Revue suisse de zoologie

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de la

SOCIETE SUISSE DE ZOOLOGIE

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MUSEUM D'HISTOIRE NATURELLE
de la Ville de Genève

tome 114
fascicule 4
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H GENEVE DECEMBRE 2007 ISSN 0035 - 418 X

REVUE SUISSE DE ZOOLOGIE

SWISS JOURNAL OF ZOOLOGY

REVUE SUISSE DE ZOOLOGIE

TOME 114— FASCICULE 4

Publication subventionnée par:
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ANNALES

de la

SOCIÉTÉ SUISSE DE ZOOLOGIE

et du

MUSEUM D'HISTOIRE NATURELLE
de la Ville de Genéve

tome 114
fascicule 4
2007

E = ;
HI GENEVE DÉCEMBRE 2007 ISSN 0035 - 418 X

REVUE SUISSE DE ZOOLOGIE

SWISS JOURNAL OF ZOOLOGY

REVUE SUISSE DE ZOOLOGIE

TOME 114— FASCICULE 4

Publication subventionnée par:
ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT)
VILLE DE GENÈVE
SOCIÉTÉ SUISSE DE ZOOLOGIE

DANIELLE DECROUEZ
Directrice du Muséum d’histoire naturelle de Genève

ALICE CIBOIS, PETER SCHUCHERT
Charges de recherche au Muséum d’histoire naturelle de Genève

Comite de lecture

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Genève et de représentants des instituts de zoologie des universités
suisses.

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le
sujet étudié.

La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie,
systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée.

Administration

MUSEUM D'HISTOIRE NATURELLE
1211 GENÈVE 6

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

PRIX DE L’ABONNEMENT:

SUISSE Fr. 225.— UNION POSTALE Fr. 250.—

(en francs suisses)

Les demandes d’abonnement doivent être adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse

REVUE SUISSE DE ZOOLOGIE 114 (4): 663-683; décembre 2007

Morphological analysis of the adult and juvenile instars of
Scutovertex minutus (Acari, Oribatida, Scutoverticidae)

Sylvia SCHÂFFER & Günther KRISPER (corresponding author)
Institute of Zoology, Karl Franzens-University, Universitätsplatz 2, A-8010 Graz,
Austria. E-mail: sylvia.schaeffer @uni-graz.at; guenther.krisper @uni-graz.at

Morphological analysis of the adult and juvenile instars of Scutovertex
minutus (Acari, Oribatida, Scutoverticidae). - Individuals of Scutovertex
minutus (Koch, 1836) were investigated originating from different regions
of eastern Austria, Italy (Southern Tyrol), and Switzerland. Because of the
fragmentary descriptions and scattered morphological data of this species it
is necessary to redescribe it in detail. Characteristics for adult S. minutus
are: Granular cerotegument; rounded rostrum; prodorsal median ridges,
which converge early and do not reach the translamella; notogastral setae
h;_3 and ps, distally slightly lanceolate, h, mostly a little broadened distally,
blunt, and spinose; mentum with irregularly interrupted sclerotized trans-
verse rib; leg chaetome (without solenidia): I (1-4-3-4-18), II (1-4-3-4-15),
III (2-2-1-3-15), IV (1-2-2-3-12). Intraspecific morphological variation was
observed in the shape of lamellar cusps and translamella, and in the length
of prodorsal median ridges, in the shape of lenticulus and in the number of
notogastral setae (10-12 pairs). Breeding experiments produced all juvenile
instars which are here described in detail for the first time. Larva and
nymphs in general show conformity in their habitus. Basic characteristics in
juveniles are the plicate surface of the hysterosoma, larva with 12 and
nymphs with 15 pairs of short gastronotic setae, reddish lateral opistho-
somal glands, short lamellar, interlamellar and exobothridial setae.
Conspicuous characters in the larval stage are the long and thick setae /,
and in the nymphs the knife-shaped lateral setae /’ and /” on tibia I.

Keywords: Taxonomy - morphology - postembryonic development - intra-
specific variation.

INTRODUCTION

Scutovertex minutus (Koch, 1836) is the earliest described and apparently the
most common representative of the genus Scutovertex Michael, 1879. The reported
findings show a Palaearctic distribution. Although this species is known since 171
years, no detailed description or redescription is available. In spite of that, this species
is often mentioned in papers dealing with the ecology and distribution of oribatid mites
living in extreme environments as rocks, roofs, salt marshes or inundation meadows
(e.g., Franz & Beier, 1948; Willmann, 1951; Weigmann, 1973).

Manuscript accepted 25.05.2007

664 S. SCHÄFFER & G. KRISPER

Moreover, our knowledge of the juvenile stages is very poor. Only Grandjean
(1946, 1949) and Haarlgv (1957) gave some information on morphological characters
and Smrz (1992) described some histological features of the juveniles of S. minutus.

The goal of this paper is to redescribe the adults of S. minutus and to describe
the juvenile stages of this species in detail. These results should provide basic data of
external morphology for further comparative investigations on other species of
Scutoverticidae.

MATERIAL AND METHODS

COLLECTING: Adults and juveniles of Scutovertex minutus can be found
throughout the year. They were extracted with Berlese-Tullgren funnels from mosses
and lichens collected on sun exposed rocks and roofs in the eastern parts of Austria.

COLLECTING SITES (numbers in parentheses refer to the numbers of specimens
examined):

Lower Austria: Asparn / Zaya: Eastern outskirt of the village; mosses and
lichens on a tiled roof; 230 m; 16.11.1996; leg. E. Ebermann — (8). Traunstein:
“Wachtstein-Camp”; mosses from rocks on the border of the parking site; 930 m;
31.05.2004 — (1). Türkensturz, Pittental: Mosses in rock crevices; 310 m; 17.10.1997;
leg. R. Schuster — (2).

Upper Austria: Urfahrer Wänd: W Linz; mosses and lichens of a rock; 300 m;
02.05.1997; leg. R. Schuster — (1).

Styria: Frauental, near the castle; mosses and lichens of a tiled roof; 340 m; May
1996; leg. E. Ebermann — (10). Gleichenberger Kogel: Quarry; lichens from rocks; 570
m; 15.06.1991; leg. R. Schuster — (2); 11.02.2007 — (2). Stiwoll: Mosses of the grave-
yard wall; S- and SW-exposed; 500 m; 23.08.2004 — (2). Oblarn: Mosses from a wall
on the riverbank of the river Enns; S-exposed; 670 m; 05.04.2006 — (2). Pogier near
Kapfenberg: Mosses from roof of an old hut; 680 m; 02.10.2006 — (6).

Carinthia: Laas: NW Hermagor; near the parking site of the hospital; mosses
from rocks; SW-SE-exposed; 830 m; 02.09.2004; leg. P. Horak - (1).

Italy, Southern Tyrol: Lüsen, 23.08.1998; ex Coll. Schatz — (7)

44 individuals from these samples were analyzed morphologically and about 50
adults were taken for breeding experiments to get larvae and nymphs.

MICROSCOPIC SLIDES FROM MUSEUM COLLECTIONS : Museum of Natural History,
Basel; Coll. Schweizer: Switzerland, Neuenstadt; slide no. 1477 (labelled as
Scutovertex sculptus) — (8 specimens in one slide).

Bavarıan State Collection of Zoology Munich (ZSM); Coll. Willmann:
Switzerland, Grindelwald, 03.10.1913, slide no. A20041022 — (1 specimen).

REFERENCE MATERIAL: 10 adult specimens, collected in Styria, stored in ethanol,
deposited in the Muséum d’histoire naturelle, Genève.

BREEDING: Adults were put in cylindrical polystyrol-boxes (diameter about
1.5 cm) with a bottom of moistened plaster of Paris. Mosses and lichens taken from the
bark of trees served as food source. To avoid the growth of fungi in the boxes, hyphae
were removed with a fine, soft paintbrush every day.

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 665

SOum

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Fics 1-4

S. minutus, adult. (1) Habitus, dorsal aspect; arrowheads indicate the position of notogastral
saccules Sj, Sj, S3. (2) Lateral part of prodorsum and rostrum; arrowheads point to the two
projecting rostral ridges; arrow indicates the ‘V-shaped’ tutorium. (3) Pedipalp in lateral view.
(4) Habitus, ventral aspect (legs omitted).

666 S. SCHÄFFER & G. KRISPER

PREPARATION: The collected specimens were preserved in 70% ethanol. A
differential interference contrast microscope (Olympus BH-2, equipped with a drawing
tube) was used for investigation in transmitted light. Some specimens were embedded
in lactic acid (as clearing agent) using concavity slides, others were mounted in Swan-
medium (a mixture of arabic gum, aqua dest., glucose, chloral hydrate and glacial
ethanoic acid) as permanent slides. Micrographs were taken with a digital camera
(Olympus Camedia C4040 zoom). For scanning electron microscopy the specimens
were dehydrated in ascending ethanol concentrations, dried in air, mounted on alu-
minium-stubs with double sided adhesive tape and coated with gold. SEM-micro-
graphs were taken at the Research Institute for Electron Microscopy, Technical
University Graz with a Zeiss Leo Gemini DSM 982.

ABBREVIATIONS used in figures: a = anterior subcapitular seta; ad = adanal seta;
ag = aggenital seta; an = anal seta; bo = bothridium; bS = sensillus; Ch = chelicere;
C_3, da, la, dm, Im, dp, Ip, hj_3, ps].3 = notogastral setae; ex = exobothridal seta; € =
famulus on tarsus I; ft”= fastigial seta; g = genital seta; G = gena; gla = opening of
lateral opisthosomal gland; h = hysterostomatic seta; in = interlamellar seta; im, iad =
lyrifissures; le = lamellar seta; Le = lenticulus; lg = labiogenal articulation; M = men-
tum; m = median subcapitular seta; Pdp = pedipalp; ro = rostral seta; RU = rutellum;
0%] > = Solenidia on tarsus; /a, 1b, Ic, 2a, 3a, 3b, 4a, 4b = epimeral setae.

RESULTS

ADULTS

DIAGNOSIS: Habitus corresponding to a typical Scutovertex. Cerotegument (on
notogaster) granulate, cuticle dark and heavily sclerotized, without foveae. Median
converging ridges of prodorsum not reaching translamella. Rostrum anterior of
translamella with circular ridge. Sensillus clavate, spinose. Mentum with irregularily
interrupted sclerotized transverse rib. 10-12 pairs of notogastral setae; setae h>_; and
ps, distally slightly lanceolate, h, often a little broadened distally, blunt, and spinose.
Leg chaetome (excluding solenidia): I (1-4-3-4-18), II (1-4-3-4-15), II (2-2-1-3-15),
IV (1-2-2-3-12).

DESCRIPTION: Habitus (Figs 1, 5): Body contour oval in dorsal view; colour dark
brown to black (in living individuals), in lactic acid and ethanol more light-coloured.

Measurements (n=40): Mean total length: 589 um (range 550-659 um). Mean
notogastral width: 347 um (range 325-380 um).

Integument (Fig. 5): Cerotegument granulate, covering entire body and leg
segments; some granules fused to irregular bars. Whole cerotegument giving the
specimens a rough surface. Granules in cavities and protected areas of the body smaller
and interconnected, forming a reticulate pattern (Fig. 9). Cuticle without foveae.

Prodorsum (Figs 1, 2): Rostrum in dorsal view characterised by a circular ridge,
its anterior part visible in lateral view (Fig. 2) as a projecting ridge. Additionally, on
the outermost part of the rostrum a second ridge projecting beyond the anterior part of
the circular one and looking like a narrow rostral lobe. Rostral setae inserting near
lateral end of this distal ridge. Distinct lamellae; lamellar cusps mostly short and broad,
sometimes long. Lamellar setae originating at the top of cusps and curving towards

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 667

Fics 5-10

S. minutus, adult. (5) Habitus, dorsal aspect; black arrow points to the lenticulus; white arrow
indicates the cuticle bar on the border between prodorsum and notogaster. (6) Sensillus and
bothridium in dorsal view. (7) Dorsal view of left lateral part of prodorsum with V-shaped tuto-
rium and base of leg I. (8) Notogastral seta h3, slightly lanceolate and spinose distally. (9)
Nodule in the humeral region containing lyrifissure ia covered with reticulated cerotegument.
(10) Camerostome and subcapitulum in ventral view; arrow indicates transverse rib of mentum.

each other. Translamella broad and straight or narrow and slightly bent caudad
connecting lamellae. Between bothridia two convergent ridges fused in the middle of
prodorsum and running rostrad, not reaching translamella. Interlamellar setae absent.

Sensillus (Fig. 6) flat, clavate and spinose. Bothridium wide and cup-like,
margin without incision; antiaxial side showing a small apophysis with a ridge running
ventrally.

668 S. SCHÄFFER & G. KRISPER

A V-shaped cuticular elevation (Figs 2, 7) rostrad of leg I placed as the tutorium.

Notogaster (Figs 1, 5): Oval; suture between notogaster and prodorsum incom-
plete medially. Lateral borders of lenticulus concave or parallel or slightly convex. No
pteromorpha, only humeral projections. Octotaxic system represented by three pairs of
very small saccules (S;-S3). 10-12 pairs of setae: c>, da, dm, dp, la, Ip, h,.; and ps} 3;
da and/or dp reduced in most cases (12 pairs observed only once); sometimes setae
reduced one sided. Setae h>_; and ps, distally slightly lanceolate and spinose (Fig. 8),
hj often a little broadened distally, blunt, and spinose. Other setae acute; setae /p the
longest, rest of setae decreasing in length rostrad and caudad. A small, projecting
cuticular nodule under the humeral projection (Fig. 9) with a weak slit (lyrifissure ia)
visible in transmitted light. Lyrifissure im latero-median on the notogaster, ip dorsally
of the line seta ps, and ps); ih and ips on the posterior lateral border of the notogaster.
Openings of lateral opisthosomal glands located in line of lyrifissures im and setae h3;
reservoir of the gland narrow but elongated.

Rostrum and camerostome (Fig. 4): Rostrophragma forming inner margin of
camerostome (Fig. 10). A longish triangular lamella originating from the posterolateral
corner of the camerostome running parallel to the inner border of the camerostome
rostrad; distal end of lamella overlapped by rostral lobe. No genal incision.

Gnathosoma: Subcapitulum diarthric (Fig. 10); rutellum pantelebasic, distally
with four teeth, first one the strongest. Genae possessing sharp lateral edges, one pair
of anterior subcapitular setae (a) and median subcapitular setae (m), both finely serrate.
An irregularly interrupted sclerotised rib running across the mentum; the simple
hysterostomatic setae (h) inserting on it. Pedipalp with five articles (Fig. 3); chaetome:
0-2-1-3-9. Solenidia: 0-0-1. Setae on each segment (femur to tarsus) spiniform and of
different lengths. The four tarsal eupathidia bacilliform, with slightly broadened basis.
Solenidion recumbend, distal end touching the basis of eupathidium acm. Porous
axillary sacculus at basis of pedipalp. Chelicerae with two setae: cha longer than chb.
Trägardh’s organ of same length as moveable digit of chela.

Epimeral region (Fig. 4): Setal formula: 3-1-2-2. All setae acute, slim and
smooth, seta /c located at basis of pedotectum I. Pedotectum I large, completely hiding
acetabulum I. Pedotectum II strongly developed, in horizontal plane “Y-shaped”.
Apodemata different, either reaching median axis or shorter, depending on the degree
of sclerotization of the individuals. Apodemata IV always absent.

Anogenital region (Fig. 4): Genital valves approximately trapezoid. Genital
setae: Six pairs; the two foremost pairs inserting next to each other, the median pair
about twice as long as the lateral pair. One pair of aggenital setae, latero-caudally of
genital opening. A transversal furrow situated closely behind genital opening, whose
ends directed towards the acetabula IV. Anal valves long, posteriorly broadened. A
ridge running along axial border of the valves, between those a groove. A shallow
groove antiaxially of this ridge. Preanal organ cup-like. Two pairs of anal setae
inserting antiaxially of the ridges; setae smooth, acute and slim, an, longer than anp.
Lyrifissures iad situated laterally near front edge of anal orifice. Adanal setae ad, and
ad; located posterior to anal orifice, ad; lateral to it; all of these setae short and spini-
form, inserting on small cuticular bumps.

Legs (Figs 11-18): Tarsi with fine-grained cerotegument except on distal end.
Apotele tridactyl; median claw dorsally weakly spinose, crescent-shaped and stronger

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 669

Fics 11-13

S. minutus, adult. (11) Left leg I (tarsus to femur), antiaxial, ventrolateral aspect. (12) Right
tarsus I, antiaxial aspect. (13) Left leg II, antiaxial aspect.

than the two lateral, clearly spinose claws. All femora with one dorsal stigma, from
there one trachea running to distal part of tibia (legs I-III) or to proximal part of tarsus
(leg IV); in legs I and IT a second short trachea running in opposite direction and ending
within femur. Additionally, one trachea within trochanter of legs III and IV; stigma
opening proximally paraxially. Chaetome and solenidia of legs see Table 1. Leg I (Figs
11, 12): Femur elongated, with short irregular ridges. Genu shortest segment with a
conspicuous ridge; solenidion (0) on a small bump. Solenidia of the tibia inserting on
a strong apophysis distally; @, very long, whip-shaped, 9, short and straight. Tarsal
solenidia (w, and ©, — second one thinner and shorter than first one), famulus (¢) and

S. SCHAFFER & G. KRISPER

670

0-1-0 NECA = SO] P = = = JMPV
0-1-0 GAVAGE? || = «Ad = = <A ydurkuoyuay,
0-1-0 NAGCLED (D) s (21) [pa lee ‚ao p ydurduojneg
0-0-0 L-0-0-0-0 (ad) (n) (d) Uf yduiAuojo1g AI FIT
0-11 CGIE = SO] P 1S0] P = > JnpV
0-1-I SIS AGE (11) (1) = — A ydw uo,
O-I-1 splelaezi 5 = = = A yduÂuoynoq
0-11 €1-C-C-C-0 De = = = ydwAuoJoag
O-I-I €1-7-7Z-Z-0 (ad) (b) s (n) (d) (21) (4) [pa sp] da p eAde] III SIT]
CITI sSI-re-r-1 su JSOT P SO] P = = JInpV
ol SI-SHr-1 (11) A A (1) ut yduAUOJAL,
GI €I-b-€-b-0 co 1 = = ydwAuoynaq
IST €1-€-€-T-0 = = = = ydwAuoJoaq
I-I-I CI-£-£-T-0 Lo (ad) (0) s (n) (d) (21) (14) Op..a,.] O P (1) «Aq P BAIBT II 397
GE) sI-rerI = JSOT P 1S0] P = = JINPV
TTI 81-S-t-t-1 (11) Cha A = aA ydwAuoyLay
GEA 91-[sIr-E-r-0 Co = = (1) yduAuoynoq
Gala] 91-b-€-T-0 = = = = yduAuo3o1q
Tabs OI-p-£-T-O los (7d) (ad) (v) s(n) (d) (2) (9) Tbp.a(ı) OP) QD BAIUT 1391
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‘UOTJELIBA [ENPIAIPUI = | ] ur SIOQUNN “erpruofos = @ “D ‘0 'adejs Suiposoid oy} 0) piedoI YIM 98uPU9 OU = — 9898 JO sured = ( )

sosoyjuared ur JI ‘s19N9] Aq pastiajoeseyo JeI9S Jo Juouudo[eA9P ISAT *SIPISUI [fe JO UONPJ9S Fa] ‘SMNUIU X9J1940/N9S "| ATAVL

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 671

Fics 14-15

S. minutus, adult; distal part of tarsus I. (14) Solenidia and famulus. (15) Basal part of solenidion
©] With short companion seta ft”.

seta ft” close to each other (Figs 14, 15). Leg II (Fig. 13): Femur stocky, with thick
cerotegument. Solenidion of genu (0) short and delicate. Tibial solenidion (@) long and
situated distally. Tarsus with two slim and long solenidia. Leg III (Figs 16, 17):
Trochanter with approximately triangular shape. Femur broad, with ribbed surface,
ventrally with lamella. Solenidion of genu (0) on an inconspicuous bump. Tibial
solenidion (9) short. Seta ft” of tarsus long, spiniform. Leg IV (Fig. 18): Trochanter
big, broad. Femur elongated, ventrally with lamella; cuticle with conspicuous short
irregular ridges. Tibia long, slim; solenidion (@) short. Seta ft” of tarsus long,
spiniform.

JUVENILE INSTARS

In general larva, proto-, deuto- and tritonymphs show conformity in their
habitus and they just vary in measurements depending on their postembryonic stage.
Therefore characters that are the same in all stages are mentioned first. Subsequently,
typical features of each juvenile stage will be described.

