Ng: New species of Sundathelphusa from southeastern Borneo
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 63: 494–502

Date of publication: 23 October 2015
http://zoobank.org/urn:lsid:zoobank.org:pub:37B01671-481A-44FA-9324-AF7E72021F5E

On a new species of Sundathelphusa Bott, 1969 (Crustacea: Brachyura:
Gecarcinucidae) from southeastern BorneoPeter K. L. Ng
Abstract. A new species of gecarcinucid freshwater crab of the genus Sundathelphusa Bott, 1969, is describedfrom southern Kalimantan in Indonesian Borneo. Although the genus is species-rich, most of the taxa are fromthe Philippines, Moluccas and Sulawesi, and only two species, S. tenebrosa Holthuis, 1979, and S. aspera Ng& Stuebing, 1989, are known from Borneo. Sundathelphusa brachyphallus, new species, differs from these two
species mainly by the distinctive structure of the male first gonopod. It also differs from S. tenebrosa by the formof the carapace, notably in the structure of the frontal median triangle.
Key words. Taxonomy, freshwater crab, Sundathelphusa, new species, Borneo, Gecarcinucidae

INTRODUCTIONTAXONOMY

The genus Sundathelphusa Bott, 1969 (type species Potamon(Geothelphusa) cassiope De Man, 1902) currently contains
33 species (Ng et al., 2008; Schubart & Ng, 2008; Mendoza& Naruse, 2010; Husana et al., 2009, 2014, 2015; Ng, 2010).
Most species occur in the Philippines, Sulawesi and Lesser
Sunda Islands, with only two species known from the islandof Borneo: S. tenebrosa Holthuis, 1979 (from northern
Sarawak and Brunei) and S. aspera Ng & Stuebing, 1989
(from northwestern Sabah). In this paper, a third species
from southeastern Borneo is described.

Family Gecarcinucidae Rathbun, 1904
Sundathelphusa Bott, 1969
Sundathelphusa brachyphallus, new species(Figs. 1, 2, 7A–E)
Material examined. Holotype: male (27.1 × 22.0 mm) (MZB
Cru 4396), Liang Hintan, Hulu Sungai Barito, Kalimantan
Tengah, Borneo, Indonesia, coll. C. Rahmadi, 16 June 2004.
Paratype: 1 male (29.3 × 23.4 mm, chelipeds missing) (ZRC
2015.0460), same data as holotype.

Specimens examined are deposited in the Muzium Zoologicum
Bogorense (MZB) in Cibinong, Java, Indonesia; The Natural
History Museum (NHM), London, United Kingdom; TheNaturalis Biodiversity Centre (previously the Rijksmuseum
van Naturrlijke Historie, RMNH), Leiden, The Netherlands;
and the Zoological Reference Collection (ZRC) of the Lee
Kong Chian Natural History Museum (previously Raffles
Museum of Biodiversity Research), National University ofSingapore. Measurements provided, in millimetres, are of
the carapace width and length, respectively. The terminology
follows that used in Ng (1988); with the G1 and G2 used
for the male first and second gonopods, respectively.

Comparative material. Sundathelphusa tenebrosa Holthuis,
1979: holotype male (23.5 × 19.1 mm) (NHM 1979.135),
Camp 2, in 2 × 2 m pool, 10 cm deep, clear cold mountainstream with pebble and gravel bottom, hill dipterocarp forest,
4°03’N 114°53’E, 500 m asl, Gunung Mulu National Park,
Sarawak, Malaysia, Borneo, coll. J. Dring, 18 March 1978;
paratype male (21.9 × 17.8 mm) (NHM 1979.136), samedata as holotype; 1 paratype female (RMNH D31972), Deer
Cave, Gunung Mulu National stream near southwest entrance,
under rock, Sarawak, Malaysia, Borneo, coll. P. Chapman,
10 March 1978; 1 paratype female (RMNH D31979), Deer
Water Cave, Gunung Mulu National Park in stream about
100 m from entrance, Sarawak, Malaysia, Borneo, coll.
P. Chapman, 25 April 1978; 2 paratype females (RMNH
D31972), Clearwater Cave, Gunung Mulu National Park in
main stream passage near river junction, coll. P. Chapman,
13 April 1978; 1 female (30.8 × 24.6 mm) (ZRC 1989.3420),
Sungai Melinau, Merimau Gorge, Gunung Mulu National
Park, Sarawak, Malaysia, Borneo, coll. R.P. Lim & G. Liew,
16 April 1978; 1 juvenile (ZRC 1989.3682), Sungai Melinau,
Merimau Gorge, Gunung Mulu National Park, Sarawak,
Malaysia, Borneo, coll. R.P. Lim, 17 April 1978; 1 male
(16.1 × 13.0 mm) (ZRC 2013.1293), Mulu Summit trail at

