominal cavity; cervical grooves

long; proximal region of fixed finger of propodus of major cheliped with 3M large molars; s6/

s7 meeting margins of a5 at a5/a6 junction.3

CUMBERLIDGE N. et al., New species of Malagasy freshwater crab

3. G2 terminal articles straight, long, not hooked over at tips; suborbital region of carapace sidewall

with large granules; postfrontal crest complete, distinct. F. goudoti (EL Milne Edwards, 1853)

G2 terminal articles distinctly curved distally, together forming distinctive heart-shape; suborbital

region of carapace sidewall smooth; postfrontal crest faint, incomplete.

. F. ambohitra Cumberlidge & Meyer, 2009

Madagapotamon humberti Bott, 1965

Madagapotamon humberti Bott, 1965: 344-346, fig. 7, pi. 4, figs 14-17. —Vuillemin 1970: 245-265.

— Ng & Takeda 1994: 162-163, table 1. — Cumberlidge & Sternberg 2002: figs 2e, 3g, 41, 5h, 6g, 7k-l,

8g, lOg-h, llh. —Ng, Guinot & Davie 2008: 165.

Material examined

MADAGASCAR: All from Antsiranana Province. - Ank arana Special Reserve, 12.92° S, 49.14° E, in

pitfall trap, all coll. F. Glaw, M. Franzen, J. Kohler & N. d’Cruze, 12 Feb. 2008: damaged, CW 21.5 (ZSM

A20145007); damaged, CW 34.6 (ZSM A20145008); damaged, legs only (ZSM A20145009); damaged,

CW 18.2 (ZSM A20145010). - Ankarana Special Reserve, Encampment des Anglais (Anilotra), 7.5 km

NW of Mahamajina, 12.91° S, 49.11° E, mostly undisturbed tsingy forest, on cliff face, 125 m asl, coll.

S.M. Goodman, 8 Apr. 2002, 1 specimen (FMNH 7577). -Baie de Sakalava, Kohler-Stelle, 12.28° S,

49.38° E, coll. F. Glaw, M. Franzen, J. Kohler & N. d’Cruze, 17 Feb. 2008, 2 adult $$ (CW 34.3, Cl

28.9, CH 13.1, FW 6.6; CW 28.9, CL 21.0) + 2 juvenile 33 (CW 21.1; CW 19.8) (ZSM A20145011).

-Passe d’OrangeaReserve, 12.23° S, 49.37° E, pitfall, coll. S. Megson, 22 Feb. 2008, subadult 3 (CW

28.8, CL 21.9); damaged subadult $ (CW 31.6, CL 21.5); damaged specimen (ZSM A20145013). -

Montagne des Fram^ais Reserve, 12.34° S, 49.35° E, in pitfall trap, coll. N. d’Cruze et al. , 19 Feb. 2008,

1 specimen, damaged (ZSM A20145012). - Analamerana Special Reserve, 8.6 km SE of Menagisy,

Foret d’Analabe, along Bobakindro River, 12.80° S, 49.37° E, partially disturbed dry deciduous forest,

on ground during day near pitfall trap number 2, 40 m asl, all coll. S.M. Goodman, 17-21 Jan. 2004: 1

specimen (FMNH 11033); adult 3 (CW 38.6) + adult $ (CW 32.6) (FMNH 11034); 1 specimen (FMNH

11035); adult 3 (CW 35.4) + adult ? (CW 27.6) (FMNH 11036); adult $ (CW 33) (FMNH 11037).

-Analamerana Special Reserve, 8.6 km SE of Menagisy, Foret d’Analabe, along Bobakindro River,

12.75° S, 49.49° E, partially disturbed dry deciduous forest, found climbing up a tree during middle

afternoon, 40 m asl, coll. S.M. Goodman, 18 Jan. 2004, 1 specimen (FMNH 11035). - Analamerana

Special Reserve, Foret d’Ankavanana, 15.8 km SE of Anivorano-Nord, 12.8° S, 49.37° E, partially

disturbed mixed dry deciduous and humid forest, found during early morning check, 200 m asl, all coll.

S.M. Goodman, 23-27 Jan. 2004: adult 3 (CW 29.6) (FMNH 11044a); adult 3 (CW 44.3) (FMNH

11047); 1 specimen (FMNH 11048); 1 specimen (FMNH 11053); adult 3 (CW 42.7) (FMNH 11055);

subadult $ (CW 29.1) (FMNH 11057).