Habitus: Oval with irregular plication in dorsal view (Fig. 19). Cuticle grey in
permanent slides, only variation in different stages from light grey to dark grey. Living
individuals also grey with silver glimmer. Weakly sclerotised, especially on notogaster
and legs. Cerotegument shows various structures in different body regions (Figs 20,
DINI):

Prodorsum: Rostrum different from that of adults. Rostral setae long, but
lamellar setae primarily very short, acute and inserting on small bumps. The latter not
in same position as in adults, situated more caudally and more medially in the first third
of prodorsum. No lamellar cusps present. Sensillus relatively long, clavate and spinose.
Bothridium cup-like and big. One pair of interlamellar setae (in) located between
bothridia; very short and acute in larval stage, tip cone-shaped in nymphal stages
(Fig. 21). Exobothridal setae near bothridium, often hardly visible because of wrinkled
cuticle. Lenticulus not distinct as in adults; a rectangular slight elevation situated
between bothridia in front of the plicate notogastral area (see Fig. 19).

Hysterosomatic region: Conspicious prodorsum and hysterosomatic region
separated by an almost straight suture. Circular reservoirs of lateral opisthosomal

672 S. SCHÄFFER & G. KRISPER

Fics 16-18

S. minutus, adult. (16) Right leg III, antiaxial aspect. (17) Left tarsus III, antiaxial aspect. (18)
Right leg IV, antiaxial aspect.

glands situated postero-laterally; their reddish content shining through the cuticle of
living individuals but this stain disappearing on permanent slides. Cupules (lyri-
fissures) always developed as faint disk-like structures; their number corresponding to
the stages (see below).

Gnathosoma: Nymphal subcapitulum with axillary saccules on basis of palps as
in adults.

Epimeral region: Number of setae different between juvenile stages. Median
area of epimera covered with fine wrinkles. Apodemata not reaching median axis.

Anogenital region: Whole anogenital region wrinkled like dorsal part of hys-
terosoma.

Legs: Cerotegumental structure (Fig. 22) consisting of mushroom-like struc-
tures. Most of these formations coated with another fine granulated layer (Fig. 23). No
tracheae in legs, but at least femora showing porose areas in the deuto- and
tritonymphal stage. Number of setae increasing from stage to stage. Chaetome and
solenidia of legs, see Table 1. Dorsal setae on genu and tibia paired with the solenidium
(Fig. 24). Various setae differing in length, thickness and serration (e.g., Fig. 25).
Lateral setae (/’ and /”) on tibia I of nymphs of special form: Thick, knife-shaped and
serrate (Fig. 26).

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 673

Fics 19-24

S. minutus. (19-21) Deutonymph. (19) Habitus, dorsal view. (20) Fine structure of cerotegument
between two wrinkles of notogastral cuticle. (21) Right interlamellar seta. (22-24) Tritonymph.
(22) Cerotegument of right tarsus I, axial aspect, near ff”. (23) Cerotegument of leg III. (24)
Solenidion @ with companion seta d of tibia II.

LARVA (Figs 27, 31): Body length (n=3): 247-250 um. Body width: 156-163
um. Translamella not discernible in this stage. Rostral setae straight and forward-
directed. 12 pairs of gastronotic setae: c;_3, da, la, dm, Im, dp, Ip, hj_3. Cupules ia and
im in dorsal aspect visible; ih anterior and ip posterior to anal orifice. Epimeral setae:
2-1-2; seta Jc on epimeron I not developed. Anal valves hardly visible because of
wrinkled cuticle. Two pairs of notogastral setae hy and h3 near anal valves. Setae h;
short and acute, inserting lateral of the valves; hy remarkably long and thick, located

674 S. SCHAFFER & G. KRISPER

Fics 25-26
S. minutus, tritonymph. (25) Left tarsus II, antiaxial aspect. (26) Right tibia I, lateral seta /’.

caudally (see Fig. 31). Legs (Figs 35-37). Apophysis on leg I approximately as long as
tibia. Claparède organs dome-shaped.

PROTONYMPH (Figs 28, 32): Body length (n=2): 350-375 um. Body width:
219-231 um. No distinct lamellae and translamella. 15 pairs of short gastronotic setae,
C 7-3 thin, the remaining ones thicker. Cupules ips lateral near front edge of anal valves;
ih and ip displaced laterally. Formula of epimeral setae: 3-1-2-1; seta 4a in the middle
of epimeron IV. One pair of genital setae on genital valves. Aggential, anal and adanal
setae not developed. Legs (Figs 38-41).

DEUTONYMPH (Figs 29, 33): Body length (n=6): 450-500 um. Body width:
281-325 um. Rostral setae slightly curved towards each other. Cuticular ridges
extenting rostrad from bothridium, apically transversally connected. The short and
acute lamellar setae close behind this transversal ridge. 15 pairs of short and slim
gastronotic setae inserting on bumps of different height; location same as in
protonymphs. Cupules iad situated in same position as in adults; ips dislocated next to
ih. Formula of epimeral setae: 3-1-2-2; seta 4b situated near the median axis. Genital
valves clearly outlined; still narrow and almond-shaped in closed condition. Two or
three pairs of genital setae present. One pair of aggential setae lateral of genital valves.
Anal valves already well-developed but without anal setae. Three pairs of adanal setae
(first appearance in juveniles), situated laterally of anal valves. Legs (Figs 42-45). Two
of the raised individuals showing intraspecific variation in their leg chaetotaxy: One
specimen with five setae on right tibia I and four on left tibia I; another one with three
setae on both tibiae III.

TRITONYMPH (Figs 30, 34): Body length (n=3): 531-613 um. Body width:
338-406 um. Prodorsal ridges resembling lamella and translamella of adults; short and
acute lamellar setae behind ridges. Rostral setae long and curved towards median axis.
Lateral opisthosomal gland poorly shining through the cuticle because of darker colour
of individuals. 15 pairs of short and acute gastronotic setae. Cupules ips displaced
posteriorly in line of ih and ip. Formula of epimeral setae: 3-1-2-2. Each genital valve
with five setae. One pair of aggenital setae latero-caudally of valves. Anal valves

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 675

4 72 TI ®
{ Ca /
N <=
) =
DÀ, ji
ZN
a

Fics 27-30
S. minutus, juveniles; habitus in dorsal view. (27) Larva. (28) Protonymph. (29) Deutonymph.
(30) Tritonymph.
narrow and elongated. Valves surrounded by two adanal plates, each of them with three
adanal setae. Setae ps), ps, and ps; laterally and caudally of anal region inserting on
short bumps. Legs (Figs 46-49): Tibia I longer than in previous stages, therefore
apophysis relatively shorter.

676 S. SCHAFFER & G. KRISPER

DISCUSSION

ADULTS

We decided to morphologically analyse the adults of Scutovertex minutus in
detail, because of the many not yet examined features on the one hand and because of
the existence of different figures of this “species” in oribatid papers and keys on the
other hand. Drawings of the dorsal view of “S. minutus” show diverging body shapes,
various sensilli, and homologous notogastral setae of different length and shape
(compare Strenzke, 1943; Schweizer, 1956; Ghiljarov & Krivolutsky, 1975; Balogh,
1972, 1992; Pérez-Inigo, 1993).

The types on which the original description by C. L. Koch is based were
collected from the moat of Regensburg (Bavaria). Our two attempts to collect addi-
tional S. minutus material in Bavaria (Großer Arber, Neu Schwanstein) were not
successful. Unfortunately, no Scutovertex-specimens of the collection of C. L. Koch
are still available for comparison; requests addressed to the Museum of Natural
History, Berlin, and to the Museum of Natural History of London brought no positive
results. The short descriptions of S. minutus (Koch, 1836) and S. ovalis (Koch, 1841)
and Koch’s tiny illustrations of these species allowed a free and broad interpretation of
their characters in the past. The description of S. sculptus Michael, 1879 seems not to
have been helpful for most acarologists in the first half of the 20% century. In 1928
Sellnick presumed that S. minutus and S. sculptus are synonyms. Van der Hammen
(1952) reported on incorrectly labelled microscopic slides of Scutovertex-species in the
collections of Berlese and Oudemans. We also found that phenomenon in three
different collections: Schweizer identified at least once S$. minutus as S. sculptus (slide
no. 1477). Furthermore, most Scutovertex-slides of the collection Willmann are
labelled as S. ovalis, although Willmann only mentioned S. minutus and S. sculptus in
his key of 1931. However, in our investigation of his slides we could not find any
noticeable differences between the specimens of S. minutus and S. ovalis. Some vials
of the collection Moritz are labelled as S. minutus, others as S. sculptus — none of them
contain §. minutus.

Although Strenzke (1943) already demonstrated some clear differences
between S. minutus and S. sculptus, his work was neglected by many authors, resulting
in different opinions on the characteristics of this species (see above). Later, Strenzke
relativised his work in a note sent to Haarlgv (1957: 47) mentioning the problem of
determination based on variation of characters and asking for an investigation of
specimens originating from a wide area and from different habitats to clarify this
problem. Haarlgv himself regarded S. minutus and S. sculptus as synonymous because
of “intermediate forms”.

In our study, which refers to material from eastern Austria, supplemented by
specimens from S-Tyrol and Switzerland, we restrict the limits of the species S. minu-
tus according to our diagnosis. Intraspecific variation includes the number of noto-
gastral setae, shape of lamellar cusps, and of prodorsal ridges. In general our results
correspond to the characters and figures given in the key of Weigmann (2006). He
reported on facultative problems to distinguish between S. minutus and other species
caused by a relatively high intraspecific variation (see also Pérez-Inigo, 1993). We do
not agree with the hypothesis of such a high extent of intraspecific variation assumed

MORPHOLOGICAL ANALYSIS OF 5. MINUTUS 677

Fics 31-34

5. minutus, juveniles; habitus in ventral view (subcapitulum not drawn). (31) Larva (note the
strong setae 43). (32) Protonymph. (33) Deutonymph. (34) Tritonymph.

by the authors mentioned above. Even though we have also found specimens not
completely corresponding to our diagnosis, we want to exclude these “diverging
specimens” from determination or description at the present state of knowledge. We

678 S. SCHÂFFER & G. KRISPER

100um

Fics 35-37

S. minutus. Larva. (35) Right leg I, antiaxial aspect. (36) Left leg II, axial aspect. (37) Right leg
III, antiaxial aspect.

assume that future results of an ongoing detailed morphological and molecular genetic
study of other European species (e.g., S. sculptus, S. alpinus) collected from various
places will clarify the taxonomic status of these “intermediate forms”. Therefore, and
due to the limited knowledge on morphological characters and their intraspecific
variation in other species, we omit a detailed comparison between S. minutus and the
remaining Scutovertex-species in order to avoid the false impression that these species
are already well-defined.

Morphological characters as camerostome, lyrifissure ia and tracheae of legs,
could play an important role in the classification of genera of Scutoverticidae:
Although Grandjean (1952) already stated that Scutovertex does not possess a genal
incision, Ghilarov & Krivolutsky (1975) as well as Sitnikova (1980) reported on an
aberrant notch on the border of the camerostome. SEM investigations of the lateral

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 679

Fics 38-41

S. minutus. Protonymph. (38) Right leg I, antiaxial aspect. (39) Left leg II, antiaxial aspect. (40)
Left leg III, antiaxial aspect. (41) Left leg IV, antiaxial aspect.

parts of the podosoma have shown that there is no (genal) incision or cleft in the rostral
margin. Nevertheless, these morphological details of the border of the camerostome
might be important in studying the relationships of Scutoverticidae.

The cuticular nodule (see Fig. 9) under the humeral projection shows in trans-
mitted light a small slit which represents the lyrifissure ia; otherwise the ia would be
missing. This nodule probably has its equivalent in a similar disc-like structure
described in Argentinovertex coineaui Fernandez & Cleva, 2002. In this case the
authors have neither found a slit on the disc-like structure, nor the lyrifissure ia. The
position and the form of this organ might be comparable to the humeral organ of other
Poronota, but histological investigations are necessary to clarify this problem.

680 S. SCHAFFER & G. KRISPER

100um

45

100um

Fics 42-45

S. minutus. Deutonymph. (42) Left leg I, antiaxial aspect. (43) Right leg II, antiaxial aspect. (44)
Left leg III, antiaxial aspect. (45) Right leg IV, antiaxial aspect.

Saccules and brachytracheae can be found in different leg segments of several
taxa of oribatid mites, but the occurrence of true tracheae in legs is very rare. They are
known only from the ameronothroid genus Aquanothrus Engelbrecht, 1975 (Norton et
al., 1997) and from the here investigated licneremaeoid genus Scutovertex. This
character might be useful for the diagnosis of the genus Scutovertex, as assumed by
Grandjean (1940), but further comparative investigations are necessary to verify that.

JUVENILE INSTARS

Data on the external morphology of the juvenile stages of Scutoverticidae are
available only in few cases. Michael (1884) gave a general description of a nymph of
Scutovertex sculptus without information on the stage. Grandjean (1954) mentioned
some characters of nymphs of an undetermined species of Scutovertex. Our

MORPHOLOGICAL ANALYSIS OF S. MINUTUS 681

100um

100um

Fics 46-49

S. minutus. Tritonymph. (46) Left leg I, antiaxial aspect. (47) Right leg II, antiaxial aspect. (48)
Left leg III, antiaxial aspect. (49) Left leg IV, antiaxial aspect.

investigations on juveniles of S. minutus generated morphological features comparable
with those described by Grandjean; therefore we assume that the latter author had used
juvenile individuals of S. minutus in his study. Haarlgv (1957) published an illustration
of a tritonymph of S. minutus showing the dorsal side, the anal and genital region. The
depicted rostral setae seem to be slimmer than those the specimens we examined.
Furthermore the tritonymphs of Arthrovertex (=Argentinovertex) coineaui (see
Fernandez & Cleva, 2002) and of Provertex delamarei Travé, 1962 are known.

682 S. SCHAFFER & G. KRISPER

Due to the lack of knowledge on the morphology of juvenile Scutoverticidae it
is difficult to compare S. minutus with congenerics and other species and to decide
which of their characters are typical for the family or a certain genus and which are
species specific. This matter becomes complicated if one follows the opinion of Woas
(2002), who stated that on the basis of characters in adults the genera Provertex and
Lamellovertex belong to the family Cymbaeremaeidae. In our opinion the available
data on juveniles (and adults) are too poor to make a clear decision on this systematic
question at the moment.

ACKNOWLEDGEMENTS

The authors thank Prof. Dr F. Hofer, head of the Research Institute for Electron
Microscopy, and his team for the realisation of the SEM micrographs. We are grateful
to Prof. Dr R. Schuster, Dr E. Ebermann, Dr H. Schatz and Dr P. Horak providing
samples and specimens. Furthermore, we want to express our thanks to the following
museums for loaning slides and specimens: Bavarian State Collection of Zoology
(Munich) (ZSM), collection Willmann; Museum of Natural History, Humboldt-
University (Berlin), collection Moritz; Natural History Museum Basel, collection
Schweizer. This work was supported by the Austrian Science Foundation (FWF,
project number P19544-B 16).

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REVUE SUISSE DE ZOOLOGIE 114 (4): 685-692; décembre 2007

Description of the male of Hersilia vicina Baehr & Baehr, 1993
from northeastern Thailand, with notes on the albomaculata-group
(Araneae, Hersiliidae)

Thanaphum CHAMI-KRANON, Chalobol WONGSAWAD

& Pakawin DANKITTIPAKUL*

Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai
50200, Thailand.

* Corresponding author, E-mail: pakawin @ gmail.com

Description of the male of Hersilia vicina Baehr & Baehr, 1993 from
northeastern Thailand, with notes on the albomaculata-group
(Araneae, Hersiliidae). - New material of Hersilia vicina Baehr & Baehr,
1993 has become available from Pa Hin Ngam and Nam Nao National
Parks, northeastern Thailand. From that material the male of Hersilia vicina
is described and illustrated for the first time. It can be recognized by the
curved embolus gradually tapering towards its tip and differs from other
males of the albomaculata-group by the presence of strong dorsal spines on
the slightly elevated ridge of the palpal tibia, and by the hook-shaped
basally excavated tegular apophysis. The newly discovered male of H.
vicina confirms that the albomaculata-group 1s a distinct species-group.

Keywords: Taxonomy - conspecific sex - albomaculata-group - Pa Hin
Ngam National Park - Khao Yai National Park - Nam Nao National Park.

INTRODUCTION

The hersiliid fauna of Thailand is poorly known. Hersilia Audouin 1826 is the
largest and most diverse genus within the Hersiliidae, comprising approximately 75
described species distributed over the tropics of Africa and Asia, and in parts of
Australia (Baehr & Baehr, 1993; Foord & Dippenaar-Schoeman, 2006; Platnick,
2007). Baehr & Baehr (1993) published a taxonomic revision of the hersiliid spiders of
the Oriental Region that included the Thai species and their known distributions at that
time. Foord & Dippenaar-Schoeman (2006) recently revised the Afrotropical species
of the genus Hersilia and added fourteen new species to the genus. Five valid species
are currently known from Thailand. These are H. vicina Baehr & Baehr, 1993 (Khao
Yai National Park), H. asiatica Song & Zheng, 1982 (Phu Kradung National Park,
Khao Yai National Park), H. simplicipalpis Baehr & Baehr, 1993 (Doi Suthep-Pui
National Park, Khao Yai National Park), H. striata Wang & Yin, 1985 (Doi Inthanon
National Park, Erawan National Park, Sam Roi Yod National Park) and H. clypealis
Baehr & Baehr, 1993 (Khao Yai National Park). The first described species from

Manuscript accepted 04.06.2007

686 T. CHAMI-KRANON ET AL.

Thailand, Hersilia siamensis Simon, 1886, collected at a ‘Canal de Bangkok, Siam’ is
regarded as anomen dubium (Baehr & Baehr, 1993: 78) because the female type was
presumably lost (not found in MNHN nor MSNG) and because the original description
given by Simon (1886) did not include satisfactory taxonomic information on the
female genitalia. Hersilia asiatica has a relatively broad distributional range covering
the southern part of China and extending into the mountainous areas of northeastern
Thailand through the Luang-Prabang Range, which is a natural boundary between
Thailand and Laos (Baehr & Baehr, 1993; Song & Zheng, 1982; Song, Zhu & Chen,
1999). Hersilia striata occurs in China, Thailand, Myanmar and beyond the Malay
Peninsula on the islands of Sumatra and Java (Baehr & Baehr, 1993; Song, 1987; Song,
Zhu & Chen, 1999; Wang & Yin, 1985). Hersilia vicina and H. clypealis appear to be
endemic to Thailand; each is known only from a single locality and no further infor-
mation has become available after their descriptions (Baehr & Baehr, 1993). Hersilia
vicina undoubtedly represents a species of its own but its exact placement in the
species-group required further taxonomic scrutiny because of the lack of males at that
time. Hersilia simplicipalpis has been reported from northern and northeastern
Thailand. Dankittipakul (2002) collected males of another hersiliid genus from Doi
Inthanon National Park but they do not belong in Hersilia because they posses rela-
tively short PLS comparing to the Hersilia species (terminal segment of PLS 1.1-1.3
times longer than the basal one). Of the nominal species reported from Thailand, four
have been recorded from the Khao Yai National Park. This indicates not only the
species richness of the Thai hersiliid fauna, it also reveals that this fauna is superficially
known and that more taxonomic work needs to be undertaken.

MATERIAL AND METHOD

External morphology was examined, measured and drawn with an Olympus
SZX-9 stereomicroscope equipped with a drawing tube. Measurements of leg
segments were taken from the dorsal side. All measurements are in millimeters.
Identifications were made by examination of male and female genitalia. Epigynes were
drawn in natural and cleared state (after immersing them in lactic acid for 10-20
minutes). Male palps were drawn in lateral and ventral view. Sizes of eyes are given as
proportions of AME (= 1.00) in the following order: AME: ALE: PME: PLE.

The material examined will be deposited in the collections of the Museum
d’histoire naturelle de la Ville de Genève, Switzerland (MHNG) and in the collections
of the Thailand Natural History Museum (TNHM) of the National Science Museum,
Pathumthani Province, Thailand. Other museum acronyms: MNHN, Museum National
d’Histoire Naturelle, Paris, France; MSNG, Museo Civico di Storia Naturale
“Giacomo Doria”, Genova, Italy.

Abbreviations used in text and in the figures are as follows: ALE, anterior
lateral eyes; AME, anterior median eyes; BS, basal segment of posterior spinneret; CO,
copulatory orifices; DMP, dorsal muscular pits; DS, dorsal spine; E, embolus; EP,
median epigynal projection; ID, insemination ducts; PLS, posterior lateral spinnerets;
PME, posterior median eyes; PLE, posterior lateral eyes; SC, socket of spine
(removed); SD, sperm ducts; SR, seminal receptacle; ST, subtegulum; T, tegulum; TA,

NEW MATERIAL OF HERSILIA VICINA FROM THAILAND 687

tegular apophysis; TB, trichobothrium; TC, tarsal spines; TS, terminal segment of
posterior spinneret.

In the text ‘Fig. refers to a figure herein, while ‘fig.’ refers to a figure published
elsewhere.

TAXONOMY

Hersiliidae Thorell, 1870
Hersilia Audouin, 1826

Hersilia vicina Baehr & Baehr, 1993 Figs 1-9
Hersilia vicina Baehr & Baehr (1993: 22, figs 18e-f), description of 9.