Lee Kong Chian Natural History Museum, Faculty of Science, National University ofSingapore, 2 Conservatory Drive, Singapore 117377, Republic of Singapore; Email:peterng@nus.edu.sg© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Fig. 1. Sundathelphusa brachyphallus, new species. A–D, holotype male (27.1 × 22.0 mm) (MZB Cru 4396); E, paratype male (29.3
× 23.4 mm) (ZRC 2015.0460). Kalimantan Tengah, Indonesia. A, overall habitus; B, E, dorsal view of carapace; C, frontal view of
cephalothorax; D, subfrontal view of cephalothorax.

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Ng: New species of Sundathelphusa from southeastern Borneo

Fig. 2. Sundathelphusa brachyphallus, new species, holotype male (27.1 × 22.0 mm) (MZB Cru 4396); Kalimantan Tengah, Indonesia. A,
left third maxilliped; B, anterior thoracic sternum and abdomen; C, sternoabdominal cavity and abdominal locking tubercles on sternite 5;
D, male abdomen; E, frontal view showing frontal median triangle, orbits, antennules, antennae and epistome; F, outer view of left chela;
G–J, outer views of right first to fourth ambulatory legs, respectively.

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Fig. 3. Sundathelphusa tenebrosa Holthuis, 1979. A–D, holotype male (23.5 × 19.1 mm) (NHM 1979.135), Gunung Mulu National Park,
Sarawak; E, F, paratype male (21.9 × 17.8 mm) (NHM 1979.136), Gunung Mulu National Park, Sarawak; G–I, female (30.8 × 24.6 mm)(ZRC 1989.3420), Gunung Mulu National Park, Sarawak. A, G, overall habitus; B, H, frontal view of cephalothorax; C, E, dorsal viewof carapace; D, F, male abdomen; I, subfrontal view of cephalothorax.

Camp 2, Gunung Mulu National Park, Sarawak, Malaysia,
Borneo, coll. A.D. Tranh, 4 November 2006. Sundathelphusa
aspera Ng & Stuebing, 1989: holotype male (42.6 × 32.8mm) (ZRC 1989.2143), clear, swift forest stream, Sungei
Rabergan, tributary of Sungei Muaya, near Mendalong,
4°57’N 115°43’E, 750 m above sea level, Sipitang District,
Sabah, Malaysia, Borneo; 1 male (32.3 × 25.4 mm), 1 female(44.1 × 33.1 mm) (ZRC 2014.845), Temburong, Amo, Batu
Apoi Forest Reserve, Ulu Temburong National Park, Brunei,
Borneo, coll. Jangorun anak Eri, 7 August 2014; 1 female(44.5 × 33.7 mm) (ZRC 2013.746), Temburong District,
Brunei, Borneo, coll. S. Salam, 6 July 2012; 2 females (29.3
× 23.0 mm, 19.7 × 15.6 mm) (ZRC 2012.1225), in primary
forest stream, in area near Sarawak border, Brunei, coll. H.
Mohammod, October, 2012.

margins (Figs. 1C, D, 2E). External orbital angle broadly
triangular, margin granulated, anterolateral margin separatedby small but distinct notch, epibranchial tooth small
but visible (Fig. 1A, B, E). Male abdominal somite 6
subrectangular, slightly longer than broad (Fig. 2B, D). G1
relatively short, stout; terminal segment 0.25 times total
length, subcylindrical, proximal part swollen, outer margin
sinuous, surfaces covered with numerous setae, tip withwide opening (Fig. 7A–D).
Description. Carapace transversely subovate, wider thanlong; regions distinct, separated by distinct grooves; dorsal
surface gently convex, glabrous, pitted or rugose, especiallyalong margins and cristae; frontal region with low granules,
orbital regions finely pitted; cervical groove deep, broad,extending from lateral edge of postorbital cristae to deepH-shaped gastric groove; branchial regions gently inflated;postorbital cristae relatively sharp, distinct, gradually
becoming lower, less distinct towards cervical groove, with