Distribution

This species was described by Bott (1965) from specimens collected in dry deciduous forests in the

Ankarana Special Reserve in Antsiranana Province in northern Madagascar. The present work adds

further material from this locality, plus several more specimens from two other localities in the

Analamerana Special Reserve in Antsiranana Province (Fig. 4). We are also able to confirm the continued

presence of M. humberti in Montagne des Frant^ais Special Reserve near the coastal city of Antsiranana

(formerly Diego-Suarez), adding to the previous record which was based on specimens collected in

1899 (MNHN-B 5034) (Cumberlidge & Sternberg 2002). This species has also been reported to occur

on the island of Nosy Be in Antsiranana Province (ZSM 1163/1; SMF 4487) (Cumberlidge & Sternberg

2002). Two new localities for M. humberti are reported here: Baie de Sakalava (Kohler-Stelle, inland

locality) and the Passe de Orangea Reserve, both of which are relatively close to the city of Antsiranana

in Antsiranana Province.

European Journal of Taxonomy 109: 1-15 (2015)

Remarks

Madagapotamon humberti is one of the most distinctive species of all of the Madagascan freshwater

crabs and this colorful, long-legged, rock-crevice and cave-dwelling species stands apart from all others

on the island. M. humberti can easily be distinguished by the absence of a flagellum on the exopod of

the third maxilliped, by the mandibular palp with a simple terminal segment, and by the egg-shaped

outline of the adult male abdomen. Living specimens from the Anlcarana Special Reserve have a yellow

carapace, pink legs, and chelipeds with white fingers, while those from Montagne des Fran 9 ais Reserve

have an all-white carapace, along with pink legs and chelipeds with white fingers.

Discussion

Conservation status of Malagasy freshwater crabs

Table 2 provides a list of the 17 species of Malagasy freshwater crabs (in eight genera), with

the conservation status of the 14 species that were assessed using the IUCN Red List protocols by

Cumberlidge et al. (2009). One half of the Malagasy freshwater crab fauna (seven out of 14 species)

were assessed as Least Concern (LC), including two species of Foza , four species of Hydrothelphusa

A. Milne-Edwards, 1872 and one species of Malagasya Cumberlidge & Sternberg, 2002 (Table 2).

Two species ( Boreathelphusa uglowi (Cumberlidge & Sternberg, 2002) and Madagapotamon humberti

Bott, 1965) were found to be vulnerable to extinction (VU) and another five (Malagasya goodmani

Cumberlidge, Boyko & Harvey, 2002; Marojejy longimerus (Cumberlidge, Boyko & Harvey, 2002);

Skelosophusa gollhardi (Bott, 1965); S. prolixa Ng & Takeda, 1994; S. eumeces Ng & Takeda, 1994)

were Data Deficient (DD) - i.e., they were too poorly known to even carry out a conservation assessment

(Cumberlidge 2008a, 2008b).

The five Malagasy DD species of freshwater crabs (35% of the fauna) introduce an element of uncertainty

into the conservation planning process because their conservation status (when assessed) may either

increase or decrease the number of threatened species (VU, EN, or CR), or it may not change current

estimates if all of these species prove to be LC. However, it is likely that most of the DD species will

be assessed as belonging to a threatened category because many of the DD species are single-locality

endemics with a very narrow distributional range, a profile typical of many of the species currently in

threatened categories worldwide (Cumberlidge et al. 2009). For example, Sri Lanka is one of the few

countries where the conservation status of the entire freshwater crab fauna has been assessed, and its

faunal list therefore includes no DD species (Bahir et al. 2005; Cumberlidge et al. 2009). These studies

show that Sri Lanka has the highest number of threatened species of freshwater crabs of any country

in the world and that many of these species are single-locality endemics (Cumberlidge et al. 2009).

Clearly, the VU and DD species of endemic Madagascan freshwater crabs present obvious foci for

future ecological fieldwork, biotic inventories, and conservation prioritization activities.

The specimens reported on here (Fig. 4) were collected within a number of protected areas in the

generally dry Antsiranana Province, all of which feature deciduous dry forest and limestone massifs

with karst formations (tsingy). These protected areas are Ankarana Special Reserve (F. manonae sp.

nov., F. ambohitra , and M. humberti), Montagne des Frangais Reserve (F. manonae sp. nov. and M.

humberti ), the Orangea Reserve (M. humberti ), and Analamerana Special Reserve (M. humberti).