HoLoTypE: 9, THAILAND, Nakhon Ratchasima Province, Khao Yai National Park,
night collecting, forest in center, 8.x1.1987, leg. C.L. & P.R. Deeleman [MHNG, examined].

OTHER MATERIAL EXAMINED: 2 d, Chaiyaphum Province, Pa Hin Ngam National Park,
creek at field of Siam tulip (Curcuma alismatifolia, Zingiberaceae), in mixed deciduous forest
with pines, 15° 38.391’N 101° 23.609’E, 750 m alt, Malaise trap, 12-18.vii.2006, P.
Dankittipakul & N. Likhitrakarn leg. [MHNG, TH-HS001; TNHM]. 2 2, Phetchabun Province,
Lom Sak and Lom Kao Districts, Nam Nao National Park, 890 m, evergreen gallery forest near
national park headquarters, 17.-18.vii.2005, P. Dankittipakul leg. [MHNG, TH-HS002; TNHM].

EXTENDED DIAGNOSIS: Males of H. vicina resemble those of H. albomaculata
(see Baehr & Baehr, 1993: figs 16c, d) and H. martensi (Baehr & Baehr, 1993: figs 17c,
d) by the curved embolus gradually tapering towards its extremity (Figs 4-6). Hersilia
vicina males can be distinguished from other males of the albomaculata-group by: The
presence of strong dorsal spines on the palpal tibia (Fig. 3); tegular apophysis hook-
shaped and basally excavated (Fig. 6) (thin and membranous, with dorsal excavation
in H. albomaculata and H. martensi). Females of H. vicina can be easily recognized
by: The digitiform median extension of the epigyne (Fig. 7); the elongated and curved
insemination ducts with a globular anterior part (Figs 8, 9); two spherical seminal
receptacles clearly separated and arising posteriorly on both sides (Figs 8, 9).

DESCRIPTION: MALE. Total length (excluding spinnerets) 5.09. Carapace length
2.38, width 2.50. Opisthosoma length 2.71, width 3.24. PLS length 4.19; BS length
0.84; TS length 3.35. Eye ratios: 1.0: 0.46: 0.86: 0.93. Clypeus 0.32 high.

Prosoma (Figs 1, 2) dorsoventrally flattened, with slightly elevated eye area and
low clypeus, pars thoracica widest. Carapace yellow, with greenish brown radial striae
radiating from deep longitudinal fovea; lateral margins black, widest posteriorly,
clothed with conspicuous setae. Clypeus (Fig. 1) pale grey, with a cross-shaped
marking medially; clypeal margin with faint black stripe. Chelicerae (Fig. 1) basally
yellow, distal part with irregular dark greenish patches; three reddish brown promar-
ginal teeth on cheliceral furrow. Sternum, labium and maxillae yellow. Eight eyes
arranged in two strongly recurved rows (Figs 1, 2); MOQ wider than long, wider in
front than behind (Fig. 1); eye area with distinct dark reddish brown pigmentation, with
red pigment encircling ALE; a red longitudinal band running between PME and fovea.
Patellae brown, other leg articles greenish brown, with green annulations on femora;
distal part of tibiae dark brown.

Opisthosoma (Fig. 2) widest in posterior third, with four pairs of dorsal mus-
cular pits, the second ones largest. Dorsum mottled with irregular white patches;

688 T. CHAMI-KRANON ET AL.

Fics 1-3

Hersilia vicina Baehr & Baehr, male from Pa Hin Ngam National Park: (1) Prosoma, frontal
view. (2) habitus, dorsal view. (3) right palpus, dorsal view. Scale bars = 1.0 mm (1, 2), 0.5 mm

(3).

NEW MATERIAL OF HERSILIA VICINA FROM THAILAND 689

cardiac impression dark brown, forming folium, extending to caudal area; anterolateral
and posterolateral borders dark. Venter pale green, mottled with small, white irre-
gularly-shaped patches, marked with a series of muscular pits arranged in a V-shaped
pattern. Posterior lateral spinnerets long; terminal segment more than two times longer
than basal one; numerous aciniform gland spigots on entire length of terminal segment;
basal segment with dorsal patch of reddish pigment distally; terminal segment with two
disconnected reddish bands.

Leg measurements:

I II II IV
Femur 5.08 5.36 1.64 4.61
Patella 112 OS) 0.50 0.85
Tibia 3.25 3.42 0.98 ZETA.
Metatarsus 1295 7.90 1.20 3.02
Tarsus 0.85 0.91 0.63 0.87
Total 20.21 20.64 4.95 12.06

Male palp (Figs 3-6): Palpal tibia provided with a slightly elevated dorsal ridge
carrying 17-20 relatively long spines (Figs 3, 6, SC, DS) with reddish basal and white
distal part; dorsal side with a row of seven retrodorsal trichobothria (TB) arranged in a
bent line and with a group of four short prodorsal-basal trichobothria (Fig. 3).
Cymbium relatively short, with two apical spines (Figs 4-6, TC). Tegulum (T) round,
pale yellow, with distinct dark brown sperm duct (Figs 4-6). Embolus (E) filiform,
curved to 3/4 of a circle, laterally with a small triangular denticule near its tip (Figs
4-6). Tegular apophysis (TA) hook-shaped, apically inserted, pointed distally, basally
excavated (Fig. 4), visible as a dark anteriorly curved spine rising from a round
membranous base when viewed from the lateral sides (Figs 5, 6).

FEMALE (holotype). Total length 4.86. Carapace length 2.11, width 2.15.
Opisthosoma length 3.05, width 2.58. PLS length 3.75; BS length 0.82; TS length 2.93.
Eye ratios: 1.0: 0.37: 0.96: 0.95. Clypeus 0.38 high.

As the male but darker in color. Opisthosoma widest in the middle. Dorsum
greenish black; cardiac impression dark green, occupying 3/4 of opisthosoma length.
DMP variable in size, third and fourth pairs circular, relatively small, second ones
largest.

All legs broken behind trochanter or femur.

Epigyne and vulva (Figs 7-9): Epigyne (Fig. 7) posteriorly excavated; epigynal
plate lightly sclerotized, with slightly elevated triangular median ridge and digitiform
posterior projection (EP); lateral borders invaginated. Copulatory orifices (CO)
anterior to epigynal projection, clearly marked with red pigmentation. Vulva (Figs 8,
9) with insemination ducts (ID) coiled and tubular, widened anteriorly to form
spherical structures. Two globular seminal receptacles (SR) lying dorsally of spherical
portion of insemination duct; thin, long receptacular stalks basally running into a single
thin fertilization duct on each side.

NEW FEMALE (from Nam Nao National Park). Total length (excluding
spinnerets) 5.57. Carapace length 2.24, width 2.18. Opisthosoma length 3.19, width

690 T. CHAMI-KRANON ET AL.

Fics 4-9

Hersilia vicina Baehr & Baehr, male (4-6) and female holotype (7-9): Right male palpus, (4)
ventral, (5) retrolateral and (6) prolateral view. Epigyne, (7) ventral view. Vulva, (8) ventral and
(9) dorsal view. Scale bars = 0.5 mm (4-6), 0.25 mm (7-9).

2.63. PLS length 4.59; BS length 0.94; TS length 3.65. Eye ratios: 1.0: 0.35: 0.89: 0.97.
Clypeus 0.35 high.
Coloration and pattern as in the male but generally darker.

Leg measurements:

I II II IV
Femur 5.40 5.03 2.91 4.80
Patella 0.98 0.82 0.65 0.81
Tibia ? 5.16 1.50 ?
Metatarsus ? 153 1.72 ?
Tarsus D 0.86 0.71 2
Total ? 19.4 7.49 2

NEW MATERIAL OF HERSILIA VICINA FROM THAILAND 691

DISTRIBUTION: Northeastern Thailand: The provinces of Nakhon Ratchasima
(type locality), Chaiyaphum (new locality) and Phetchabun (new locality). Baehr &
Baehr (1993: 23) erroneously stated that the female holotype of H. vicina was collected
from southern Thailand.

DISCUSSION

Baehr & Baehr (1993) established the albomaculata-group to accommodate
three Hersilia species from Nepal, southeastern China and Thailand. The species-group
comprises H. albomaculata (male and female known), H. martensi (only male known)
and H. vicina (female and now also male known). The following characteristics of this
species-group were given by Baehr & Baehr (1993). Members of the albomaculata-
group are generally small, extremely long-legged species, with leg I slightly longer
than leg II, eye area and clypeus are relatively low. The males of this group are distin-
guishable by: The embolus narrowed and curved; the simple tegular apophysis exca-
vated, situated distally on the tegulum; and by the absence of a spinose ridge on the
male palpal tibia. The females are distinguished by a simple epigyne without pecu-
liarity, the vulva with coiled insemination ducts and one or two pairs of well separated
seminal receptacles.

Previously the taxonomic affinity of the albomaculata-group was obscure. The
female of H. vicina was only tentatively included in this species-group because the
female of H. martensi has not yet been described and the female of H. albomaculata
from China was not known to Baehr & Baehr (1993), who obtained their information
from the original description and illustration given by Wang & Yin (1985). With the
discovery of the male of H. vicina described in here, the albomaculata-group is suffi-
ciently distinguishable from other species-groups. The putative monophyly of the
species-group is here supported by morphological characters of males which are con-
sidered synapomorphic. These characteristics are the curved and narrowed embolus
which forms a twisted circle, and the simple tegular apophysis which is situated
distally on the tegulum.

ACKNOWLEDGEMENTS

We thank Dr Peter Schwendinger (MHNG) for the loan of the type specimen.
He and Dr Barbara Baehr (Queensland Museum) kindly provided constructive
comments on an earlier version of the manuscript.

The Graduate School of Chiang Mai University supported P. Dankittipakul
during his study. The Royal Forest Department gave permission to collect spider
specimens in national parks and other protected areas. Special thanks go to the super-
intendents of Nam Nao and Pa Hin Ngam National Parks and to the park rangers who
organized and assisted the field trips. We are grateful to Dr Angoon Lewvanich (The
Royal Academy of Thailand) and Dr Wipada Vungsilabutr (Department of Entomology
& Zoology, Ministry of Agriculture) for their generous support.

692 T. CHAMI-KRANON ET AL.

REFERENCES

AUDOUIN, V. 1826. Explication sommaire des planches d’arachnides de l'Egypte et de la Syrie
publiées par Jules-César Savigny; offrent un exposé des charactères naturels des genres,
avec la distinction des espèces (pp. 99-186, pl. 1-9). In: SAVIGNY, J. C. (ed.). Description
de l'Egypte. Histoire Naturelle (Paris) 1 (4).

BAEHR, M. & BAEHR, B. 1993. The Hersiliidae of the Oriental Region including New Guinea.
Taxonomy, phylogeny, zoogeography (Arachnida, Araneae). Spixiana, Supplement 19:
1-95.

DANKITTIPAKUL, P. 2002. Diversity, distribution and occurrence of spiders in Doi Inthanon
National Park, Chiang Mai Province. Unpublished M.Sc. thesis, Chiang Mai University,
Chiang Mai, 325 pp.

FOORD, S. H. & DIPPENAAR-SCHOEMAN, A. S. 2006. A revision of the Afrotropical species of
Hersilia Audouin (Araneae: Hersiliidae). Zootaxa 1347: 1-92.

PLATNICK, N. I. 2007. The world spider catalog, version 7.5. American Museum of Natural
History, online at http://research.amnh.org/entomology/spiders/catalog/ index.html (ac-
cessed February 15, 2007).

SIMON, E. 1886. Arachnides recueillis par M. A. Pavie (sous-chef du service des postes au
Cambodge) dans le royaume de Siam, au Cambodge et en Cochinchine. Actes de la
Societe linneenne de Bordeaux 40: 137-166.

Song, D. X. 1987. Spiders from agricultural regions of China (Arachnida: Araneae). Agriculture
Publishing House, Beijing, 376 pp.

SONG, D. X. & ZHENG, S. X. 1982. A new spider of the genus Hersilia from China (Araneae:
Hersiliidae). Acta Zootaxonomica Sinica 7: 40-42.

SONG, D. X., ZHU, M. S. & CHEN, J. 1999. The Spiders of China. Hebei Science & Technology
Publishing House, Shijiazhuang, 640 pp.

THORELL, T. 1870. On European spiders. Nova Acta Regiae Societatis Scientiarum Upsaliensis
7(3): 109-242.

WANG, J. F. & YIN, C. M. 1985. Two new species of spiders of the genus Hersilia from China
(Araneae: Hersiliidae). Acta Zootaxonomica Sinica 10: 45-49.

REVUE SUISSE DE ZOOLOGIE 114 (4): 693-702; décembre 2007

Redescription of Proteocephalus sulcatus (Klaptocz, 1906)
(Cestoda: Proteocephalidea), a poorly known parasite of Clarotes
laticeps (Pisces: Siluriformes) in the Sudan

Alain DE CHAMBRIER!, Toma’ SCHOLZ?, Moges BELETEW?

& Zuheir N. MAHMOUD*

! Department of Invertebrates, Natural History Museum, PO Box 6434,
CH-1211 Geneva 6, Switzerland. E-mail: alain.dechambrier @ ville-ge.ch

2 Institute of Parasitology, Biology Centre of the Czech Academy of Sciences,
BraniSovska 31, 370 05 Ceské Budéjovice, Czech Republic.

3 Department of Biology, Faculty of Science, Addis Ababa University, PO Box 1176,
Ethiopia.

* Department of Zoology, Faculty of Science, University of Khartoum, Khartoum,
Sudan.

Redescription of Proteocephalus sulcatus (Klaptocz, 1906) (Cestoda:
Proteocephalidea), a poorly known parasite of Clarotes laticeps (Pisces:
Siluriformes) in the Sudan.- The proteocephalidean cestode Proteo-
cephalus sulcatus (Klaptocz, 1906), a poorly known parasite described from
widehead catfish Clarotes laticeps (Rüppell) and saddled bichir Polypterus
endlicheri Heckel from the White Nile in the Sudan, Africa, is redescribed
on the basis of new material collected in C. laticeps caught near the type-
locality. New data on the morphology of P sulcatus are provided, based on
the first scanning electron microscopical observations and histological
sections. The most characteristic features of P sulcatus are: (i) pyriform
embryophore; (ii) scolex in form of a four-side truncated cone, with four
suckers deeply embedded within lobes with wrinkled posterior margins; (111)
no apical organ; (iv) a low number (1-3) of mature proglottides; (v) a high
number (115-171) of testes. Clarotes laticeps is considered the only suitable
host for Proteocephalus sulcatus.

Keywords: Proteocephalus - Proteocephalidae - cestodes - catfish parasite -
Clarotes laticeps - Africa - taxonomy.

INTRODUCTION

Klaptocz (1906) described Ichthyotaenia sulcata [(= Proteocephalus sulcatus
(Klaptocz, 1906) La Rue, 1911)] from several tapeworms found in the intestine of
widehead catfish Clarotes laticeps (Rüppell, 1829) (Siluriformes: Bagridae) and
saddled bichir Polypterus endlicheri Heckel, 1847 (Polypteriformes: Polypteridae)
from the White Nile in the Sudan. The author provided a morphological description
supplemented by 10 not very detailed illustrations (one sketch of the total view of the

Manuscript accepted 04.09.2007

694 A. DE CHAMBRIER ET AL.

tapeworm, three figures of the scolex in dorsoventral view, three figures of the scolex in
apical view, one sketch of the oval eggs, one figure of the cirrus-sac and one illustration
of the last three proglottides, with details of internal morphology of the two last ones).
However, differential diagnosis of P. sulcatus from any other species of Ichthyotaenia
(= Proteocephalus) was not included in the original description (Klaptocz, 1906).

Although Khalil (1963) provided some additional data on P. sulcatus (see
Table 1), the morphology of P. sulcatus remains insufficiently known (Freze, 1965) and
no data based on histological sections and scanning electron microscopy (SEM) are
available. New specimens of P. sulcatus found in the intestine of C. laticeps from the
Sudan made it possible to redescribe the tapeworm and to provide new data on its
morphology.

MATERIALS AND METHODS

Widehead catfish (vernacular name in the Sudan “abu misaika”), Clarotes lati-
ceps, were caught by local fishermen in the White Nile at Kosti, about 265 km south
of Khartoum, Sudan, on 25-27 March 2006 (other two C. laticeps from the fish market
in Khartoum examined on 21 March 2006 were free of infection). The fish were
examined immediately after their capture, the parasites found were isolated from the
host intestine and fixed with hot 4% neutral formaldehyde solution and subsequently

TABLE |. Comparative measurements (in millimetres unless otherwise stated) of Proteocephalus
sulcatus from the Sudan.

Host Polypterus Clarotes © Clarotes Clarotes
endlicheri niloticus niloticus niloticus

Reference Klaptocz (1906) Khalil (1963) Present study

Total length 48-68 60 65-95 36-57

Maximum width - - 1.2-1.8 1.48

Scolex

length - - 1.2-1.3 0.66-1.07

width 0.75-1.72 0.67-1.28 1.4-2.0 0.80-1.26

Diameter of suckers 0.25 - - 0.28-0.45

Mature proglottids

length 2. 1.54 1.6-2.3 0.47-1.31
width 1.5 1.9 1.0-1.9 0.92-1.41
Number of testes about 200 117-130 115-171

Cirrus-sac (in um)

length = - 290-310 190-300

width - = 160-180 120-150

Uterine diverticula = - 11-13 15-22

Eggs (in um)
length 29 - 26-28
width 16 - 16-18

REDESCRIPTION OF PROTEOCEPHALUS SULCATUS FROM SUDAN 695

stored in 70% ethanol. The specimens were then stained with Mayer’s hydrochloric
carmine solution, dehydrated in an ethanol series, cleared with eugenol (clove oil) and
mounted in Canada balsam.

Pieces of strobila were embedded in paraffin wax, sectioned at 12-15 um (cross
sections of the strobila and longitudinal and sagittal sections of two scoleces), stained
with Weigert’s haematoxylin and counterstained with 1% eosin B (Scholz &
Hanzelovä, 1998; de Chambrier, 2001). Eggs were studied in distilled water. Three
specimens (scolex with the anterior part of the strobila) were used for SEM
observations using the procedure outlined by de Chambrier et al. (2008).

All measurements are given in micrometres unless otherwise indicated.
Abbreviations used in descriptions are as follows: x = mean, n = number of measure-
ments, OV = ratio of ovary width versus proglottis width (in %), PP = position of
genital pore (cirrus pore) in % of proglottis length, PC = ratio of cirrus-sac length
versus proglottis width (in %). MHNG INVE = Natural History Museum, Invertebrate
Collection, Geneva, Switzerland; IPCAS = Institute of Parasitology, Ceské
Budéjovice, Czech Republic.

Proteocephalus sulcatus (Klaptocz, 1906) La Rue, 1911 Figs 1-21
Syn.: Ichthyotaenia sulcata Klaptocz, 1906

Redescription: Proteocephalidea, Proteocephalidae. Testes, ovary, vitelline
follicles, uterus with uterine stem and diverticula medullary. Strobila with slightly
craspedote proglottides (Fig. 7), 36-57 mm long and up to 1.48 mm wide (Table 1).

Proliferative zone about 700-1100 long and 360-775 wide. Strobila consisting
of 94-120 proglottides: 43-70 immature (up to appearance of spermatozoa in vas
deferens), only 1-3 mature (up to appearance of eggs in uterus), 24-36 pregravid (up to
appearance of hooks in oncospheres) and 26-38 gravid.

Scolex unarmed, wider than neck, about 810-1260 in diameter, and about 665-
1070 long. Scolex a four-sides truncated cone. Suckers uniloculate, 280-450 in
diameter, paired on opposite sides of cone, separated by deep incisions; two sides
without sucker also divided by deep incisions. Distinct apron of tissue posterior to
suckers, longitudinally wrinkled (Figs 1-6). Apical organ absent (Figs 13-15).

Scolex, proliferative zone and anterior proglottides covered uniformly with
relatively short and wide filiform microtriches, only slightly differing from each other
in their shape, length and density (Figs 8-12), except for those of proliferation zone,
which are notably smaller.

Longitudinal internal musculature well developed, anastomosing, forming wide
band of dense muscle bundles; muscle bundles becoming slightly wider near lateral
margins, at level of lateral vitelline follicles (Figs 19, 20). Numerous dorsoventral
muscle fibres present. Subtegumental muscles well developed.

Ventral osmoregulatory canals wide, thin-walled, without anastomoses, over-
lapping lateralmost testes. Dorsal osmoregulatory canal thick-walled, overlapping
dorsally vitelline follicles (Figs 16, 20).

Testes medullary, spherical to oval, 55-120 in diameter, numbering 115-171
(n = 9, x = 143; about 200, according to Klaptocz, 1906), in two or three incomplete
layers, forming two fields confluent anteriorly, with more testes laterally (Figs 16, 19).