Diagnosis. Carapace with dorsal surfaces rugose, cervicalgrooves deep, broad (Fig. 1A–C). Frontal median triangle
distinct, dorsal margin cristate, contiguous with lateral497

Ng: New species of Sundathelphusa from southeastern Borneo

Fig. 4. Sundathelphusa aspera Ng & Stuebing, 1989, overall habitus. A, holotype male (42.6 × 32.8 mm) (ZRC 1989.2143), Sabah; B,
male (32.3 × 25.4 mm) (ZRC 2014.845), Temburong, Brunei; C, female (44.1 × 33.1 mm) (ZRC 2014.845), Temburong, Brunei; D,
female (44.5 × 33.7 mm) (ZRC 2013.746), Temburong, Brunei.

separate short, low crista just before anterolateral margin;
postorbital crista separated from epigastric crista by shallow
depression; epigastric cristae distinct, not sharp, prominently
rugose, margins sloping anteriorly, separated mediallyby deep longitudinal groove which extends to just before
mesogastric area (Fig. 1A, B, E). Frontal margin slightly
sinuous to almost straight, slightly deflexed, frontal median
triangle broad, distinct, dorsal crista distinct, completely
merging with lateral parts; inner edge of supraorbital margin,
next to outer edge of frontal margin, with low crista adjacent,parallel to it (Figs. 1, 2E). External orbital angle low, broadly
triangular, outer margin length ca. 4 times longer than inner
margin, outer margin almost straight, lined with small
granules, curving to meet anterolateral margin, forming small
triangular notch; epibranchial tooth low but just visible evenwhen eroded; anterolateral margin gently convex, lined withsmall granules, adjacent surfaces covered with numerous
rows of strong oblique striae; not clearly demarcated fromposterolateral margin; posterolateral margin gently concave,
adjacent surfaces lined with oblique striae, converging toalmost straight or gently posterior carapace margin (Fig.
1A, B, E). Pterygostomial, suborbital, sub-hepatic and subbranchial regions distinctly rugose or lined with obliquegranules (Figs. 1C, D, 2E). Posterior margin of epistome
with triangular median lobe, tip rounded; lateral margin withbroadly triangular lobe, separated by deep fissures (Figs. 1C,
D, 2E). Third maxilliped quadrate; ischium subrectangular,
distinctly longer than broad, sulcus deep, oblique, closer
to inner margin; merus squarish, lateral and distal margins
cristate, medially depressed; exopod slender, reaching tohalf length of merus, with distinct triangular projection on

Fig. 5. Sundathelphusa aspera Ng & Stuebing, 1989, frontal andsubfrontal views showing frontal median triangle. A, holotype male
(42.6 × 32.8 mm) (ZRC 1989.2143), Sabah; B, C, male (32.3 ×
25.4 mm) (ZRC 2014.845), Temburong, Brunei; D, E, female (29.3
× 23.0 mm) (ZRC 2012.1225), Brunei.

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Fig. 6. Sundathelphusa aspera Ng & Stuebing, 1989. A, B, holotype male (42.6 × 32.8 mm) (ZRC 1989.2143), Sabah; C, male (32.3 ×
25.4 mm) (ZRC 2014.845), Temburong, Brunei. A, anterior thoracic sternum and male abdomen; B, C, male abdomen.

inner margin of distal part, flagellum long, extending acrosswidth of merus (Fig. 2A). Terminal segment of mandibular
palp bilobed, inner lobe twice length of outer lobe, flap-like.

5 trapezoidal with gently concave to straight lateral margins;somite 6 subrectangular, slightly longer than broad, proximal
margin slightly shorter than distal margin; telson triangularwith concave lateral margins and rounded tip (Fig. 2B, D).

Chelipeds subequal; outer surfaces of articles rugose (Fig.
1A). Basis-ischium fused, with low tooth on anterior margin(Fig. 2B). Merus trigonal in cross-section, margins uneven,covered with striae and low granules; anterior margin withlow subterminal tooth. Upper surfaces of carpus with broad,shallow longitudinal depression, inner angle with prominent
sharp spine, base with sharp tubercle (Fig. 1A). Fingers
of chela subequal to length of palm; cutting edges withnumerous sharp teeth and denticles, tips of fingers strongly
curved (Fig. 2F).