The conservation status of F. ambohitra , F. manonae sp. nov., and one species of Glabrithelphusa

Meyer et al ., 2014 has not yet been assessed because these taxa were described since the IUCN global

conservation assessment was carried out (Cumberlidge et al. 2009). The specimens ofM humberti (VU)

reported here from near the Baie de Salcalava and from the Orangea Reserve are new locality records

for this species, and the specimens from the Montagne des Fran 9 ais Reserve confirm its continued

presence after more than 110 years. These new data may well influence the current conservation status of

CUMBERLIDGE N. et al., New species of Malagasy freshwater crab

Table 2. Summary of the freshwater crabs found in Madagascar, with their conservation status assessed

using the IUCN Red List protocols (based on data from Cumberlidge et al. 2009). Non-threatened

categories: LC = Least Concern, NT = Near Threatened. Threatened categories: VU = Vulnerable, EN

= Endangered, CR = Critically Endangered. Other categories: DD = Data Deficient. * = not assessed.

Species

LC NT

EN CR DD

Boreathelphusa uglowi

Foza raimundi

Loza goudoti

Foza ambohitra *

Foza manonae sp. nov. *

Glabrithelphusa angene *

Hydrothelphusa agilis

Hydrothelphusa madagascariensis

Hydrothelphusa bombetokensis

Hydrothelphusa vencesi

Madagapotamon humberti

Malagasya antongilensis

Malagasya goodmani

Marojejy longimerus

Skelosophusa gollhardi

Skelosophusa eumeces

Skelosophusa prolixa

Species per Red List category

7 0

0 0 5

M. humberti (VU, Cumberlidge 2008a). The presence of F. ambohitra in the Ankarana Special Reserve

is a new locality record for this species (Cumberlidge & Meyer 2009), and will be useful when the

conservation status of this species is eventually assessed.

Acknowledgements

We thank Dr Frank Glaw and his colleagues for collecting some of the specimens included here that were

passed on to the second author (SK) for further study. The first author (NC) prepared the photographs,

the second author (SK) prepared the maps, and the fourth author (JCK) prepared Figs 1 and 2. We also

thank Dr Steven M. Goodman of the Field Museum, Chicago, for some of the specimens included here,

and we are grateful to Marty Prydzia, Joachim Gerber, and Dr Janet Voight of the Field Museum for

kindly lending some of the specimens used in this study. SK was supported by a Fellowship for Young

International Scientists of the Chinese Academy of Sciences (no. 2013Y1SA0006).

References

BahirM.M.,NgP.K.L., Crandall K. & Pethiyagoda R. 2005. A conservation assessment of the freshwater

crabs of Sri Lanka. The Raffles Bulletin of Zoology, Suppl. 12: 121-126.

Bott R. 1960. Crustacea (Decapoda): Potamonidae. In: Hanstrom B., Brinck P. & Ruderbeck G. (eds)

South African Animal Life: Results of the Lund University Expedition in 1950-1951, vol. 7: 13-18.

Almqvist & Wiksells, Uppsala.

European Journal of Taxonomy 109: 1-15 (2015)

Bott R. 1965. Die SiiBwasserkrabben von Madagaskar. Bulletin du Museum national d ’Histoire naturelle

37 (2): 335-350.

Cumberlidge N. 1999. The Freshwater Crabs of West Africa, Family Potamonautidae. Faune et Flore

Tropicales 35, IRD, Paris.

Cumberlidge N. 2008a. Madagapotamon humberti. In: IUCN 2013. IUCN Red List of Threatened

Species. Version 2013.2. www.iucmedlist.org . Downloaded 13 May 2014.

Cumberlidge N. 2008b. Boreas uglowi. In: IUCN 2013. IUCN Red List of Threatened Species. Version

2013.2. www.iucmedlist.org . Downloaded 13 May 2014.

Cumberlidge N. 2014. An overview of the Afrotropical freshwater crab fauna: Diversity, biogeography,

and conservation (Brachyura, Potamoidea, Potamonautidae and Potamidae). In: Yeo D., Cumberlidge

N. & Klaus S. (eds.) Crustaceana Monographs: Advances in Freshwater Decapod Systematics and

Biology: 71-99. Koninklijke Brill, The Netherlands, http://dx.doi.org/10.1163/9789004207615_006

Cumberlidge N. & Meyer K.S. 2009. A new species of Foza Reed & Cumberlidge, 2006, from northern

Madagascar (Decapoda, Brachyura, Potamoidea, Potamonautidae), with a redescription of F. goudoti (H.