696 A. DE CHAMBRIER ET AL.

Fics 1-12

Proteocephalus sulcatus (Klaptocz, 1906) from Clarotes laticeps, Kosti, Sudan (MHNG INVE
54146: 1, 2, 12; MHNG INVE 54141: 3-5; MHNG INVE 54140: 6-11). Scanning electron
micrographs. (1) Scolex, dorsoventral view. (2) Scolex, apical view. (3) Scolex, detail of a
sucker, showing the plicated tissue posterior to sucker. (4) Scolex, dorsoventral view. (5) Scolex,
apical view. (6) Scolex, lateral view. (7) First immature, markedly craspedote proglottides. (8)
Filiform microtriches on apex of scolex. (9) Filiform microtriches on anterior external margin of
suckers. (10) Filiform microtriches on internal surface of suckers. (11) Filiform microtriches
between suckers. (12) Filiform microtriches on external surface of proliferation zone (neck).
Scale bars: 1, 2, 4-6 = 200 um; 3, 7 = 100 um; 8 = 1 um; 9-12 = 2 um.

REDESCRIPTION OF PROTEOCEPHALUS SULCATUS FROM SUDAN 697

External vas deferens strongly coiled, reaching midline of proglottis, never
crossing it aporally. Internal vas deferens thin-walled, very short; ejaculatory duct
thick-walled, long, forming several loops; cirrus long, may occupy complete length of
cirrus-sac. Cirrus-sac elongate, thick-walled, 190-300 (x = 242 + 22; n = 42; CV = 9%),
120-150 wide (L/W ratio = 1.65-2.15, x = 1.85). PC = 16-23% (x = 20 + 1.5%; n = 42;
CV = 8%). Genital pore irregularly alternating, pre-equatorial, situated at 27-46%
(x = 36 + 4%; n = 42; CV = 11.4%) of proglottis length. Genital atrium present
(Figs 16, 17).

Ovary bilobed, medullary, with numerous lobules extending ventrally (Figs 16,
20). OV = 64-71% (x = 68 + 2%; n = 42; CV = 3%). Mehlis’ glands about 100-180 in
diameter, representing 8-17% of proglottis width. Vagina thick-walled, always
posterior to cirrus-sac (n = 150), with higher concentration of chromophilic cells in its
distal (terminal) part, with terminal ring-like vaginal sphincter, difficult to observe in
mature proglottides. Vitelline follicles in two longitudinal bands on both sides of
proglottis, occupying almost its total length; bands interrupted at level of terminal
genitalia on ventral side, with few follicles on dorsal side (Figs 16-18).

Uterus medullary, with development of type 1 according to de Chambrier et al.
(2004), defined as follows: In immature proglottides, uterine stem present as longitu-
dinal concentration of chromophilic cells along median line. Lumen of uterus appears
in last premature proglottides, gradually extending to form tubular structure. Eggs
appear simultaneously with formation of lateral, thick-walled diverticula lined with
chromophilic cells. In gravid proglottides, lateral diverticula remain thick-walled,
15-22 in number on each side, occupy up to 77% of proglottis width (Fig. 18).

Eggs with hyaline, ring-like outer envelope, about 35-42 in diameter; thick,
pyriform embryophore 26-28 long by 16-18 wide, consisting of two layers; outer layer
thicker than nuclei-containing envelope; oncospheres spherical to oval, 9-10 by 12-13
in diameter, with six embryonic hooks 5-6 long (Fig. 21).

TAXONOMIC SUMMARY

Type-host: not designated (see Discussion).

Type-locality: White Nile, between Khartoum and Fachoda (Kodok).

Definitive hosts: widehead catfish Clarotes laticeps (Rüppell) and saddled
bichir Polypterus endlicheri Heckel.

Material studied: 35 specimens collected by A. de C. & T. S. in 8 C. laticeps
from the White Nile at Kosti, Sudan (MHNG INVE 54139, 54140, 54141, 54145,
54146, 54147; IPCAS C-470/1). Five specimens from C. laticeps from the Sudan
(Khartoum) collected by L. F. Khalil in 1959 (MHNG INVE 34798).

Site of infection: intestine.

Prevalence: 73% (infected 8 of 11 fish examined).

Intensity of infection: 1-10 tapeworms (mean 4.4).

Distribution: White Nile in the Sudan. Reports of P. sulcatus from Chrysichthys
sp. from Belgian Congo (now Democratic Republic of the Congo — de Beauchamp,
1914; Mahon, 1954) represent in fact Proteocephalus beauchampi Fuhrmann & Baer,
1925 (Fuhrmann & Baer, 1925; Khalil, 1971).

698 A. DE CHAMBRIER ET AL.

Fics 13-21

Proteocephalus sulcatus (Klaptocz, 1906) from Clarotes laticeps, Kosti, Sudan (MHNG INVE
54145: 13; MHNG INVE 54147: 14; MHNG INVE 34798: 15; IPCAS C 470/1: 16, 17; MHNG
INVE 54139: 18; MHNG INVE 54140: 19-21). (13, 15) Scolex, dorsoventral view. (14) Scolex,
sagittal section. (16) First pregravid proglottis, dorsal view. (17) Terminal genitalia, dorsal view.
(18) Gravid proglottis, dorsal view. (19, 20) Cross sections. (21) Eggs (egg on the right of the
upper line figured from en face view). Scale bars: 13-15, 18-20 = 500 um; 16 = 1000 um; 17 =
250 um; 21 = 20 um.

REDESCRIPTION OF PROTEOCEPHALUS SULCATUS FROM SUDAN 699

DISCUSSION

The present study, which included for the first time SEM observations and
histological sections, confirmed the validity of Proteocephalus sulcatus. The species
possesses several characteristics missing or rare in other proteocephalideans, including
those parasitic in African freshwater fish (Beauchamp, 1914; Fuhrmann & Baer, 1925;
Khalil, 1960, 1963; Lynsdale, 1960; Freze, 1965; Jones, 1980; de Chambrier er al.,
2008). The most typical characters of P. sulcatus are: (1) pyriform embryophore of the
eggs; (ii) scolex a four-sides truncated cone, with deep incisions between suckers and
posterior longitudinal wrinkles; and without apical organ; (iii) only 1-3 mature proglot-
tides present; (iv) a fairly high number of testes (115-171), arranged in 2-3 incomplete
layers.

De Beauchamp (1914) reported P. sulcatus from Chrysichthys sp. from Kilewa
Bay of Lake Tanganyika, Belgian Congo (currently Democratic Republic of the
Congo), but a new species, Proteocephalus beauchampi, Was proposed to accom-
modate this tapeworms (Fuhrmann & Baer, 1925). Mahon (1954) also reported P. sul-
catus from Chrysichthys sp. from Congo, but it is possible that she probably found P
beauchampi as well, because P. sulcatus has not been found in congeneric fish hosts
(Chrysichthys sp.) from the White Nile, examined by the present authors in 2006
(unpublished data). Khalil (1963) provided a brief description of the morphology of
several specimens of P. sulcatus he found in Clarotes laticeps from Khartoum area in
the Sudan (Table 1) and illustrated rather schematically an allegedly mature proglottis
(possibly pregravid or gravid one — see fig. 1 in Khalil, 1963). The present study
confirmed most of Khalil’s (1963) measurements (see Table 1), except for the number
of uterine diverticula, which is in fact much higher than reported by Khalil (1963)
(15-22 vs 11-13 on each side).

Besides P. sulcatus, only two proteocephalidean cestodes possess pyriform
embryophores similar to those of P. sulcatus, namely Proteocephalus beauchampi
Fuhrmann & Baer, 1925 from Chrysichthys catfishes in Africa and Amphoteromorphus
piriformis Carfora, de Chambrier & Vaucher, 2003, a parasite of pimelodid catfish
Brachyplatystoma flavicans (Lichtenstein) in the Neotropical Region (de Beauchamp,
1914 — as P. sulcatus; Freze, 1965; Carfora et al., 2003). Morphology of the eggs has
been proved suitable for species differentiation in some proteocephalidean tapeworms
(Scholz, 1999; Gil de Pertierra & de Chambrier, 2000; Carfora ef al., 2003) and we
strongly recommend that morphological descriptions contain detailed data on egg
morphology.

Proteocephalus beauchampi, a parasite of bagrid catfish of the genus Chry-
sichthys in Africa, resembles P sulcatus in possessing a somewhat similar scolex
without an apical organ, but it differs in the absence of posterior longitudinal wrinkles
(Fuhrmann & Baer, 1925). De Beauchamp (1914) observed in his material of P.
beauchampi (designated as P. sulcatus) also pyriform embryophores of similar shape
and size (25 by 18 um). However, the two taxa differ from one another in strobilar
morphology, especially position of the vagina [always posterior in P. sulcatus and
anterior (44%) and posterior (56%) in Khalil’s material of P. beauchampi] and the
number of testes (57 figured according to fig. 1 of de Beauchamp, 1914). We were not
able to locate the type material of both species. Previous records of P. sulcatus from

700 A. DE CHAMBRIER ET AL.

other catfish, such as Chrysichthys brachynema and Chrysichthys sp. (Prudhoe, 1951;
Mahon, 1954; Khalil & Polling, 1997), are so considered to belong to P. beauchampi.

SEM observation of the surface of the scolex, neck and anterior part of the
strobila of P. sulcatus has shown the presence of relatively short and wide filiform
microtriches, with only slight differences in their shape, size and density between
individual body regions. Only the anterior part of the strobila shows somewhat smaller
filiform microtriches, but of the same shape. This uniformity in the shape of
microtriches differs from that found mainly in Neotropical members of the Proteo-
cephalidea studied using SEM, such as species of the genera Nomimoscolex Woodland,
1934 and Monticellia La Rue, 1911 (de Chambrier & Vaucher, 1999; Rego et al., 1999;
Gil de Pertierra, 2002, 2004, 2005; de Chambrier er al., 2005, 2006).

Proteocephalus sulcatus was described on the basis of tapeworms from two
unrelated fish hosts, a catfish and a bichir, because Klaptocz (1906) considered slight
differences between them, especially in the shape of the scolex, to be accounted for by
intraspecific variability. The type-host was not designated, although Khalil (1963 —
p. 309) considered saddled bichir Polypterus endlicheri to be the type-host because it
was listed first in the text of Klaptocz’ (1906) original description. However, the
following data indicate that widehead catfish Clarotes laticeps should be considered
the actual type-host of P. sulcatus: (i) the original description was mainly based on
larger tapeworms from widehead catfish Clarotes niloticus (see Klaptocz, 1906 —
p. 123); (ii) saddled bichir harboured less and smaller tapeworms (Klaptocz, 1906 —
p. 123); (111) Khalil (1963) did not find any P. sulcatus tapeworms in 322 P. endlicheri,
P. bichir Lacépède and P. senegalus Cuvier from the White Nile in the Sudan (see also
Jones, 1980); (iv) the present authors also did not find P. sulcatus in eight specimens
of Polypterus, mainly P. senegalus. On the other hand, eight out of nine widehead
catfish Clarotes laticeps examined in Kosti were infected.

On the basis of the above-listed facts, Clarotes laticeps is considered to be the
type- and most probably the only actual definitive host of P. sulcatus. Klaptocz’ (1906)
finding of P. sulcatus in saddled bichir may have represented an accidental infection
via predation of widehead catfish and the bichir served as a postcyclic host (see
Odening, 1976 for terminology of host categories). This latter assumption is supported
by the fact that saddled bichir is essentially piscivorous and that the infected P
endlicheri specimen was extremely large (total length 622 mm according to Klaptocz,
1906, thus reaching to the maximum standard length reported for the species, which is
630 mm — Froese & Pauly, 2007).

Klaptocz (1906) described another species of Proteocephalus, P. pentastoma
(syn. Ichthyotaenia pentastoma Klaptocz, 1906), from Nile bichir, Polypterus bichir
Lacépède, 1803, from the White Nile in the Sudan. This cestode is a specific and
relatively common parasite of bichirs in the Sudan (Jones, 1980), but it is markedly
different from P. sulcatus, especially in possessing a muscular apical sucker and
vitelline follicles arranged in a transverse band posterior to the ovary with anterior
longitudinal arms along the lateral margins of proglottides (Jones, 1980).

Proteocephalus sulcatus, which is considered here to be a specific parasite of
Clarotes niloticus, has hitherto been found only in the Nile River in the Sudan.
However, it probably also occurs in the Nile River basin in other countries of north-
eastern Africa.

REDESCRIPTION OF PROTEOCEPHALUS SULCATUS FROM SUDAN 701

ACKNOWLEDGEMENTS

The authors thank Dia-Eldin Elnaiem (Davis, California) for help in
organization of the stay of two of the present authors (A. de C. and T.S.) in the Sudan,
Andre Piuz for providing SEM photomicrographs, Janik Pralong for technical help and
Florence Marteau and Gilles Roth (all Geneva) for their help with drawings. Research
stay in the Sudan would not have been possible without the invaluable help of Ali
Adam and Sayed (University of Khartoum), Khalid Bashir Abaker and Ammar Osmar
(White Nile Fisheries Research Station in Kosti). The support of the Embassy of
Switzerland in Khartoum (Chargee d’Affaires Andrea Reichlin) is also acknowledged.
A. de C. is also deeply indebted to the “Donation Georges et Antoine Claraz” for
supporting this study. T.S. acknowledges financial support of the Grant Agency of the
Czech Republic (project No. 524/04/0342) and the Institute of Parasitology (projects
Nos. Z60220518 and LC 522); M.B. is grateful to Patrick Mugny, of the Département
de la Culture, Geneva City, for financial support to realize his research stay in
Switzerland in 2007.

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REVUE SUISSE DE ZOOLOGIE 114 (4): 703-719; décembre 2007

Description of Ambanus jaegeri sp. n. and of the male of A. euini
(Paik) from Korea (Arachnida: Araneae: Amaurobiidae)

Byung-Woo KIM
Department of Life Science, College of Natural Sciences, Hanyang University,
Seoul 133-791, Korea. Email: bwkim00@hotmail.com

Description of Ambanus jaegeri sp. n. and of the male of A. euini (Paik)
from Korea (Arachnida: Araneae: Amaurobiidae). - Two species of the
genus Ambanus from Korea are described and illustrated in detail. Ambanus
jaegeri Sp. n. and the male of Ambanus euini (Paik, 1976) are described for
the first time. The new species is distinguished by its male palp with
cymbial furrow longer than half the cymbium length, dorsal apophysis of
conductor elongated and bent distally, median apophysis small, semi-
circular, with sharp apical edge, and by its epigynum with widely triangular
atrium and broadly curved copulatory ducts with transparent membranes.

Keywords: Taxonomy - Coelotinae - new species - Korea.

INTRODUCTION

The Holarctic spider subfamily Coelotinae is one of the most common spider
taxa in the region from North America to East Asia. Wang (2002) revised the
Coelotinae at the generic level based on 31 characters and 22 taxa, together with two
outgroup taxa (Tamgrinia, Amaurobius). The subfamily comprises at least 373 species
(Platnick, 2007; Wang, 2002, 2007). Of these, the genus Ambanus Ovtchinikov, 1999
includes 18 species from several Asian countries (Korea, 10 species; Russia, 4; China,
3; Japan, 1). These taxa are characterized by the absence of a femoral apophysis, the
presence of dorsal apophysis of the conductor, a large epigynal atrium, and broadly
expanded, posteriorly originating copulatory ducts. More than nine species of
Ambanus were described only from male or female specimens and they have been
previously included in the genus Coelotes Blackwall, 1841 (Paik, 1974, 1976, 1978;
Song et al., 1993; Kim & Jung, 1993; Ovtchinikov, 1999). Although Namkung (2001,
2003) presented simple illustrations of nine Ambanus species (A. bifidus [Paik, 1976],
A. dimidiatus [Paik, 1974], A. euini [Paik, 1976], A. kayasanensis [Paik, 1972], A. kimi
[Paik, 1974], A. lunatus [Paik, 1976], A. ovatus [Paik, 1976], A. paikwunensis [Kim &
Jung, 1993], A. quadrativulvus [Paik, 1974]), these cannot be reliably identified from
his illustrations. Kim & Lee (2006) revised two poorly known species, A. lunatus and
A. coreana, and the latter was transferred to become the type species of the monotypic
genus Alloclubionoides Paik, 1992 (Clubionidae). Furthermore, the female paratype of
Ambanus paikwunensis and female specimens identified as A. lunatus from Korea are
in fact the females of A. coreana. Kim & Lee (2007) transferred A. kayasanensis to the

Manuscript accepted 03.08.2007

704 B.-W. KIM

genus Draconarius on the basis of several characters: The presence of a patellar apo-
physis, the hooked dorsal apophysis of the conductor, the position of the spermathecal
head, etc.

During a survey of the spider fauna of Korea spiders of the genus Ambanus
were collected by means of pitfall traps in natural forests and caves. In this paper the
male of Ambanus euini is described for the first time and Ambanus jaegeri is described
as a new species. The main goal of this paper is to provide data for a future revisional
study of the endemic Korean spider genus Ambanus.

MATERIAL AND METHODS

The Korean National Park of Mt Odae (KNPO) is situated in Pyungchang-gun
and Hongcheon-gun, Gangwon-do. Three collection sites (Woljeong temple, GPS: N
37°43’48”, E 128°35’43”; Sangwon temple, GPS: N 37°47’00”, E 128°34 10”; and
Maebong mountain peak, GPS: N 37°45°34.7”, E 128°42’57.4”) were chosen. All are
located beside small streams, with vegetation characterized by a mixture of dead trees,
giant fir trees (Abies holophylla) and broadleaf species (Quercus mongolica). At each
station two pitfall traps (plastic cups, height 6.3 cm, diameter 8 cm) were set 10 m apart
and filled with ethylene glycol (see Greenslade & Greenslade, 1971). Specimens were
preserved in 70% ethanol. Measurements of all parts are in millimeters unless noted
otherwise and are given for one specimen of each sex. Specimens examined in this
paper will be deposited in the National Institute of Biological Resources (NIBR), in the
collections of the Arachnological Institute of Korea (AIK), the Laboratory of Bio-
diversity, Hanyang University (LBHU) and in the Muséum d’histoire naturelle,
Genève (MHNG).

The descriptive terminology follows that of Kim & Lee (2006, 2007). Abbre-
viations: a, apical; AER, anterior eye row; ALE, anterior lateral eye; ALS, anterior
lateral spinneret; AME, anterior median eye; CDA, dorsal apophysis of conductor; d,
dorsal; ITA, intermediate tibial apophysis; KNPO, Korean National Park of Mt Odae;
p, prolateral; PER, posterior eye row; PLE, posterior lateral eye; PLS, posterior lateral
spinneret; PME, posterior median eye; PMS, posterior median spinneret; r, retrolateral;
RTA, retrolateral tibial apophysis; I, II, III, IV, first, second, third, forth legs.

TAXONOMY

Ambanus euini (Paik, 1976) Figs 1-3, 7A-C

Coelotes euini Paik, 1976: 78, figs 6-8 (description of female); Paik 1978: 341, fig. 151.1-2.
Ambanus euini: Ovtchinnikov 1999: 64 (transferred from Coelotes).

MATERIAL EXAMINED: 1 female, 1 male (AIK), 3 November, 1984, Mt Yebong,
Gyeonggi-do, leg. K.S. Lee; 1 female (LBHU), 13 October, 2001, Mt Gyebang, Gangwon-do,
leg. T.S. Kwon; 1 male, 1 female (NIBR), 1 May, 2005, Woljeong temple, 1 male (NIBR), 10
June, 2005, Gwangmijang, 55 males, 3 females (LBHU), 11 June, 2005, Sangwon temple, 100
males, 12 females (LBHU), 11 June, 2005, Mabong mountain peak, 5 males (NIBR), 22 July,
2005, Woljeong temple, 2 males (NIBR), 4 September, 2005, Woljeong temple, female (NIBR),
28 September, 2005, Sangwon temple, 10 males, 7 females (LBHU), 11 October, 2005,
Woljeong temple, 2 females, 2 males (MHNG), 8 November, 2005, Woljeong teple, KNPO,
Gangwon-do, leg. B.W. Kim.

KOREAN AMBANUS SPIDERS 705

DIAGNOSIS: This species is similar to A. jaegeri sp. n., A. ovatus and A. quadra-
tivulvus in having the epigynal atrium very broadly oval and situated posteriorly near
the epigastric furrow; atrial septum and atrial hood absent or indistinct; epigynal teeth
absent; copulatory ducts broadly curved with transparent membranes on both lateral
margins; male palpal organ with large embolus with curved distal part; dorsal apoph-
ysis of conductor present; conductor hook-like, with a rounded distal end. The female
of Ambanus euini can be distinguished by its long oval genital atrium expanded to both
spermathecal stalks, 3.5 times as wide as long (length 0.2 mm, width 0.7 mm); copu-
latory ducts broadly curved, with transparent membranes, overlapped slightly for one
third of the outline of the longest duct. Males are distinguished by patellar apophysis
being very small; cymbial furrow (1.1 mm) shorter than half of cymbium length (2.5
mm); dorsal apophysis of conductor horn-like, expanded to conductor and situated on
the side of conductor; median apophysis developed as a small thin semicircular pro-
jection.