G1 relatively short, stout, directed obliquely outwards;
subterminal segment tapering gradually from broad base,sinuous on proximal part of outer margin, rest of margin
gently concave; terminal segment demarcated from
subterminal segment but grooves shallow, 0.25 times lengthof G1, subcylindrical, proximal part swollen, outer margin
sinuous, surfaces covered with numerous setae, tip withwide opening, groove for G2 marginal or slightly dorsal in
position, dorsal and ventral folds slightly overlapping (Fig.
7A–D). G2 slightly longer than Gl, flagellum well developed,
about half length of basal segment (Fig. 7E).

Ambulatory legs not elongated, third pair longest; outersurfaces gently rugose (Figs. 1A, 2G–J). Merus stout, anterior
margin slightly crested, uneven appears weakly serrated,dorsal subterminal angle just visible but without spine or
tooth; propodus with 2 rows of downward pointing spines onposterior margin; dactylus almost straight or gently curved,
margins with numerous well developed spines pointing
obliquely distally (Fig. 2G–J).

Etymology. The name alludes to the diagnostic G1 of thespecies, which is relatively short and stout. The name isused as a noun in apposition.
Remarks. Ng & Stuebing (1989: 17) discussed the systematic
status of the two Sundathelphusa species from Borneo,
commenting that they are distinct among congeners in having
carapaces which are not strongly inflated, with the dorsal
surfaces prominently rugose, possession of prominent andrugose epigastric and postorbital cristae, a frontal median
triangle which is more developed, with the dorsal crista
distinct; and adult male chelipeds which are symmetrical(Figs. 1, 2E, 3A–C, E, G, H, I, 4, 5; Holthuis, 1979: pl.
7; Ng & Stuebing, 1989: fig. 1A). In almost all other nonBornean Sundathelphusa species, the carapaces are generally
higher and/or more inflated, with the surfaces generallysmoother; the epigastric and postorbital cristae are lessprominent; the frontal median triangle is incomplete, withthe dorsal crista distinct, not very sharp and separated fromthe lateral margins; and the adult male chelipeds are often
distinctly asymmetrical. Ng & Stuebing (1989) believedthat S. minahassae (Schenkel, 1902) (from Sulawesi)

Surface of thoracic sternum smooth to punctate; sternites 1
and 2 completely fused, forming triangular plate; separated
from sternite 3 by gently concave suture; sternites 3 and 4completely fused except for lateral clefts; sternoabdominal
cavity almost reaching imaginary line connecting anterioredges of coxae of chelipeds (Fig. 2B, C). Sternoabdominal
cavity deep; abdominal locking mechanism on sternum
composed of rounded tubercle on submedian part of sternite
5 (Fig. 2C).
Abdomen distinctly T-shaped; all somites and telson free;somite 1 longitudinally narrow, extending across width ofposterior carapace margin, reaching bases of coxae of lastambulatory legs; somite 2 as wide as somite 1; somite 3
subrectangular, with convex lateral margins; somites 4 and499

Ng: New species of Sundathelphusa from southeastern Borneo

Fig. 7. A–E, Sundathelphusa brachyphallus, new species, holotype male (27.1 × 22.0 mm) (MZB Cru 4396); Kalimantan Tengah,Indonesia; F–H, Sundathelphusa tenebrosa Holthuis, 1979, holotype male (23.5 × 19.1 mm) (NHM 1979.135), Gunung Mulu National
Park, Sarawak; I–K, Sundathelphusa aspera Ng & Stuebing, 1989, male (32.3 × 25.4 mm) (ZRC 2014.845), Temburong, Brunei. A–D,
I–K, left G1; E, left G2; F–H, right G1 (laterally inverted to allow for better comparisons; left G1 missing). A, F, I, left G1 (ventralview); B, left G1 (dorsal view); C, G, J, distal part of left G1 (ventral view); D, H, K, distal part of left G1 (dorsal view); E, left G2.
Scale bars: A, B, E = 1.0 mm; C, D = 0.5 mm.