Milne Edwards, 1853) comb, n., and comments on Skelosophusaprolixa Ng & Talceda, 1994. ZooKeys

18: 77-89. http://dx.doi.org/10.3897/zookeys. 18.102

Cumberlidge N. & Sternberg R. 2002. The freshwater crabs of Madagascar (Crustacea, Decapoda,

Potamoidea). Zoosystema 24: 41-79.

Cumberlidge N., Boyko C.B. & Harvey A. W. 2002. Anew genus and species of freshwater crab (Deca¬

poda, Crustacea, Potamoidea) from northern Madagascar, and a second new species associated with

Pandanusleaf axils. Journal ofNatural History 3 6:65-77. http://dx.doi.org/10.1080/00222930010003800

Cumberlidge N., Reed S.K. & Boyko C.B. 2004. Distribution patterns of the Malagasy freshwater

crabs (Crustacea: Decapoda: Brachyura). Journal ofNatural History 38: 1133-1157. http://dx.doi.

org /10,1080/0022293 031000068419

Cumberlidge N., Fenolio D.B., Walvoord M.E. & Stout J. 2005. Tree-climbing crabs (Potamonautidae

and Sesarmidae) from phytotelmic microhabitats in rainforest canopy in Madagascar. Journal of

Crustacean Biology 25: 302-308. http://dx.doi.org/10.1651/C-2532

Cumberlidge N., Marijnissen S.A.E. & Thompson J. 2007. Hydrothelphusa vencesi , a new species of

freshwater crab (Brachyura: Potamoidea: Potamonautidae) from southeastern Madagascar. Zootaxa

1524: 61-68.

Cumberlidge N., Ng P.K.L., Yeo D.C.J., Magalhaes C., Campos M.R., Alvarez F., Naruse T., Daniels

S.R.,EsserL.J.,AttipoeF.Y.K., Clotilde-BaF.-L.,DarwallW.R.T.,McIvorA.,RamM. &CollenB. 2009.

Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges.

Biological Conservation 142: 1665-1673. http://dx.doi.Org/10.1016/j.biocon.2009.02.038

Meyer K.S., Cumberlidge N. & Koppin J.C. 2014. A new genus and species of freshwater crab from

Madagascar (Decapoda, Brachyura, Potamoidea, Potamonautidae). Zootaxa 3884: 65-72. http://dx.doi.

org/10.11646/zootaxa.3884.1.5

Ng P.K.L. & Takeda M. 1994. Skelosophusa (Crustacea, Decapoda, Brachyura), a new genus of

potamonautid freshwater crab from Madagascar, with descriptions of two new species. Bulletin of the

National Science Museum, Series A (Zoology) 20: 161-172.

Ng P.K.L., Guinot D. & Davie P. 2008. Systema Brachyuorum: Part I. An annotated checklist of extant

brachyuran crabs of the world. The Raffles Bulletin of Zoology, Suppl. 17: 1-286.

CUMBERLIDGE N. et al.. New species of Malagasy freshwater crab

Reed S.K. & Cumberlidge N. 2006. Foza raimundi , anew genus and species ofpotamonautid freshwater

crab (Crustacea: Decapoda: Potamoidea) from northern Madagascar. Proceedings of the Biological

Society of Washington 119: 58-66. http://dx.doi.org/10.2988/0006-324Xt20Q6) 119%5B58:FRANGA%

5D2,O.CO:2

Sternberg R. & Cumberlidge N. 2001. Notes on the position of the true freshwater crabs within the

brachyrhynchan Eubrachyura (Crustacea: Decapoda: Brachyura). Hydrobiologia 449: 21-39. http://

dx.doi.org/10.1023/A: 1017560113027

Vuillemin S. 1970. Observations ecologiques et biologiques sur Madagapotamon humberti Bott, 1965

(Crustace, Decapode, Potamonide) du massif de l’Ankarana (Madagascar). Annales de I’Universite de

Madagascar 7: 245-266.

Manuscript received: 26 August 2014

Manuscript accepted: 4 November 2014

Published on: 15 January 2015

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; National History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, De nm ark