MEASUREMENTS: Male (female in parentheses): Body length 9.3 (11.5); cheli-
cera length 2.3 (2.9), chelicera width 1.2 (1.6), cheliceral fang length 1.2 (1.5); clypeus
height 0.5 (0.4); carapace length 4.9 (5.9), carapace width 3.4 (3.6), carapace height 2.6
(2.9); maxillae length 1.6 (1.8), maxillae width 0.8 (1.0); labium length 0.8 (1.0),
labium width 0.7 (0.9); sternum length 2.5 (2.7), sternum width 2.0 (2.2); AER 0.9
(1.0), PER 1.2 (1.4), AME 0.2 (0.1), ALE 0.3 (0.3), PME 0.2 (0.2), PLE 0.2 (0.3). Eye
formula ALE>PLE=PME=AME (ALE=PLE>PME>AME). Palp 5.3 (5.6) [1.7 (1.9),
02089) 0:4. (lal) s 2259-8) ], Firstlesd3 0432) 3:B731:0.1.7,29:E:0)341
(C0) 5281128); second leg -12.2 (12:2), [3:4 (3.4)s 1:6241-8);, 2:5 (2:5) 3:01 (9) 17
RO) |eithirdlegs A6) 262) 15416) 21001)32 G.1) 7 (4/6)]Mourth
leg 15.4 (15.4) [4.0 (4.1), 1.6 (1.7), 3.3 (3.3), 4.5 (4.4), 2.0 (1.9)]. Leg formula IV I II
IH (IV III III). Abdomen length 4.7 (6.0), abdomen width 2.9 (3.8), abdomen height
PAGES).

DESCRIPTION OF MALE (from Woljeong temple, KNPO, 1 May, 2005 [NIBR]):
Medium-sized spider, shorter than female. Carapace elongate, 1.4 times as long as
wide, moderately narrowed in thorax area, with distinctly longitudinal fovea (Fig. 1A).
AER straight and PER slightly procurved in frontal view; ALE larger than other eyes,
AME separated by slightly less than their diameter, and longest eye row width to cara-
pace width ratio 35 (Fig. 1A). Clypeus height 2.5 times as long as AME diameter; a
pair of eyebrow-shaped chila present (Fig. 1B). Chelicerae with numerous long setae;
lateral condyle yellowish brown; three promarginal teeth on groove, middle one
largest, and two retromarginal teeth of subequal size (Fig. 1C). Maxillae reddish
brown, widest at mid-part. Labium rectangular, slightly longer than wide (Fig. 1D).
Sternum shield-shaped, widest between second coxae, not produced between fourth
coxae (Fig. 1E). Palp, see Figure 1G-J; tibia with 12 trichobothria in three rows
(Sd-6d-2r), tarsus with five trichobothria in one row (5d); femur with three spines, tibia
with four spines (one, 1-0 dorsally; three, 2-1 prolaterally), tarsus with eight spines
(one, 1-0-0 dorsally; three, 1-1-1 prolaterally; two, 0-2 retrolaterally; two, 0-2 ven-
trally). Legs (Fig. 1F) yellowish brown; patella + tibia of first leg always shorter than
carapace length; trochanters not notched; tibiae with 20-23 trichobothria in four rows

706 B.-W. KIM

FIG. 1

Ambanus euini (Paik, 1976), male (NIBR; 1 May, 2005) from Woljeong temple, KNPO. (A)
Habitus, dorsal view. (B) Eye area and clypeus, frontal view. (C) Chelicerae, posterior view. (D)
Maxillae and labium, ventral view. (E) Sternum, ventral view. (F) Left leg IV, prolateral view.
(G-J) Left palp, prolateral view (G), ventral view (H), retrolateralview (I), dorsal view (J).

Note: CDA, dorsal apophysis of conductor; CF, cymbial furrow; CO, conductor; EM, embolus;

ITA, intermediate tibial apophysis; MA, median apophysis; PA, patellar apophysis; RTA, retro-
lateral tibial apophysis.

KOREAN AMBANUS SPIDERS 707

(5p-6d-5d-6r on first leg, 5p-6d-6d-6r on second, 4p-6d-5d-Sr on third, 5p-6d-6d-6r on
fourth), metatarsi with seven to eight trichobothria in one row (eight on first and fourth
leg, seven on second and third leg), tarsi with eight to nine trichobothria in one row
(nine on first and fourth leg, eight on second and third leg); tarsal organ situated close
to distal end of tarsus, slightly anteriorly part of distal trichobothrium; tarsi with three
claws, upper claws with 9-13 teeth (13 on first leg, 12 on second, nine on third, 10 on
fourth), lower claw with zero to one tooth (zero on first, second and third leg, one on
fourth). Leg spination (see Table 1): Leg I: Femur with four spines, tibia with seven
spines (one, 0-0-1 prolaterally; six, 2-2-2a ventrally), metatarsus with seven spines
(one, 0-0-1 prolaterally; six, 2-2-2 ventrally), tarsus without spine; leg II: Femur with
four spines, tibia with five spines (one, 0-0-1 prolaterally; four, 1-1-2a ventrally),
metatarsus with nine spines (three, 1-0-2 prolaterally; six, 2-2-2 ventrally), tarsus
without spine; leg III: Femur with six spines, tibia with 10 spines (four, 1-1 prolaterally
and retrolaterally; six, 2-2-2a ventrally), metatarsus with 16 spines (ten, 1-2-2 pro-
laterally and retrolaterally; six, 2-2-2 ventrally), tarsus with three spines (two, 0-1-1
prolaterally; one, 0-1-0 retrolaterally); leg IV: Femur with five spines, tibia with 11
spines (one, 1-0-0 dorsally; four, 1-1 prolaterally and retrolaterally; six, 2-2-2a
ventrally), metatarsus with 17 spines (one, 1-0-0 dorsally; ten, 1-2-2 prolaterally and
retrolaterally; six, 2-2-2 ventrally), tarsus with six spines (six, 0-1-1 prolaterally, retro-
laterally and ventrally). Abdomen ovoid, with scattered brownish yellow spots and
chevrons on dorsal side (Fig. 1A). Cribellum absent.

Palp (Figs 1G-J, 7A-B): Patellar apophysis very small; RTA modified with
small ITA; cymbial furrow (1.1 mm) shorter than half the cymbium length (2.5 mm);
tegular sclerite weakly sclerotized and situated longitudinally on tegulum; conductor
broadly hook-like, bent in a clockwise direction (left palp), its distal end rounded;
dorsal apophysis of conductor horn-like, expanded towards and facing the conductor;
embolus long slender, broadly wound clockwise (left palp) and penetrating into the
middle of the tegulum; median apophysis a small thin semicirclar projection.

TABLE 1. Spination of leg segments in Ambanus euini (Paik, 1976) from Korea. Male (female in
parentheses). Note: a, apical part; metat., metatarsus; [ ], according to original description by
Paik (1976).

dorsal ventral prolateral retrolateral
Ist leg femur 110 (1 1) 0 (0) 002 (002) 0 (0)
tibia 0 (0) 222 (222a) 001 (001) 0 (0)
metat. 0 (0) 2221222) 001 (011) 0 (0)
tarsus 0 (0) 0 (0) 0 (0) 0 (0)
2ndleg femur 11(11) 0 (0) 011 (011) 0 (0)
tibia 0 (0) 112a (222a) 001 (011) 0 (0)
metat. 0 (0) 222 (222) 102 (012) 0 (010)
tarsus 0 (0) 0 (0) 0 (0) 0 (0)
3rd leg femur 121 (121) 0 (0) JA (LU) 0 (0)
tibia 0 (0 [1 1]) 222a (222a) 11(11) 11(01[11])
metat. 0 (0) 222 (222) 122 (122) 122 (122)
tarsus 0 (0) 0 (0) O11 (011) 010 (010)
4th leg femur 111 (111) 0 (0) 101 (1 1) 0 (0)
tibia 100 (101) 222a (222a) 11(11) 11(11)
metat. 100 (110) 222 (222) 122 (112) 222 (122)

708 B.-W. KIM

DESCRIPTION OF FEMALE (from Woljeong temple, KNPO, 1 May, 2005 [NIBR]):
Medium-sized spider, longer than male. Carapace elongate, 1.6 times longer than wide,
moderately narrowed in thorax area, with distinctly longitudinal fovea on middle (Fig.
2A). AER almost straight and PER slightly procurved in frontal view; AME smaller
than other eyes, separated by as much as their diameter, and longest eye row width to
carapace width ratio 28 (Fig. 2A). Clypeus height four times as long as AME diameter;
distinct chilum present (Fig. 2B). Chelicerae with numerous long setae; lateral condyle
yellowish brown; with three promarginal teeth on groove, middle one largest, and two
retromarginal teeth of subequal size (Fig. 2C). Maxillae reddish brown, widest at mid-
part. Labium rectangular, slightly longer than wide (Fig. 2D). Sternum shield-shaped,
widest between second coxae, 1.2 times as long as wide and slightly projecting
between 4th coxae (Fig. 2E). Palp: Claw with seven teeth; tibia with 15 trichobothria
in three rows (6d-6d-3r), tarsus with six trichobothria in one row (5d); femur with three
spines, tibia with five spines (two, 1-1 dorsally; three, 0-1-2 prolaterally), tarsus with
16 spines (one, 1-0 dorsally, six, 3-2-1 prolaterally; five, 2-2-1 retrolaterally; four,
0-0-4 ventrally). Legs (Fig. 2F) yellowish brown, without ring patterns; patella + tibia
of first leg always shorter than carapace length; trochanters not notched; tibiae with
22-25 trichobothria in four rows (6p-6d-6d-7r on first leg, 6p-6d-6d-6r on second,
5p-6d-6d-5r on third, 5p-7d-6d-6r on fourth), metatarsi with seven to nine trichobothria
in one row (seven on first leg, eight on second and third, nine on fourth), tarsi with
eight to 10 trichobothria in one row (nine on first and second leg, eight on third, 10 on
fourth); tarsal organ situated close to distal end of each leg tarsus, slightly anteriorly of
distal trichobothrium; tarsı with three claws, upper claws with 9-12 teeth (12 on first
leg, 12 on second, 10 on third and nine fourth), lower claw with one to two (one on first
and second leg, two on third and fourth leg). Leg spination (see Table 1): Leg I: Femur
with three spines, tibia with seven spines, one small spine on inner ventral side half as
long as others (one, 0-0-1 prolaterally; six, 2-2-2a ventrally), metatarsus eight (two,
0-1-1 prolaterally; six, 2-2-2 ventrally), tarsus without spine; leg II: Femur with four
spines, tibia with eight spines, one small spine on inner ventral side half as long as oth-
ers (two, 0-1-1 prolaterally; six, 2-2-2a ventrally), metatarsus with 10 spines (three, 0-
1-2 prolaterally; one, 0-1-0 retrolaterally; six, 2-2-2 ventrally), tarsus without spine; leg
III: Femur with six spines, tibia with nine spines (two, 1-1 prolaterally; one, 0-1 retro-
laterally; six, 2-2-2a ventrally), metatarsus with 16 spines (ten, 1-2-2 prolaterally and
retrolaterally; six, 2-2-2 ventrally), tarsus with three spines (two, 0-1-1 prolaterally;
one, 0-1-0 retrolaterally); leg IV: Femur with five spines, tibia with 12 spines, two slen-
der spines on dorsal (two, 1-0-1 dorsally; four, 1-1 prolaterally and retrolaterally; six,
2-2-2a ventrally), metatarsus with 17 spines, two spines on dorsal side (two, 1-1-0 dor-
sally; four, 1-1-2 prolaterally; five, 1-2-2 retrolaterally; six, 2-2-2 ventrally), tarsus
with six spines (six, 0-1-1 prolaterally, retrolaterally and ventrally respectively).
Abdomen ovoid, with scattered brownish yellow spots and chevrons on dorsal side
(Fig. 2A). Cribellum absent.

Epigynum (Figs 2G, H, 3A-B, 7C): Epigynal teeth absent; epigynal opening
long oval, expanded to both spermathecal stalks, 3.5 times as wide as long (length 0.2
mm, width 0.7 mm); atrial septum indistinct, originating on posterior plate; copulato-
ry pores deep, round on both sides; without atrial hood; copulatory ducts broadly

KOREAN AMBANUS SPIDERS 709

Fic. 2
Ambanus euini (Paik, 1976), female (NIBR; 1 May 2005) from Woljeong temple, KNPO. (A)
Habitus, dorsal view. (B) Eye area and clypeus, frontal view. (C) Chelicerae, posterior view. (D)
Maxillae and labium, ventral view. (E) Sternum, ventral view. (F) Left leg IV, prolateral view.
(G) Epigynum, ventral view. (H) Corse of copulatory duct, right part, ventral view.

710 B.-W. KIM

FIG. 3
Ambanus euini (Paik, 1976), female (NIBR; 1 May 2005) from Woljeong temple, KNPO. (A)
Epigynum, ventral view. (B) Vulva, dorsal view.

Note: CD, copulatory duct; EF, epigastric furrow; FD, fertilization duct; GO, genital opening;
SH, spermathecal head; SS, spermathecal stalk.

KOREAN AMBANUS SPIDERS Zan

curved, with transparent membranes, slightly overlapped for one third of the outline of
the longest duct (from genital opening to upper returning part); spermathecal heads
developed as small cylindrical processes situated at anterior end of spermathecae;
spermathecae large, curved inwards, with distinct stalks and bases: fertilization ducts
small, arising from the posterior ends of the spermathecae.

DISTRIBUTION: Korea (Mt Gyebang, Mt Odae, Mt Yebong).

REMARKS: The female of A. euini collected from Mt Odae is redecribed with a
multitude of characters (leg spination, trichobothrial patterns, etc.) neglected in the
original description. The specimens examined were found wandering on the ground
among stones and leaf litter.

Ambanus jaegeri sp. n. Figs 4-6, 7D-F

Ambanus euini: Namkung, 2002: 398, fig. 28.12a-b (description of female), 2003: 400, fig.
28.12a-b; Kim and Cho, 2002: 176, photos 355-360 (description of male and female).
Misidentification.

MATERIAL EXAMINED: Male holotype (NIBR), 11 June, 2005, Sangwon temple, KNPO,
Gangwon-do, leg. B.W. Kim. Paratypes: 1 female (NIBR), 11 June, 2005, Sangwon temple,
KNPO, Gangwon-do; | male (AIK), 23 August, 2003, Mt Samak, Gangwon-do, B.W. Kim;
1 female (MHNG), 1 male (MHNG), 11 males (LBHU), 2 females (LBHU), 11 June, 2005,
Sangwon temple, 1 male (LBHU), 11 June, 2005, Woljeong temple, 1 female (LBHU), 11 June,
2005, Maebong mountain peak, 1 female (LBHU), 3 September 2005, Dongpi valley, 1 female
(AIK), 4 September, 2005, Sangwon temple, KNPO, Gangwon-do, leg. B.W. Kim.

ETYMOLOGY: The specific name is a patronym in honor of Dr Peter Jäger, the
German arachnologist who supported the early stages of my taxonomic studies with

dedicated assistances and encouragement.

DIAGNOsIs: This species is similar to A. euini, A. ovatus and A. quadrativulvus
in having the epigynal atrium very broadly oval, situated posteriorly near the epigastric
furrow; atrial septum and atrial hood absent or indistinct; epigynal teeth absent; copu-
latory ducts broadly curved, with transparent membranes on both lateral margins; male
palpal organ with large embolus with curved distal part; dorsal apophysis of conductor
present; conductor hook-like, with a rounded distal end situated in centre of papal
organ. The female of Ambanus jaegeri sp. n. can be distinguished by its trianglar epi-
gynal atrium which is heart-shaped and two times as wide as long (length 0.3 mm,
width 0.6 mm); copulatory ducts broadly curved, with transparent membranes, slightly
overlapped for half of the outline of the longest duct. Males are distinguished by
patellar apophysis absent; cymbial furrow (1.6 mm) longer than half the cymbium
length (2.8 mm); dorsal apophysis of conductor slender, horn-like, expanded, curving
towards top of cymbium and facing the conductor; median apophysis developed as a
small, thin, semicirclar projection with sharp apical part.

MEASUREMENTS: Male (female in parentheses): Body length 9.4 (11.2);
chelicera length 2.6 (2.7), chelicera width 1.2 (1.3), cheliceral fang length 1.4 (1.4);
clypeus height 0.3 (0.3); carapace length 5.5 (5.3), carapace width 3.6 (3.4), carapace
height 1.9 (1.7); maxillae length 1.8 (1.6), maxillae width 0.8 (0.9); labium length 1.0
(0.9), labium width 0.8 (0.8); sternum length 2.7 (2.6), sternum width 2.0 (2.1); AER
0.8 (0.9), PER 1.2 (1.3), AME 0.1 (0.1), ALE 0.2 (0.2), PME 0.1 (0.2), PLE 0.2 (0.2).

712 B.-W. KIM

TABLE 2. Spination of leg segments of Ambanus jaegeri sp. n. from Korea. Male (female in
parentheses). Note: a, apical part; metat., metatarsus.

dorsal ventral prolateral retrolateral
Ist leg femur 110 (2 1) 0 (0) 002 (002) 0 (0)
tibia 0 (0) 222a (222a) 001 (011) 0 (0)
metat. 0 (0) 222 (222) 011 (101) 0 (001)
tarsus 0 (0) 0 (0) 0 (0) 0 (0)
2nd leg femur 110 (110) 0 (0) 1 1(011) 0 (0)
tibia 0 (0) 222a (112a) 011 (011) 0 (0)
metat. 0 (0) 222 (222) 012 (012) 0 (011)
tarsus 0 (0) 0 (100) 0 (0) 0 (0)
3rd leg femur 122 (122) 0 (0) 101 (101) 0 (0)
tibia 0 (100) 222 (2224) 11(11) KON)
metat. 0 (0) 222222) 122 (122) 122 (122)
tarsus 0 (0) 010 (010) 011 (011) 010 (010)
4th leg femur 112 (112) 0 (0) 10 (0) 0 (0)
tibia 0 (0) 222 (222a) 11(11) 11(11)
metat. 100 (0) 2222222) 1227122) 122 (122)
tarsus 0 (0) 010 (010) 011 (013) 011 (011)

Eye formula ALE=PLE>PME=AME (ALE=PLE>PME>AME). Palp 5.8 (5.1) [1.9
(427), 0.7 (0:7); 04 (10), 2.8 (1:71: First leg 15.5 (12.1) [4.3 6:35) 1710) 8335928)
3:81(2:8), 2.2 (1.6)], second leg 142 11.3) BI @2), 17 LO) ZI CHIBIOCHZA
(1.6)], third leg 13.3 (10.9) [3.5 (3.0), 1.7 (1.4), 2.4 (1.9), 3.7 (2.9), 2.0 (1.5)], fourth
leg 17.4 (14.4) [4.6 (3.5), 1.7 (1.7), 3.6 (3.1), 5.2 (4.2), 2.3 (1.9)]. Leg formula IV I II
IH (IV III III). Abdomen length 4.9 (6.5), abdomen width 2.9 (4.1), abdomen height
2.8 (3.9).

DESCRIPTION OF MALE (holotype): Medium-sized spider, shorter than female.
Carapace elongate, 1.3 times as long as wide, moderately narrowed in thorax area, with
distinctly longitudinal fovea (Fig. 4A). AER straight and PER slightly procurved in
frontal view; AME smaller than other eyes, separated by slightly less than their
diameter, longest eye row width to carapace width ratio 33 (Fig. 4B). Clypeus height
three times as long as AME diameter; a pair of eyebrow-shaped chila present (Fig. 4B).
Chelicerae with numerous long setae; lateral condyle yellowish brown; three pro-
marginal teeth on groove, middle one largest, and two retromarginal teeth of subequal
size (Fig. 4C). Maxillae reddish brown, widest at mid-part. Labium rectangular,
slightly longer than wide (Fig. 4D). Sternum shield-shaped, widest between second
coxae, not produced between fourth coxae (Fig. 4E). Palp, see Figure 4G-J; tibia with
14 trichobothria in three rows (5d-3d-6r), tarsus with five trichobothria in one row (5d)
and femur with three spines, tibia with three spines (1-2 prolaterally), tarsus with four
spines (four, 0-0-0-2 prolaterally and retrolaterally). Legs (Fig. 4F) yellowish brown;
patella + tibia of first leg always shorter than carapace length; trochanters not notched;
tibia with 20-24 trichobothria in four rows (Sp-6d-5d-6r on first leg, 6p-6d-6d-6r on
second, 4p-6d-5d-5r on third, 5p-7d-6d-4r on fourth), metatarsi with seven to nine
trichobothria in one row (nine on first and fourth leg, eight on second, seven on third),
tarsı with seven to nine trichobothria in one row (nine on first, third and fourth leg,
seven on second); tarsal organ situated close to distal end of tarsus, slightly anteriorly

KOREAN AMBANUS SPIDERS

713

Fic. 4

Ambanus jaegeri sp. n., male holotype. (A) Habitus, dorsal view. (B) Eye area and clypeus,
frontal view. (C) Chelicerae, posterior view. (D) Maxillae and labium, ventral view. (E) Sternum,
ventral view. (F) Left leg IV, prolateral view. (G-J) Left palp, prolateral view (G), ventral view
(H), retrolateralview (I), dorsal view (J). Note: CDA, dorsal apophysis of conductor; CF, cymbial

furrow; CO, conductor, EM, embolus; ITA, intermediate tibial apophysis; MA, median
apophysis; RTA, retrolateral tibial apophysis.