and S. halmaherensis (De Man, 1902) (from Halmahera)
also belonged to the group on the basis of the externalfeatures. However, external carapace features aside, their
G1 structures are quite different. The two Bornean species
are unique among Sundathelphusa species in having the G1
terminal segment relatively cylindrical, with the tip broad
and the opening relatively wide; and the surfaces covered
with numerous setae (Fig. 7F–K; Holthuis, 1979: fig. 8b,c; Ng & Stuebing, 1989: fig. 2B–E; Ng, 2004: fig. 11E). In
all other Sundathelphusa species, including S. minahassae

and S. halmaherensis, the G1 terminal segment is conicaland tapering to a sharp tip (e.g., see Chia & Ng, 2006:
figs. 39–44). In S. brachyphallus, the G1 terminal segment
is actually subcylindrical as the distal part does taper andthe tip is proportionately less broad compared to its two
Bornean congeners (Fig. 7A–D). However, the G1 terminal
segment is still relatively stouter than other Sundathelphusa
species, with the opening at the tip wider and the surfaces
are covered with numerous long setae (Fig. 7A–D), as in
the other two Bornean Sundathelphusa species (Fig. 7F–K).
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In a molecular phylogenetic study of the family by Klaus et al.
(2009), Sundathelphusa was shown to be polyphyletic, withits members in several clades. Sundathelphusa minahassae
came out with a clade from Java and Peninsular Malaysia,
and possibly including Sulawesi (Klaus et al., 2009: 520, 523)
while S. halmaherensis was in a clade primarily comprising
species from the Philippines and Moluccas (Klaus et al., 2009:
520, 522). Significantly, the only Bornean species tested, S.
tenebrosa, came out in a monophyletic clade by itself, farfrom other congeners. The Bornean species will need to be
removed to their own genus at a later date. Certainly, thedistinctive form of their G1 structures easily distinguishes
them from all other Sundathelphusa species.

notch (Fig. 3A, C, E, G ). The rugosity of the carapaces
differs to some degree. The holotype of S. aspera has the
dorso-lateral surfaces relatively less rugose (Fig. 4A) and isslightly weaker in smaller adult specimens of S. tenebrosa(Fig. 3A, C). Other specimens of S. aspera (from Brunei),
however, have their dorsal surfaces distinctly more rugose(Fig. 4B–D) than any specimen of S. tenebrosa. The formof the male abdomen for specimens is useful, but only
when they are of comparable sizes. Large specimens ofmany gecarcinucids often have the abdomen relatively more
slender, often with somite 6 proportionately longer whencompared to smaller ones. For S. aspera, the type is thelargest known specimen (42.6 × 32.8 mm), and its maleabdominal somite 6 is relatively elongate (Fig. 6A, B). In
an adult, but smaller male specimen from Brunei (32.3 ×
25.4 mm), somite 6 is relatively broader and less elongate(Fig. 6C). For S. tenebrosa, smaller males (e.g., 21.9 × 17.8mm) still have a relatively elongated abdominal somite 6(Fig. 3F), which in adults, is proportionately longer (Fig.
3D; Holthuis, 1979: fig. 8a).

The known distributions of S. tenebrosa and S. aspera areinteresting. The type locality of S. aspera in Mendalongin Sipitang, Sabah, is some 150 km northeast of the type
locality of S. tenebrosa in Gunung Mulu in northern Sarawak.
These species are found in montane habitats, and as such areprobably easily isolated. The specimens from Temburong in
Brunei are in a location between Mendalong and Gunung
Mulu, but in almost all features, including the form of theG1, they are clearly S. aspera.

Sundathelphusa brachyphallus, new species, can be separated
from S. aspera in having the dorsal carapace surfaces, notably
posterolateral regions relatively less rugose (Fig. 1A–C)(prominently rugose with strong striae in S. aspera; Fig. 4;
Ng & Stuebing, 1989: fig. 1A); the external orbital tooth isvery broad (Fig. 1A–C) (relatively narrower in S. aspera;
Fig. 4; Ng & Stuebing, 1989: fig. 1A); the cervical groove
is relatively less broad and distinct (Fig. 1A–C) (relatively
broader in S. aspera; Fig. 4); the third maxilliped is relatively
more slender (Fig. 2A) (versus more quadrate in S. aspera;
Ng & Stuebing, 1989: fig. 2A); the male abdominal somite
6 is as wide as long (Fig. 2D) (versus longer than wide in
S. aspera; Fig. 6; Ng & Stuebing, 1989: fig. 1B); and theG1 is relatively stouter, with the terminal segment shorterand more swollen (Fig. 7A–D) (versus G1 relatively slender,
with the terminal segment more elongate in S. aspera, at
0.27 times the length of the G1; Fig. 7I–K; Ng & Stuebing,
1989: fig. 2B–E).