714 B.-W. KIM

of distal trichobothrium; tarsi with three claws, upper claws with 10-13 teeth (13 on
first and second leg, 12 on third, 10 on fourth), lower claw without tooth. Leg spination
(see Table 1): Leg I: Femur with four spines, tibia with seven spines, one small spine
on inner ventral side half as long as others (one, 0-0-1 prolaterally; six, 2-2-2a
ventrally), metatarsus with eight spines (one, 0-1-1 prolaterally; six, 2-2-2 ventrally),
tarsus without spine; leg II: Femur with four spines, tibia with eight spines, one small
spine on inner ventral and median prolateral side half as long as others (two, 0-1-1 pro-
laterally; four, 2-2-2a ventrally), metatarsus with nine spines (three, 0-1-2 prolaterally;
six, 2-2-2 ventrally), tarsus without spine; leg III: Femur with seven spines, tibia with
10 spines (four, 1-1 prolaterally and retrolaterally; six, 2-2-2a ventrally), metatarsus
with 16 spines (ten, 1-2-2 prolaterally and retrolaterally; six, 2-2-2 ventrally), tarsus
with four spines (two, 0-1-1 prolaterally; two, 0-1-0 retrolaterally and ventrally); leg
IV: Femur with five spines, tibia with 10 spines (four, 1-1 prolaterally and retro-
laterally; six, 2-2-2 ventrally), metatarsus with 17 spines (one, 1-0-0 dorsally; ten,
1-2-2 prolaterally and retrolaterally; six, 2-2-2 ventrally), tarsus with five spines (four,
0-1-1 prolaterally and retrolaterally; one, 0-1-0 ventrally). Abdomen ovoid, with
scattered brownish yellow spots and without distinct chevrons on dorsal side (Fig. 4A).
Cribellum absent.

Palp (Figs 4G-J, 7D-E): Patellar apophysis absent; RTA modified with ITA;
cymbial furrow (1.6 mm) longer than half the cymbium length (2.8 mm); tegular
sclerite weakly sclerotized and situated longitudinally on tegulum; conductor broadly
hook-like, bent in a clockwise direction (left palp), distal end rounded; dorsal apo-
physis of conductor slender, horn-like, expanded towards top of cymbium and facing
the conductor; embolus long, slender, broadly wound clockwise (left palp) and
penetrating the middle of the tegulum; median apophysis a small thin semicircular
projection with sharp apical part.

DESCRIPTION OF FEMALE (paratype from Sangwon temple, KNPO,11 June, 2005
[NIBR]): Medium-sized spider, longer than male. Carapace elongate, 1.4 times longer
than wide, moderately narrowed in thorax area, with distinctly longitudinal fovea
(Fig. 5A). AER straight and PER slightly procurved in frontal view; AME smaller than
other eyes, separated by as much as their diameter, and longest eye row width to cara-
pace width ratio 38 (Fig. 5B). Clypeus height three times as long as AME diameter; a
pair of eyebrow-shaped chila present (Fig. 5B). Chelicerae with numerous long setae;
lateral condyle yellowish brown; three promarginal teeth on groove, middle one
largest, and two retromarginal teeth of subequal size (Fig. SC). Maxillae reddish
brown, widest at mid-part. Labium rectangular, 1.1 times as long as wide (Fig. 5D).
Sternum shield-shaped, widest between second coxae, 1.2 times as long as wide, and
slightly projecting between 4th coxae (Fig. 5E). Palp: Claw with seven teeth; tibia with
15 trichobothria in three rows (6d-6d-3r), tarsus with seven trichobothria in one row
(7r); femur with three spines, tibia with five spines (three, 1-2 prolaterally; two, 1-1
retrolaterally), tarsus with 14 spines (one, 1-0-0 dorsally; six, 3-2-1 prolaterally; five,
2-2-1 retrolaterally; four, 0-0-4 ventrally). Legs (Fig. 5F) yellowish brown, without
annulation; patella + tibia of first leg always shorter than carapace length; trochanters
not notched; tibia with 22-25 trichobothria in four rows (5p-2-3d-6r on first leg, Sp-6d-
5d-6r on second, 4p-6d-5d-Sr on third, 5p-4d-5d-5r on fourth), metatarsi with six to

KOREAN AMBANUS SPIDERS TANS

Fic. 5
Ambanus jaegeri sp. n., female paratype (NIBR; 11 June, 2005) from Sangwon temple, KNPO.
(A) Habitus, dorsal view. (B) Eye area and clypeus, frontal view. (C) Chelicerae, posterior view.
(D) Maxillae and labium, ventral view. (E) Sternum, ventral view. (F) Left leg IV, prolateral
view. (G) Epigynum, ventral view. (H) Course of copulatory duct, right part, ventral view.

716 B.-W. KIM

BES, À
i Sat

Fic. 6

Ambanus jaegeri sp. n., female paratype (NIBR; 11 June, 2005) from Sangwon temple, KNPO.
(A) Epigynum, ventral view. (B) Vulva, dorsal view.

Note: CD, copulatory duct; EF, epigastric furrow; FD, fertilization duct; GO, genital opening;
SH, spermathecal head; SS, spermathecal stalk.

KOREAN AMBANUS SPIDERS TA

pitti,

Inga @9X near ED

“aber ERW.

FIG. 7

SEM photographs of Ambanus euini (Paik, 1976) (A-C) and A. jaegeri sp. n. (D-F) from Korea.
(A, D) Left male palp, retrolateral view. B, E. Median apophysis of left palpal organ, retrolater-
al view. (C, F) Epigynum, ventral view.

eight trichobothria in one row (eight on first, second and fourth leg, six on third), tarsi
with seven to nine trichobothria in one row (nine on first and fourth leg, eight on
second, seven on third); tarsal organ situated close to distal end of tarsus, slightly
anteriorly of distal trichobothrium; tarsi with three claws, upper claws with 9-12 teeth
(12 on first and second leg, 10 on third, nine on fourth), lower claw with two teeth (on
all legs). Leg spination (see Table 2): Leg I: Femur with five spines, tibia with eight

718 B.-W. KIM

spines, one small spine on inner ventral side half as long as others (two, 0-1-1 pro-
laterally; six, 2-2-2a ventrally), metatarsus with nine spines (two, 1-0-1 prolaterally;
one, 0-0-1 retrolaterally; six, 2-2-2 ventrally), tarsus without spine; leg II: Femur with
four spines, tibia with six spines (two, 0-1-1 prolaterally; four, 1-1-2a ventrally),
metatarsus with nine spines (three, 0-1-2 prolaterally; six, 2-2-2 ventrally), tarsus
without spine; leg III: Femur with seven spines, tibia with 11 spines (one, 1-0-0
dorsally; two, 1-1 prolaterally; two, 0-1-1 retrolaterally; six, 2-2-2a ventrally), meta-
tarsus with 16 spines (ten, 1-2-2 prolaterally and retrolaterally; six, 2-2-2 ventrally),
tarsus with four spines (two, 0-1-1 prolaterally; one, 0-1-0 retrolaterally; one, 0-1-0
ventrally); leg IV: Femur with four spines, tibia with 10 spines (four, 1-1 prolaterally
and retrolaterally; six, 2-2-2a ventrally), metatarsus with 16 spines (10, 1-2-2 pro-
laterally and retrolaterally; six, 2-2-2 ventrally), tarsus with seven spines (four, 0-1-3
prolaterally; two, 0-1-1 retrolaterally; one, 0-1-0 ventrally). Abdomen ovoid, with
scattered brownish yellow spots and without distinct chevrons on dorsal side (Fig. 5A).
Cribellum absent.

Epigynum (Figs 5G, H, 6A-B, 7F): Epigynal teeth absent; genital opening
heart-shaped, two times as wide as long (length 0.3 mm, width 0.6 mm); atrial septum
indistinct, originating on posterior plate; copulatory pores deep, round on both sides;
atrial hood absent; copulatory ducts broadly curved, with transparent membranes,
slightly overlapped for half of the outline of the longest duct (from genital opening to
upper returning part); spermathecal heads developed as small cylindrical processes
situated at anterior end of spermathecae; spermathecae large, curved inwards, with
distinct stalks and bases; fertilization ducts small, arising from the posterior ends of the
spermathecae.

DISTRIBUTION: Korea (Mt Odae, Mt Samak).

REMARKS: The specimens examined were found wandering on the ground
among stones and leaf litter.

ACKNOWLEDGEMENTS

The author wishes to express his sincere thanks to Prof. W. Lee of Hanyang
University, Seoul and Prof. J. P. Kim of Dongguk University, Seoul, to Dr D. X. Song
of Hebei University, China, to Dr X. Xu of the Chinese Academy of Sciences, to Dr
Y. M. Marusik of the Russian Academy of Sciences, to Dr X. P. Wang of the University
of Florida, U.S.A, to Dr P. Jager of the Senckenberg Museum, Germany, and to Dr
P. J. Schwendinger of the Muséum d’histoire naturelle, Genève for many valuable
comments and for providing several important papers. This research was financially
supported by the Korea Research Foundation Grant (KRF-2006-005-J01901).

REFERENCES

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of Bioscience and Biotechnology, Daejeon. 424 pp. (In Korean)

Kim, J. P. & Jung, C. H. 1993. A new species of the genus Coelotes (Araneae: Agelenidae) from
Korea. Korean Arachnology 9: 1-6.

KOREAN AMBANUS SPIDERS 719

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PAIK, K. Y. 1978. Illustrated Flora and Fauna of Korea. Volume 21 (Araneae). Samwha Press,
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PAIK, K. Y. 1992. A new genus of the family Clubionidae (Arachnida, Araneae) from Korea.
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SONG, D. X., ZHU, M. S., Gao, S. S. & GUAN, J. D. 1993. On new species of the genera Coelotes
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WANG, X. P. 2002. A generic-level revision of the spider subfamily Coelotinae (Araneae,
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REVUE SUISSE DE ZOOLOGIE 114 (4): 721-728; décembre 2007

Some new earthworms of the genus Amynthas (Oligochaeta:
Megascolecidae) with male discs from Bogildo Island, Korea

Yong HONG
Institute of Agricultural Science, College of Agriculture, Sangju National University,
Sangju 742-711, Republic of Korea. Email: geoworm@hanmail.net

Some new earthworms of the genus Amynthas (Oligochaeta: Mega-
scolecidae) with male discs from Bogildo Island, Korea. - Three new
Amynthas are described from Bogildo Island, Korea: Amynthas angulatus
sp. nov., Amynthas jamesi sp. nov., and Amynthas yunseondoi sp. nov.
Amynthas angulatus sp. nov. have two pairs of spermathecal pores interseg-
mental in 5/6, and 6/7. The other 2 species have three pairs of spermathecae
in VI, VII, and VII, the former with spermathecal pores intersegmental in
5/6, 6/7, and 7/8, the latter intrasegmental in VI, VII, and VIII. These
species have disc-shaped male pore region and simple intestinal caeca.
Descriptions of the new species are provided.

Keywords: Earthworms - Amynthas - Megascolecidae - Oligochaeta -
Korea - taxonomy.

INTRODUCTION

Amynthas has been long known to have more species than any other genus of
the Pheretima-complex (Sims & Easton, 1972). Korean Megascolecidae also belong to
Amynthas and many new species have been described recently further expanding this
large genus (Hong & James, 200 1ab; Hong & Lee, 2001; Hong et al., 2001). Beginning
with Kobayashi (1934), Korean earthworms have been studied repeatedly, but the work
is not yet completed since much area remains to be collected. There is little overlap
between the area sampled for this paper and areas covered by previous studies of
Korean Amynthas. In all probability, other mountains and other islands have diverse
faunas, thus the number of endemic species known in Korea will increase even more.

Genital papillae of the male pore region, especially the male disc, are useful for
classification of Amynthas and have been used throughout the history of the genus.
However, not many species have male discs, a particularly large form of genital
papillae or male porophore. Species with male discs are also represented by small
numbers of individuals. In this paper, I describe 3 species with male discs: Amynthas
angulatus sp. nov., Amynthas jamesi sp. nov., and Amynthas yunseondoi sp. nov.
Material was found from litter layers and soils in forests by hand sorting.

The Bogildo Island, one of the six major counties, is located at the southern end
of Wando-gun. Among its big mountains are Jeogja peak (435 m), Gwangdae peak

Manuscript accepted 14.09.2007

722 Y. HONG

(311 m), and Mangwol peak (350 m), while most areas have an altitude less than 300
m. The mean temperature during the coldest winter month January was 0.5 °C. Summer
temperatures reach a mean of 28.8 °C in August and thereafter decrease. The annual
mean precipitation is 1456.8 mm (by Korea Meteorological Administration). Holotype
and paratypes are deposited in the Korean Bioresources Collection (KB), National
Institute of Biologocal Resources.

DESCRIPTIONS
Amynthas angulatus sp. nov. Figs 1A-C

MATERIAL: Holotype: Clitellate specimen: Korea, Jeollanam-do province, Wando-gun,
Bogildo Isl., litter layers in forest, 19 July 2000, Y. Hong coll. Paratype: one clitellate: Same da-
ta as for holotype. Other material: Same data as for holotype, 1 aclitellate specimen.

ETYMOLOGY: The name angulatus is Latin for “angular” here referring to the
shape of the genital papillae.

DIAGNOSIS: Paired spermathecal pores in 5/6, 6/7; male pores XVIII at posterior
corner of large angular papillae, each pad extending to 17/18 and 18/19, each papilla
hardened.

DESCRIPTION: Worm unpigmented. Dimensions 86 (aclitellates 71-86) mm by
4.5 mm at segment X, 5.2 mm at XXX, 4.7 mm at clitellum; body cylindrical
throughout, segments 85. Setae regularly distributed around segmental equators,
numbering 71 at VII, 92 at XX; 2-3 between male pores; setal formula AA:AB: YZ:ZZ
= 1:1:1.5:2 at XIII. Female pore single in XIV, 0.3 mm. First dorsal pore in 12/13.
Clitellum annular in XIV-XVI; setae visible externally.

Male pores in XVIII at posterior corner of large angular papillae with
dimensions 1.7 x 2.0 mm with big ring 2.0-3.0 mm outer dimensions, each pad
extending to 17/18 and 18/19, each papilla hardened, with seminal groove of variable
shape in clitellates, oval, round. Paired spermathecal pores in 5/6 and 6/7, un-
recognizable. Genital markings lacking.

Septa 5/6 thin, 6/7 thick, 7/8 thin, 8/9, 9/10 absent, 10/11-13/14 thin. Gizzard in
VIH-X. Intestine begins in XV. Typhlosole as simple fold from XX VII. Intestinal caeca
simple, originating in XXVII, and extending anteriorly about to XXIII, each consisting
of a finger-shaped sac. Esophageal hearts four pairs in X-XIII, [X lateral.

Ovaries in XIII. Paired spermathecae in VI, VII; each ampulla as a flattened,
blunt, oval pouch, duct shorter than ampulla, diverticulum coiled and kinked with short
muscular stalks; no nephridia on spermathecal ducts. Male sexual system holandric,
testes and funnels enclosed in ventral paired sacs in X and XI. Seminal vesicles, two
pairs in XI and XII; those of XI enclosed in testis sacs. Prostates in X VIII extending to
XVU-XXI; both glandular portions consist of three main lobes, deeply divided in
slender leaflets; vas deferens not muscular. Genital papillae glands lacking.

REMARKS: Amynthas angulatus sp. nov. keys to the morrisi group in Sims &
Easton (1972), which is composed of 30 species. Among them, the following Korean
species are reported: A. fibulus fibulus (Kobayashi, 1936), A. fibulus ranunculus
(Kobayashi, 1936), A. kobayashii (Kobayashi, 1938), and A. koreanus (Kobayashi,
1938). In the shape of the male discs Amynthas angulatus sp. nov. is similar to A.

NEW SPECIES OF EARTHWORMS FROM KOREA 723

FIG. 1

Amynthas angulatus sp. nov. (A) Ventral view. (B) Male pore region in XVIII. (C) Sperma-
thecae. Scale bars = 5 mm (A), 2 mm (B, C).

kobayashii, but it differs in the male pore region. A. kobayashii has clear male pore, but
in Amynthas angulatus sp. nov. it is unrecognizable on the male discs. A. fibulus fibulus
and A. fibulus ranunculus have different male discs. After Sims & Easton (1972), two
species of the morrisi group were recorded from Korea, A. geojeinsulae (Song & Paik,
1970b) and A. draconis Hong & James, 2001. The new species appears to be related to
A. draconis, but is distinguishable easily by the shape of the male pore region. Its male
discs are angular, while A. draconis has round discs. Also, Amynthas angulatus sp. nov.
differs from A. draconis in having more setae in VII and XX, shorter body, more
closely spaced ventral and dorsal gap, and coiled diverticula.

Chen (1933, 1936, 1938, 1946) recorded 12 species of morrisi group from
China. Among them, two species have discs-shaped male pore regions: A. hainanicus
(Chen, 1938) and A. oculatus (Chen, 1938), but these discs are quite differently shaped
than in Amynthas angulatus sp. nov.

Amynthas jamesi sp. nov. Figs 2A-C
MATERIAL: Holotype: Clitellate specimen: Korea, Jeollanam-do province, Wando-gun,
Bogildo Isl., Buyong-ri, Jeokjabong, litter layers in forest, 19 July 2000, Y. Hong coll. Other
material: Same data as for holotype, | aclitellate specimen.
ETYMOLOGY: Named after Dr Samuel W. James, who made great contributions
to systematics of earthworms.

724 Y. HONG

FIG. 2

Amynthas jamesi sp. nov. (A) Ventral view. (B) Male pore region in XVIII. (C) Spermathecae.
Scale bars = 5 mm (A), 3 mm (B, C).

DIAGNOSIS: Three pairs of spermathecal pores in 5/6-7/8; male pores at lateral
margins of ventrum in XVIII; disc shape resembles a droplet placed with its narrow
end laterally, central axis with a diagonal seminal groove; distance between male pores
1.8 mm.

DESCRIPTION: Worms unpigmented. Dimensions 59-63 by 2.5-2.3 mm at
segment X, 3.0-3.5 mm at XXX, 2.8-3.0 mm at clitellum; body cylindrical throughout,
segments 88-102. Setae regularly distributed around segmental equators, numbering 32
at VII, 46 at XX; 3-5 between male pores; setal formula AA: AB: YZ:ZZ = 2:1:2:3.5 at
XIII. Female pore single in XIV, 0.3 mm, oval. First dorsal pore in 12/13. Clitellum
annular in XIV-XVI; setae at XVI visible externally within clitellum.

Male pores at lateral margins of ventrum in XVIII, as superficial bright spots
near lateral margins at the median ends of seminal grooves within small male discs;
disc shape resembles droplet placed with narrow end laterally, their central axis with a
diagonal seminal groove, distance between male pores 1.8 mm. Three pairs of sper-
mathecal pores in 5/6-7/8, inconspicuous, small, ventral. Genital markings absent.

Septa 5/6-7/8 thin, 8/9 absent, 9/10 present as sac enclosing testes and funnels;
10/11-13/14 thin. Gizzard globular in VIII-IX. Intestine begins in XV. Typhlosole
medium, from XXVII. Intestinal caeca simple, originating in XX VII, extending ante-
riorly to about XXV, each consisting of one finger-shaped lobe. Hearts X-XIII
esophageal, IX lateral.

Ovaries in XIII. Three pairs of spermathecae in VI, VII and VIII; each ampulla
as a small broad oval pouch, ducts shorter than ampulla; diverticula slender, stalked,
with long sausage-shape chamber, longer than ampulla; no nephridia on spermathecal

NEW SPECIES OF EARTHWORMS FROM KOREA 25

ducts. Male sexual system holandric, testes and funnels in paired sacs in X and XI.
Seminal vesicles, 2 pairs in XI and XII, slightly developed. Prostates in XVIII,
extending to XVII-XXII; ducts long, muscular, both glandular portions consisting of
two main lobes, each lobe divided into leaflets.

REMARKS: The present species keys to hawayanus (gracilis) group by the three
spermathecal pores in 5/6-7/8. After Sims & Easton (1972), the Korean species of this
group are A. acinctus (Goto & Hatai, 1899), A. agrestis (Goto & Hatai, 1898), A.
carnosus (Goto & Hatai, 1899), A. hilgendorfi (Michaelsen, 1892), A. kamitai
(Kobayashi, 1934), A. serratus (Kobayashi, 1936), and A. vallis (Kobayashi, 1936).
These species all have quite different characters from Amynthas jamesi sp. nov., such
as manicate caecae and different male field configurations. Later on, two more species
of the group were recorded in Korea, A. palgongensis Hong, 2001 and A. minjae Hong,
2001.

The new species is similar to A. minjae Hong, 2001 in the male pore region and
number of spermatheca, but has a different shape of the spermathecal diverticulum, a
longer diverticulum relative to the ampulla, obvious seminal grooves on male discs,
and fewer setae per segment at VII and XX.

Chen (1933, 1936, 1938, 1946) recorded 10 species of this group in China.
Amynthas jamesi sp. nov. is similar to the Chinese A. muticus (Chen, 1938) and A.
magnificus (Chen, 1936) by the male pore region, but is separated easily by the disc-
shape and diverticulum.

Amynthas yunseondoi sp. nov. Figs 3A-C

MATERIAL: Holotype: Clitellate specimen: Korea, Jeollanam-do province, Wando-gun,
Bogildo Isl., Buyong-dong, Bojuksan, litter layers in forest, 19 July 2000, Y. Hong coll.

ETYMOLOGY: Yun Seon-Do (1587-1671) who was the master of the Korean
literary circle in the age of Chosun Dynasty built a pavilion at his place of residence in
Bogildo Island.

DIAGNOSIS: Spermathecal pores in VI, VIT and VIII close to 5/6, 6/7 and 7/8;
male pores superficial at median ends of seminal grooves within paired discs; disc
shape resembles droplet placed with its narrow end posterior, central axis with longi-
tudinal seminal groove; distance between male pores 6.5 mm.

DESCRIPTION: Worm unpigmented. Dimensions 156 mm by 7.0 mm at segment
X, 7.0 mm at XXX, 8.2 mm at clitellum; body cylindrical throughout, segments 126.
Setae regularly distributed around segmental equators, numbering 51 at VII, 69 at XX;
14 between male pores, size and distance regular; setal formula AA:AB:YZ:ZZ =
5:3:4:5 at XIII. Female pore single in XIV, 1.0 mm wide, oval, slightly invaginated.
First dorsal pore in 12/13. Clitellum annular in XIV-XVI; setae invisible externally in
the clitellum.

Male pores superficial at median ends of seminal grooves within paired,
elevated hardened male discs; disc shape resembles droplet placed with narrow end
posterior, raised above body wall level, central axis with longitudinal seminal groove,
lateral end posterior to medial end, distance between male pores 6.5 mm. Spermathecal
pores in VI, VII and VIII close to 5/6, 6/7 and 7/8, above mid-lateral level, on slightly
elevated conspicuous small bumps, pore opening appears black.

726 Y. HONG

FIG. 3

Amynthas yunseondoi sp. nov. (A) Ventral view. (B) Male pore region in XVIII. (C) Sper-
mathecae. Scale bars = 5 mm (A), 3 mm (B), 2 mm (C).

Septa 5/6-7/8 thick, 8/9, 9/10 absent, 10/11 moderately muscular, 11/12, 12/13
thick, 13/14 thin. Gizzard globular in VIII-X. Intestine begins in XV, small paired
lymph glands from XXVIII along dorsal vessel. Typhlosole medium as a simple fold
from XXVII. Hearts in X-XIII esophageal; in IX lateral, on the left side larger.
Intestinal caeca simple, originating in XXVII, and extending anteriorly about to XXIV,
each consisting of a finger-shaped sac with many small pouches on vertical margin.

Ovaries in XIII. Paired spermathecae in VI, VII and VIII; ampulla as a small
pouch, ducts shorter than ampulla, of medium thickness; diverticula slender, consisting
of digitate chamber with narrow stalk, shorter than ampulla. Male sexual system
holandric, testes and funnels in ventral paired sacs in X and XI. Seminal vesicles two
pairs in XI and XII, well developed. Prostates in X VIII, divided in many long slender

NEW SPECIES OF EARTHWORMS FROM KOREA 124];

lobes extending to XVII-XIX, many pieces; vas deferens muscular, clearly single.
Genital marking glands lacking.

REMARKS: The species is similar to A. deogyusanensis Hong & James, 2001,
with respect to the shape of male pore region. It differs from A. deogyusanensis in the
orientation of the male disc’s droplet shape head direction and location of the male
pores. Amynthas yunseondoi sp. nov. also differs from A. deogyusanensis in having
more setae per segment at VII and XX, greater length, and more widely spaced ventral
and dorsal gap. Amynthas yunseondoi sp. nov. has three pairs of spermathecal pores in
VI-VII, but A. deogyusanensis has two pairs of spermathecal pores in VI-VII, also
lacks genital markings but has conspicuous genital patches, while the new species has
neither genital markings nor genital patches. The sampling locality of Amynthas
yunseondoi sp. nov. is an island of southern Korea, but A. deogyusanensis was
collected from Mt. Deogyu of the central mainland Korea.

Amynthas yunseondoi sp. nov. keys to the bournei group with three pairs of
spermathecal pores intrasegmental in VI-VIII. This group is composed of four species;
A. bournei (Rosa, 1890), A. domosus (Chen, 1946), A. mucrorimus (Chen, 1946), and
A. sulcatus (Gates, 1932). After Sims & Easton (1972), one species recorded in Korea,
A. pagyeiensis Hong, 2001, but it differs by the seminal grooves within the male discs
(Hong et al., 2001).

ACKNOWLEDGEMENTS

This work was supported by Korea Research Foundation Grant (KRF-2004-
050-C00019). I would like to express appreciation to Dr Samuel W. James, University
of Kansas, USA, who kindly shared valuable bionomical information and reviewed the
taxonomic descriptions in the manuscript.

REFERENCES

CHEN, Y. 1933. A preliminary survey of the earthworms of the Lower Yangtze valley.
Contributions of the Biology Laboratory Sciences Society of China (Zoology), Nanking
9: 178-296.

CHEN, Y. 1936. On the terrestrial Oligochaeta from Szechwan II with the notes on Gates’ types.
Contributions of the Biology Laboratory Sciences Society of China (Zoology), Nanking
11: 269-306.

CHEN, Y. 1938. Oligochaeta from Hainan, Kwangtung. Contributions from Biological
Laboratory of Science Society of China (Zoological series) 12: 375-427.

CHEN, Y. 1946. On the terrestrial Oligochaeta from Szechwan III. Journal of the West China
Border Research Society 16: 83-141.

GATES, G. E. 1932. The earthworms of Burma III, the Megascolecidae. Records of the Indian
Museum, Calcutta 34: 357-549.

GATES, G. E. 1936. On some earthworms from the Cameron Highlands, Pahang. Bulletin of the
Raffles Museum, Singapore 12: 87-117.

Goro, S. & HATAI, S. 1898. New or imperfectly known species of earthworms. No. 1. Anno-
tationes Zoologicae Japonenses Tokyo 2: 65-78.

Goro, S. & HATAI, S. 1899. New or imperfectly known species of earthworms. No. 2. Anno-
tationes Zoologicae Japonenses Tokyo 3: 13-24.

HATAI, S. 1930. Note on Pheretima agrestis (Goto & Hatai), together with the description of four

new species of the genus Pheretima. Scientific Reports of Tohoku Imperial University,
Tokyo 5: 651-667.

728 Y. HONG

HONG, Y. & JAMES, S. W. 2001a. New species of Korean Amynthas Kinberg, 1867 (Oligochaeta,
Megascolecidae) with two pairs of spermathecae. Revue suisse de Zoologie 108: 65-93.

HONG, Y. & JAMES, S. W. 2001b. Five new earthworms of the genus Amynthas Kinberg
(Megascolecidae) with four pairs of spermathecae. Zoological Studies 40: 269-275.

HONG, Y. & LEE, W. K. 2001. Description of three new Korean earthworms of the genus
Amynthas Kinberg, 1867 (Oligochaeta, Megascolecidae) with multiple genital markings.
Revue suisse de Zoologie 108: 283-290.

HONG, Y., LEE, W. K. & Kim, T. H. 2001. Four new species of the genus Amynthas Kinberg
(Oligochaeta: Megascolecidae) from Korea. Zoological Studies 40: 263-268.

KOBAYASHI, S. 1934. Three new Korean earthworms belonging to the genus Pheretima, together
with the wider range of the distribution of Pheretima hilgendorfi (Michaelsen). Journal
of the Chosen Natural History Society, Keijo 19: 1-11.

KOBAYASHI, S. 1936. Earthworms from Koryo, Korea. Scientific Reports of Tohoku Imperial
University, Tokyo 11: 139-184.

KOBAYASHI, S. 1937. Preliminary survey of the earthworms of Quelpart Island. Scientific Reports
of Tohoku Imperial University, Tokyo 11: 333-351.

KOBAYASHI, S. 1938. Earthworms of Korea I. Scientific Reports of Tohoku Imperial University,
Tokyo 3: 89-170.

MICHAELSEN, W. 1892. Terricolen der Berliner Zoologischen Sammlung, II. Archiv fiir Natur-
geschichte, Berlin 58: 209-261.

Rosa, D. 1890. Viaggio di Leonardo Festa in Birmanica e regioni vicine, XX VI. Perichetidi.
Annali del Museo di Storia Naturale di Genova 10: 107-122.

SIMS, R. W. & Easton, E. G. 1972. A numerical revision of the earthworm genus Pheretima auct.
(Megascolecidae: Oligochaeta) with the recognition of new genera and an appendix on
the earthworms collected by the Royal Society North Borneo Expedition. The Biological
Journal of the Linnean Society, London 4: 169-268.

SONG, M. J. & Paik, K. Y. 1970a. Earthworms from Chejoo-do Island, Korea. Korean Journal of
Zoology 13: 9-14.

SONG, M. J. & Paik, K. Y. 1970b. On a small collection of earthworms from Geo-je Isl., Korea.
Korean Journal of Zoology 13: 101-111.

SONG, M. J. & Paik, K. Y. 1973. Earthworms from Mt. Sopaik, Korea. Korean Journal of Zoology
1659-12:

REVUE SUISSE DE ZOOLOGIE 114 (4): 729-733; décembre 2007

A taxonomic revision of the family Oncopodidae VII. A new
Oncopus species (Opiliones, Laniatores) from eastern Kalimantan

Peter J. SCHWENDINGER
Museum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland.
E-mail: peter.schwendinger@ville-ge.ch

A taxonomic revision of the family Oncopodidae VII. A new Oncopus
species (Opiliones, Laniatores) from eastern Kalimantan. - Oncopus
kaltim sp. n. is described from males collected in the Indonesian part of
Borneo. This species belongs to the Oncopus hosei species-group and
further supports the assumption that O. expatriatus Schwendinger &
Martens, 2004 occurs on Borneo and not in Thailand.

Keywords: Taxonomy - penis morphology - Borneo - Indonesia.

INTRODUCTION

After having completed a revision of the available Oncopus Thorell material in
co-operation with Jochen Martens (see Schwendinger & Martens, 2004), I received
two additional male specimens collected by Louis Deharveng and Anne Bedos in the
eastern part of Borneo. These specimens are presumably conspecific with five juveniles
earlier reported from the same area (Schwendinger & Martens, 2004: 171) and
represent a new species that is described and illustrated here.

METHODS

Methods and terminology follow Martens & Schwendinger (1998: fig. 1) and
Schwendinger & Martens (2004: 140). All measurements are in mm. Leg and palp
measurements are here given as: Total length (length of trochanter + femur + patella +
tibia + [metatarsus +] tarsus).

TAXONOMY

Oncopus kaltim sp. n. Figs 1-15

MATERIAL: INDONESIA, East Kalimantan Province, Sangkulirang, Pengadan, Baai
Forest, d holotype (deposited in the Museum Zoologicum Bogoriense, Cibinong, Java,
Indonesia) and d paratype (deposited in the Muséum National d'Histoire Naturelle, Paris,
France), collected by hand from forest litter by L. Deharveng, A. Bedos, C. Rahmadi & Y.
Suhardjono on 14.-16.VIII.2004.

ETYMOLOGY: The specific epithet, a noun in apposition, refers to the Indonesian
name of the province in which the types were collected. "Kaltim" is the locally used
short form of Kalimantan Timur.

Manuscript accepted 01.11.2007

730 P. J. SCHWENDINGER

DrAGNOsis: Similar to O. megachelis Schwendinger, 1992, distinguished by:
Smaller size; interocular area less rounded; paramedian tubercles present on dorsal
scutal area VIII; anteroproximal processes on coxae III shorter, stouter, parallel to each
other; genital operculum narrower; truncus penis with narrower, widely notched distal
margin; median plate of glans penis shorter and with straight distal margin; lateral
sclerites of glans longer, with truncate tips.

DESCRIPTION: d (holotype, except for penis morphology). Colouration: Body
amber, with dark reticulation on carapace region, chelicerae and palps; dark margin on
dorsal scutum and dark pattern on its opisthosomal part (Fig. 1); legs mostly dark,
except for lighter distal portion of metatarsi I-IV, lighter metatarsus III and tarsi III-IV,
and cream tarsi I-II.

Dorsal scutum 7.3 long and 4.6 wide, its carapace region large and raised, 2.4
long and 2.9 wide, interocular area a low, rounded, slightly forward-directed hump
(Figs 1-2). Opisthosomal areas of dorsal scutum indistinctly keeled, with pairs of small
paramedian tubercles on areas VI-VIII, the latter slightly protruding from posterior
margin of scutum (Figs 1-3); ventral scutal areas distinctly keeled, with transversal
bands of white enclosures (Figs 2-3). Palpal coxa with small, conical ventral process;
ventral side of leg coxa I with small anterolateral process, low central mound and sub-
proximal bulge; coxa II with small scale-like anteroproximal apophysis and low
rounded process posterior to it; coxa III with fairly large anteroproximal process.
Genital operculum rounded, slightly longer than wide (Fig. 10).

Chelicerae (Figs 1, 11-12) strong; proximal article with small, rounded retro-
ventral tubercle and anteriad-inclined prodorsal boss (Fig. 12); second article with
small ventrodistal tubercle, mobile finger without ventral process; cutting edge of
mobile finger with one rounded proximal tooth situated slightly more distally than
indistinct proximal tooth on cutting edge of fixed finger; both cutting edges slightly
worn (Fig. 11).

Palps (Fig. 13): Trochanter with small ventral process; femur with only an
indistinct ventroproximal hump, otherwise unarmed; patella without proventral
process; tibia with low retroventral proximal process. Measurements: 6.1 (0.9 + 1.8 +
0.9 + 0.8 + 1.7).

Legs 3124 (from shortest to longest). Measurements: Leg I 8.1 (0.8 + 2.5 + 1.1
+ 1.2 + 2.1 + 0.4); leg II 11.3 (1.0 + 3.3 + 1.5 + 1.9 + 3.2 + 0.4); leg II 8.0 (0.9 + 2.0
+ 1.2 + 1.2 + 2.3 + 0.4); leg IV 11.8 (1.0 + 2.8 + 1.7 + 2.0 + 3.8 + 0.5). Femur I with
small dorsoproximal tubercle. All leg tarsi with dorsal pore organ (see Schwendinger,
2006).

Penis of paratype (Figs 4-7): Truncus fairly stout, proximal half narrower than
distal half, widened between glans and distal margin, there carrying subterminal lateral
setae arranged in an irregular recurved row on each side; distal margin of truncus with
a widely V-shaped indentation forming two triangular sockets carrying one seta on one
side and two setae on the other. Glans penis narrower than truncus at that point; mem-
branous socket flanked by two setae on each side; lateral sclerites bent outwards and
away from the truncus, their bases with knee-like lateral projections, their apices
truncate; median plate short, with wide and straight distal margin; membranous tubes
distinctly protruding beyond median plate.

2. Unknown.

ONCOPUS FROM KALIMANTAN 731

Fics 1-3
Oncopus kaltim sp. n., 4 holotype. - Habitus, (1) dorsal, (2) ventral and (3) lateral view.

REMARKS: The penis of the holotype (Fig. 8) has collapsed below the glans,
which gives the apex penis a narrower appearance than in the paratype (Fig. 6). The
penis of the paratype is therefore described above.

VARIATION: Measurements of & paratype: Dorsal scutum 7.4 long and 4.4 wide,
its carapace region 2.3 long and 2.9 wide. Interocular area of paratype slightly more
pointed, cheliceral fingers shorter and their cutting edges more strongly worn (without
teeth), ventral process on palpal trochanter longer (Fig. 14), dorsoproximal tubercle on
femur II larger (Fig. 15) than in holotype. The holotype penis has only one seta on each
side of its distal margin (Fig. 8), which is presumably normal (as in all other hosei-
group species), whereas the paratype has two setae on one side (Fig. 6). Other
differences between the illustrations of both penes (Figs 6-7, cf. Figs 8-9) are due to
the collapsed state of the holotype penis.

RELATIONSHIPS: Genital and somatic characters clearly show that the new
species belongs to the hosei-group (see Schwendinger & Martens, 2004: 165), which
now comprises O. hosei Pocock, 1897 (eastern Sarawak), O. megachelis (eastern
Sabah), O. kaltim sp. n. (eastern Kalimantan) and O. expatriatus (doubtful record from
Bangkok, Thailand). The discovery of a third species in this group in eastern Borneo
is another piece of circumstantial evidence that Roewer's locality data for the holotype
of O. expatriatus are incorrect (see Schwendinger & Martens, 2004: 170).

DISTRIBUTION: Known only from the type locality.

732 P. J. SCHWENDINGER

15 10

Fics 4-15

Oncopus kaltim sp. n., (8-13) 4 holotype and (4-7, 14-15) d paratype. - Total penis, (4) dorsal
and (5) lateral view. Apex of penis, (6, 8 [partly collapsed]) dorsal and (7) lateral view. (9) Glans
penis, lateral view. (10) Anterior part of body, ventral view. (11) Left chelicera, retrolateral view.
(12) Proximal article of left chelicera, retrodorsal view. (13) Left palp, retrolateral view. (14)
Left palpal trochanter, retrolateral view. (15) Left trochanter II and proximal part of femur II,
retrolateral view. - Scale lines: (4-5; 10-15) 1.0 mm, (6-7; 8-9) 0.1 mm.

ACKNOWLEDGEMENTS

Louis Deharveng and Anne Bedos (Muséum National d'Histoire Naturelle,
Paris) kindly provided the material examined, Jochen Martens (Universität Mainz)
commented on the manuscript, John Hollier checked the English and helped scan the
drawings, and Florence Marteau (Muséum d'histoire naturelle, Genève) helped
compose the figure plates.

ONCOPUS FROM KALIMANTAN 733

REFERENCES

MARTENS, J. & SCHWENDINGER, P. 1998. A taxonomic revision of the family Oncopodidae I. New
genera and new species of Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de
Zoologie 105 (3): 499-555.

Pocock, R. I. 1897. Descriptions of some new Oriental Opiliones recently received by the
British Museum. Annals and Magazine of Natural History (6) 19: 283-310.

SCHWENDINGER, P. J. 1992. New Oncopodidae (Opiliones, Laniatores) from Southeast Asia.
Revue suisse de Zoologie 99 (1): 177-199.

SCHWENDINGER, P. J. 2006. A taxonomic revision of the family Oncopodidae VI. Martensiellus,
a new genus from Borneo, and the discovery of a tarsal pore organ in Oncopodidae
(Opiliones: Laniatores). Zootaxa 1325: 255-266.

SCHWENDINGER, P. J. & MARTENS, J. 2004. A taxonomic revision of the family Oncopodidae IV.

The genus Oncopus Thorell (Opiliones, Laniatores). Revue suisse de Zoologie 111 (1):
139-174.

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REVUE SUISSE DE ZOOLOGIE 114 (4): 735-741; décembre 2007

Description of a new species of Entedonomphale (Hymenoptera:
Eulophidae) from Bulgaria, with notes on E. carbonaria

Peter S. BOYADZHIEV! & Serguei V. TRIAPITSYN?

! Department of Zoology, University of Plovdiv “Paisii Hilendarski”,
24 Tsar Asen St., 4000 Plovdiv, Bulgaria. E-mail: boyadz@pu.acad.bg

2 Entomology Research Museum, Department of Entomology, University of
California, Riverside, California, 92521, USA. E-mail: serguei.triapitsyn @ucr.edu

Description of a new species of Entedonomphale (Hymenoptera: Eulo-
phidae) from Bulgaria, with notes on E. carbonaria. - A new distinctive
species of the genus Entedonomphale Girault (Hymenoptera: Eulophidae:
Entedoninae), E. bulgarica Boyadzhiev & Triapitsyn sp. n., is described
from the mountains of southwestern and southeastern Bulgaria. The
European species E. carbonaria (Erdös) is newly recorded from the
Nearctic region (USA). An identification key to both sexes of the
Palaearctic species of Entedonomphale is provided.