Holthuis (1979: 39) reported that the two type males of S.
tenebrosa were 25 × 21 mm and 22 × 19 mm in size, but
the specimens on hand (NHM 1979.135 and 136) measure
23.5 × 19.1 mm and 21.9 × 17.8 mm, respectively. The
difference is not substantial.
The most distinct differences between S. tenebrosa and S.
aspera are in the structures of the frontal median triangle andG1. In S. tenebrosa, the dorsal crista of the frontal median
triangle is distinct, but is not completely fused with the lateral
margins. There is either a small but distinct gap between thecristae, or it joins by a series of uneven flattened granules(Fig. 3B, H, I) (see also Ng, 1988: fig. 38C). In S. aspera,
the dorsal margin of the frontal median triangle is contiguouswith the lateral margins, even if the two margins can still be
discerned by a groove (Fig. 5A–E). This is obvious even insmall specimens and females (Fig. 5C, D). The G1 terminal
segment of S. tenebrosa is relatively shorter than that of S.
aspera (0.21 versus 0.27 times the total G1 length), withthe distal opening relatively smaller (Fig. 7F–H versus Fig.
7I–K). In addition, the distal half of the terminal segment
is relatively more sinuous and elongated in S. aspera (Fig.
7I–K) compared to that of S. tenebrosa (Fig. 7F–H). These
gonopodal differences are evident even in smaller specimensof each species examined.

Compared to S. tenebrosa, S. brachyphallus has the dorsal
margin of the frontal median triangle completely contiguouswith the lateral margins (Fig. 1C, D) (versus with a small
gap or joining as a series of low granules; Fig. 3B, H, I);
it has a male abdominal somite 6 which is squarish, being
as wide as long for specimens of similar or smaller sizes(Fig. 2D) (versus subrectangular, longer than wide with thelateral margins gently concave in S. tenebrosa; Fig. 3D, F;
Holthuis, 1979: fig. 8a); and the G1 is distinctly stouter,with the terminal segment proportionately shorter and moreswollen (Fig. 7A–D) (versus G1 relatively slender, with theterminal segment longer in S. tenebrosa, at 0.21 times thelength of the G1; Fig. 7F–H; Holthuis, 1979: fig. 8b, c).

There are other differences between the two species but
these are size dependent or may vary. The cervical grooveof S. aspera (Fig. 4) is distinctly broader than S. tenebrosa(Fig. 3A, C, E, G) when specimens of similar sizes arecompared. The epibranchial tooth of S. aspera is usuallyvisible, even if small, and there is usually a distinct small
V-shaped notch separating it from the anterolateral margin(Fig. 4). In S. tenebrosa, the epibranchial tooth is very
low and hardy demarcated from the anterolateral margin,
with the two structures separated only by a small shallow

The proportions of the G1 terminal segment to overall length
are deceptive, being 0.21 in S. tenebrosa, 0.27 in S. asperaand 0.25 times in S. brachyphallus, even though the G1 of S.
brachyphallus actually appears shorter. This is because in S.
brachyphallus, the overall G1 structure is already short andstout, as is the terminal segment (Fig. 7A, B). In S. tenebrosa501

Ng: New species of Sundathelphusa from southeastern Borneo
and S. aspera, the terminal segment proportions are moreindicative of differences as their subterminal segments havethe same structure (Fig. 7F, I).

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Zoological Journal of the Linnean Society, London, 154:
211–221.

The specimens of S. brachyphallus were collected fromhighlands in southeastern Kalimantan in the karst formationsof the Barito River basin (see Rahmadi, 2006; Rahmadi &Suhardjono, 2004).
ACKNOWLEDGEMENTS
I am grateful to the intrepid cave explorer, Cahyo Rahmadi,
for collecting this interesting material, and to Daisy Wowor(MZB) for passing me the specimens for study. Thanks are
due to Charles Fransen and the late Lipke Holthuis (RMNH)
for permission to examine their specimens so many years ago;
and to Paul Clark (NNM) for the loan of his material. The
author is grateful to Célio Magalhães (Instituto Nacional dePesquisas da Amazonia, Manaus, Brazil) and Jose Mendoza(ZRC) for their suggestions.
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