Keywords: Hymenoptera - Eulophidae - Entedoninae - Entedonomphale -
taxonomy - Bulgaria - screen-sweeping net.

INTRODUCTION

The rarely collected genus Entedonomphale Girault, 1915 (Hymenoptera:
Eulophidae: Entedoninae) was recently revised on a worldwide basis by Triapitsyn
(2005). The species of Entedonomphale are larval parasitoids of Phlaeothripidae
(Thysanoptera: Tubulifera) (Triapitsyn, 2005). In Europe, this genus is represented by
two species, E. bicolorata (Ishii, 1933) and E. carbonaria (Erdös, 1954) (Triapitsyn,
2005).

Here we diagnose, describe, and illustrate a new, very distinctive species of
Entedonomphale that was collected by the senior author first by screen sweeping at low
elevation on the Pirin Mt. in southwestern Bulgaria; later 3 males of the same species
were collected on the Sakar Mt. in southeastern Bulgaria. Additionally, we provide new
information on the distribution of E. carbonaria, which was recently discovered in the
Nearctic region (Oregon, USA). An identification key to both sexes of the Palaearctic
species of Entedonomphale is also provided.

MATERIAL AND METHODS

The type specimens were collected with a modified net (Fig. 10) for screen
sweeping (after Noyes, 1982) and fixed in 96% alcohol. In laboratory they were dried
from ethanol using a critical point drier and point-mounted, and then the female and
one of the males (from the same locality as the female) were cleared in 10% KOH,

Manuscript accepted 13.07.2007

736 P. S. BOYADZHEV & S. V. TRIAPITSYN

dissected, and slide-mounted in Canada balsam. The slide-mounted specimens were
examined under a Zeiss Axioskop 2 plus compound microscope (using Nomarski
differential interference contrast optics) and photographed using a Sony DSC-S75
digital still camera. Morphological terminology follows Gibson (1997); measurements
(as length or length:width for the wings) are given in micrometers (um). The examined
specimens are deposited in the collections indicated by the following acronyms:
MHNG, Muséum d’histoire naturelle, Geneva, Switzerland; PUPB, Department of
Zoology, University of Plovdiv “Paisii Hilendarski”, Plovdiv, Bulgaria; UCDC, The
R.M. Bohart Museum of Entomology, Department of Entomology, University of
California, Davis, California, USA; and UCRC, Entomology Research Museum,
Department of Entomology, University of California, Riverside, California, USA. An
abbreviation used in the key and description is: F = antennal funicular segment.

RESULTS
Genus Entedonomphale Girault

KEY TO THE PALAEARCTIC SPECIES

[male of E. bicolorata (Ishii) is unknown]

1 Female (antenna with a 2-segmented funicle and an entire clava).......... y
- Male (antenna either with a 3-segmented funicle and an entire clava or

with a 2-segmented funicle and a 3-segmented clava).................. +
2(1) Forewing disc completely hyaline, without a transverse dark band in the

md a) OS asec cere ee orgs eee MASSE ne oe Neer ee ee oo E. bicolorata (Ishii)

Forewing disc with a conspicuous transverse dark band in the middle...... 3

3(2) FI almost as long as F2 (only slightly shorter); postmarginal vein
2.4-2.6 x length of stigmal vein; ovipositor 0.6 x length of metatibia
eA a PRE Res E. bulgarica Boyadzhiev & S. Triapitsyn sp. n.
- F1 markedly shorter than F2; postmarginal vein about as long (1.0 x) as
stigmal vein; ovipositor 1.1 x length of metatibia...... E. carbonaria (Erd6s)
4(1) Antenna with a 2-segmented funicle and a 3-segmented clava
UNSERE Sn a E. bulgarica Boyadzhiev & S. Triapitsyn sp. n.
- Antenna with a 3-segmented funicle and an entire clava . E. carbonaria (Erdös)

Entedonomphale bulgarica Boyadzhiev & S. Triapitsyn, sp. n. Figs 1-9

TYPE MATERIAL: Holotype: female on slide [MHNG]: BULGARIA, Blagoevgrad Region,
Pirin Mt. 2 km W of Hadzhidimovo, 41°31’15”N, 23°50’40”E, 495 m, 8.ix.2006, PS.
Boyadzhiev. Paratypes: 1 male on slide [PUPB], same data as the holotype. 3 males on points:
BULGARIA, Stara Zagora Region, Sakar Mt., 1 km NE of Madrets Village: 42°07’55”N,
26°05’38”E, 140 m, 26.iv.2007, PS. Boyadzhiev [2 males, PUPB, UCRC]; 42°08’06”N,
26°06’ 11” E, 160 m, 26.iv.2007, A.M. Stojanova [1 male, PUPB].

DIAGNOSIS: Both sexes of this new species can be easily distinguished from the
other two Palaearctic species of Entedonomphale using the morphological characters
indicated in the key above. The male of E. bulgarica sp. n. is somewhat similar to that
of the Australian species E. zakavyka Triapitsyn, 2005 as both taxa have the funicle
2-segmented and the clava 3-segmented. However, the male forewing of E. zakavyka

A NEW SPECIES OF ENTEDONOMPHALE FROM BULGARIA 737

/ A 2

2/7,

1,010

4

Fics 1-4

Entedonomphale bulgarica sp. n. (holotype female): (1) Antennae. (2) Mesosoma and meta-
soma. (3) Forewing. (4) Hind wing. Scale lines = 0.1 mm.

738 P. S. BOYADZHEV & S. V. TRIAPITSYN

Fics 5-9

Entedonomphale bulgarica sp. n. (paratype male from Pirin Mt., Bulgaria): (5) Antenna. (6)
Mesosoma and petiole. (7) Forewing. (8) Hind wing. (9) Genitalia. Scale lines = 0.1 mm.

A NEW SPECIES OF ENTEDONOMPHALE FROM BULGARIA 739

10

Plastic diaphragm with perforations (6-10 mm)
Press-button

Plastic holder with inner thread (29 mm)
Sa Plastic bottle (100 ml) covered with adhesive tape

Fic. 10
Schematic diagram of the modified net for screen sweeping.

50 cm

has a conspicuous transverse dark band in the middle, and F2 of its female antenna is
slightly wider than long (Triapitsyn, 2005), whereas the male forewing of E. bulgarica
sp. n. is only slightly infumate, and F2 of its female antenna is distinctly longer than
wide.

DESCRIPTION: FEMALE (holotype): Body shining black with a slight violet
tinge except petiole whitish (particularly distally) to light brown. Scape light brown
(distally) to brown (basally), pedicel and flagellum brown to dark brown. Coxae and
trochanters whitish to light brown, femora and tibiae brown to dark brown, tarsı mostly
light brown.

Vertexal suture faint. Antenna (Fig. 1) with numerous setae, these denser on
flagellar segments; scape slender, about 6 x as long as wide, only a little expanded in
basal half; both funicular segments longer than wide, F1 slightly shorter and much
narrower than F2, F1 with 1 longitudinal sensillum, F2 with at least 3 (possibly with
4) longitudinal sensilla; clava much longer than funicle, about 3 x as long as wide, with
numerous (at least 12) longitudinal sensilla.

Mesosoma a little shorter than gaster (Fig. 2), mostly smooth (except pronotum
lightly sculptured). Midlobe of mesoscutum with 1 pair of setae; anterior margin of
scutellum almost straight. Scutellum shorter than mesoscutum, with 1 pair of setae and
placoid sensilla at lateral margins. Forewing (Fig. 3) 3.2 x as long as wide; post-
marginal vein very long for the genus, much longer than stigmal vein (2.4-2.6 x);
longest marginal setae a little less than two fifths of maximal forewing width; disc with

740 P. S. BOYADZHEV & S. V. TRIAPITSYN

a dark, transverse band behind stigmal and most of marginal vein (reaching posterior
margin), more or less evenly setose in apical three fifths of forewing (setae short). Hind
wing (Fig. 4) 8.0 x as long as wide; longest marginal setae about 1.1 x maximal hind
wing width; disc with short setae, slightly infumate at apex of venation. Coxae weakly
sculptured (hind coxa more so).

Petiole conspicuous, more or less cylindrical, 1.2 x as long as wide. Ovipositor
short, occupying a little more than one third length of gaster, not exserted beyond
gastral apex; ovipositor 0.6 x length of metatibia.

Measurements: Body (length of the dry specimen before slide-mounting): 859;
head (length of the dry specimen before slide-mounting): 150; mesosoma: 455; petiole:
64; gaster: 470; ovipositor: 170. Antenna: scape (including radicle): 200; pedicel: 77;
F1: 55; F2: 58; clava: 162. Forewing: 861:267; longest marginal seta: 100. Hind wing:
750:94; longest marginal seta: 106.

MALE (paratypes): Body length (of the dry specimens before slide-mounting):
673-863. Head and mesosoma shining dark brown to black, petiole light brown,
antenna and gaster brown, legs light brown to brown. Antenna (Fig. 5) with a 2-seg-
mented funicle (Fl subequal to F2, both a little longer than wide) and a 3-segmented
clava with a long apical spicula, claval segments more or less subequal to funicular
segments in size; scape 3.1 x as long as wide; all flagellar segments with numerous
long setae exceeding each segment’s width and without longitudinal sensilla. Midlobe
of mesoscutum with 2 pairs of setae (Fig. 6). Forewing (Fig. 7) 3.5 x as long as wide;
longest marginal setae about three fifths maximal forewing width; disc considerably
less pigmented in the middle, only slightly infumate (more so behind stigmal and
marginal veins), with cubital row of setae sinuate. Hind wing (Fig. 8) 9.3 x as long as
wide; longest marginal setae about 1.6 x maximal forewing width; disc slightly
infumate. Petiole more or less trapezoidal, 1.2 x as long as wide. Genitalia as in Fig. 9,
typical for the genus.

ETYMOLOGY: The specific name (an adjective, gender feminine) refers to the
country (Bulgaria) where this species occurs.

Host: Unknown.

NOTE: The type specimens were collected on boggy grass communities and at
the edge of a mowed agricultural field.

COMMENTS: The female of the new species was also compared with a recently
identified specimen of E. zakavyka with the following label data: Australia,
Queensland, car net between Goomeri and Petris on Hwy 17 87, 9.1.1986, E.A. Sugden
[1 female, UCDC].

Entedonomphale carbonaria (Erdös)
Entedonomphale carbonaria (Erdös): Triapitsyn 2005: 285-286 (taxonomic history, diagnosis,
figures, distribution).

MATERIAL EXAMINED: USA, Oregon, Tillamook Co., 4 km W of Sandlake, 2.vii.1991,
S.L. Heydon [1 female, UCDC].

DISTRIBUTION: Europe (records from Bulgaria, Germany, Hungary, Russia,
Slovakia, Slovenia, Sweden) (Triapitsyn, 2005) and North America (USA) [new
record].

A NEW SPECIES OF ENTEDONOMPHALE FROM BULGARIA 741

COMMENTS: As noted by Triapitsyn (2005), the Nearctic species E. kaulbarsi
(Yoshimoto) might be a wingless (female only) form of E. carbonaria, as their other
morphological features are identical; besides, their fully winged males are also iden-
tical. The discovery of the fully winged E. carbonaria in Oregon, USA, is puzzling.
Further research using molecular methods will be necessary to address the limits of
these species.

ACKNOWLEDGEMENTS

We thank Vladimir V. Berezovskiy (UCRC) for making excellent slide-mounts
of the type specimens of the new species described in this communication, Dr Anelia
M. Stojanova (PUPB) for collecting one of the specimens, and Dr Steve L. Heydon
(UCDC) for the loan of material. The collecting trips in Bulgaria were supported by the
Fund of the Scientific Research, University of Plovdiv “Paisii Hilendarski”.

REFERENCES

GIBSON, G. A. P. 1997. Chapter 2. Morphology and terminology (pp. 16-44). In: GIBSON,
G. A. P., HUBER, J. T. & WOOLLEY, J. B. (eds). Annotated keys to the genera of Nearctic
Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, Ontario, Canada, 794 pp.

Noyes, J. S. 1982. Collecting and preserving chalcid wasps (Hymenoptera: Chalcidoidea).
Journal of Natural History 16: 315-334.

TRIAPITSYN, S. V. 2005. Revision of Ceranisus and the related thrips-attacking entedonine genera
(Hymenoptera: Eulophidae) of the world. African Invertebrates 46: 261-315.

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REVUE SUISSE DE ZOOLOGIE 114 (4): 743-769; décembre 2007

Le specie del genere Orphnebius Motschulscky, 1858, nel Borneo
(Coleoptera, Staphylinidae)*

Roberto PACE
Via Vittorio Veneto, 13, 1-37032 Monteforte d’Alpone (Verona), Italia.
E-mail: pace.ent@tiscali.it

The species of the genus Orphnebius Motschulscky, 1858, from Borneo
(Coleoptera, Staphylinidae). - This study on the genus Orphnebius from
Borneo contains the illustration and the revision of the followings six holo-
types or lectotypes: O. bigladiosus (Bernhauer), O. bakerianus Bernhauer,
O. splendens Bernhauer, O. anguliceps Cameron, O. antennarius Bernhauer
and O. bakeri Bernhauer. The lectotypes of O. bakeri Bernhauer and of O.
bakerianus Bernhauer, are designated. The genus Deroleptus Bernhauer and
the subgenus Mesocephalobius Bernhauer have been synonymized with
Orphnebius and Orphnebius quadrigladiosus Pace, 1986, with Orphnebius
quadricuspidatus Bernhauer, 1929. Deroleptus bigladiosus (Bernhauer) and
Deroleptus arachnoides Bernhauer are transferred to Orphnebius. The
followings 13 species are described as new: O. borneanus, O. scalaris, O.
directus, O. minor, O. biapicalis, O. ideogramma, O. penangensis, O.
bajauorum, O. curticornis, O. uniformis, O. perpenetrans, O. krypticola and
O. crassus. Habitus and male and female genitalia of the new species are
illustrated. A key to the species is provided.

Keywords: Coleoptera - Staphylinidae - Aleocharinae - taxonomy -
Borneo.

INTRODUZIONE

Il genere Orphnebius Motschulscky, 1858, è stato descritto per una specie della
Colombia (0. ventricosus Motschulscky, 1858). Pochi decenni dopo è stato ricono-
sciuto essere presente anche nella regione orientale, sottoregione malese compresa, in
cui rientra il Borneo. Cinque sono le specie note del genere Orphnebius per il Borneo:
O. anguliceps Cameron, 1943, del Brunei, e O. bakeri Bernhauer, 1929, O. bakerianus
Bernhauer, 1929, O. quadricuspidatus Bernhauer, 1929, O. splendens Bernhauer, 1929
e O. antennarius Bernhauer, 1929, tutti di Sandakan. Gli holotypi e i sintypi di tutte
queste cinque specie sono stati da me esaminati e dissezionati per disegnare edeago e/o
la spermateca. Nel 1915 Bernhauer descrisse il genere Deroleptus per la specie
D. bigladiosus (Bernhauer, 1915), del Borneo. Anche questa specie è stata da me esa-
minata e dissezionata. I risultati di queste mio esame sono esposti nel presente studio.

* 210° Contributo alla conoscenza delle Aleocharinae.
Manoscritto accettato il 07.08.2007

744 R. PACE

MATERIALE E METODO

La presente revisione delle specie note del genere Orphnebius e Deroleptus del
Borneo si è resa necessario al fine di determinare gli esemplari adulti raccolti preva-
lentemente nel Parco Nazionale del Monte Kinabalu dal Dr. Ale’ Smetana di Ottawa
durante le sue spedizioni nel 1987 e 1988, dalla spedizione Burckhardt & Löbl del
Museo di Storia Naturale di Ginevra del 1987-1988 e Bright del 1988 e dal materiale
riconosciuto nell’esame delle serie tipiche di Bernhauer al Field Museum of Natural
History di Chicago, da un esemplare del DEI e da esemplari affidatimi in esame dal
collega Dr. Volker Assing di Hannover.

La tassonomia delle nuove specie del Borneo viene qui mutata grazie all’esame
dei caratteri dell’organo copulatore maschile e della spermateca. Prima della pubblica-
zione del presente studio nessun esame a fini tassonomici di questi importanti organi è
stato portato a termine da vecchi autori. Gli holotypi delle specie note sono stati da me
disegnati dopo esame e inseriti nella chiave qui data dopo quella del 1929 data da
Bernhauer. Tutti gli esemplari sono stati dissezionati. Gli organi genitali sono stati
montati in balsamo del Canadà su piccoli rettangoli trasparenti di materiale plastico,
infilzati sullo spillo dell’esemplare. Le strutture genitali sono state studiate usando un
microscopio composto e disegnate mediante oculare a reticolo. Gli habitus sono stati
disegnati con l’uso di un oculare micrometrico di un microscopio binoculare. Tutti i
disegni sono dell’autore fino alla fase finale. Il sicuro riconoscimento dei generi e delle
specie da parte del lettore è qui affidato prevalentemente alla parte illustrativa che ha
linguaggio internazionale. Per questo motivo le descrizioni sono brevi, limitate a porre
in evidenza ciò che non è riproducibile graficamente come il colore, la reticolazione e
la granulosità. D'altronde per le specie della sottofamiglia Aleocharinae la sola descri-
zione anche molto accurata e lunga non dà quasi mai la certezza di un’esatta identi-
ficazione delle varie specie. È l'osservazione del disegno dell’edeago e/o della sper-
mateca insieme a quello dell’habitus che aiuta molto a risolvere problemi interpretativi
dati dalla sola descrizione.

Gli holotypi delle nuove specie sono depositati nel Museo di Storia Naturale di
Ginevra (MHNG), nel Field Museum of Natural History di Chicago (FMNH), in
collezione del D.E.I di Eberswalde (DEI) e all’ Institut Royal des Sciences Naturelles
de Belgique di Bruxelles (IRSB).

Paratypi sono conservati in collezione Smetana e nell’Institut Royal des
Sciences Naturelles de Belgique di Bruxelles.

ESAME DI MATERIALE TIPICO

Orphnebius bigladiosus (Bernhauer, 1915), comb. n. Figg. 5-7

Astilbus (Deroleptus) bigladiosus Bernhauer, 1915: 150
Deroleptus bigladiosus; Bernhauer, 1929: 143; Hammond, 1984: 209

Holotypus maschio etichettato come segue. I° Cartellino: Madang 1000 F., 2.11.19,
Sarawak, ded. Moulton, II°: 41. III°: Deroleptus bigladiosus Brnh. TYPUS, IV°: Chicago NH
Mus., M. Bernhauer collection.

Nota: Quando nel 1915 Bernhauer descrisse bigladiosus probabilmente non
conosceva il genere Orphnebius. Se lo avesse conosciuto non avrebbe attribuito questa

ORPHNEBIUS NEL BORNEO 745

FIGG. 1-7

Habitus, edeago in visione laterale e ventrale e spermateca. (1-4) Orphnebius borneanus sp. n.
(5-7) Orphnebius bigladiosus (Bernhauer), holotypus maschio.

746 R. PACE

specie ad Astilbus, ma a Orphnebius. La netta differenza della forma dell’addome di
bigladiosus lo ha indotto a creare Deroleptus, sottogenere di Astilbus. Nel 1929
Bernhauer dimostra di conoscere il genere Orphnebius. Dà una chiave per separare, tra
l’altro, Deroleptus da Orphnebius. Il carattere distintivo dato è «Hals hôchstens ein
Fiinftel so breit als der Kopf» per Deroleptus e «Hals mindestens ein Viertel so breit
als der Kopf» per Orphnebius. Un po’ poco per separare i due generi, tanto più che il
capo nelle specie di Orphnebius varia molto in larghezza sicché il rapporto larghezza
collo/larghezza del capo varia molto anche se si è in presenza di specie tra loro assai
affini, così riconosciute in base all’esame dell’edeago e della spermateca. L’edeago di
bigladiosus ha struttura che si ritrova in molte specie di Orphnebius. Risulta evidente
che Bernhauer si è arrampicato sugli specchi al fine di salvare dalla sinonimia il suo
Deroleptus. L'esame dell’holotypus mi ha permesso di constatare, soprattutto nella
forma dell’edeago, l'appartenenza di bigladiosus al genere Orphnebius. Pertanto viene
qui stabilita la seguente sinonimia:

Orphnebius Motschulsky, 1858: 253 (genotypus: O. ventricosus Motschulsky,
1858, della Colombia)

Deroleptus Bernhauer, 1915: 150 (genotypus: Deroleptus bigladiosus
(Bernhauer, 1915), del Borneo), syn. n.

Un mio esame di diverse specie di Orphnebius della Colombia, mi permette di
confermare la presenza di questo genere anche nella regione orientale.

Orphnebius bakeri Bernhauer, 1915 Figg. 39-41
Orphnebius (Mesocephalobius) bakeri Bernhauer, 1915: 150; Hammond, 1984: 211

Lectotypus maschio etichettato come segue. Cartellino I°: Singapore, Coll. Baker (a
stampa), II°: Orphnebius bakeri Brnh. TYPUS, HI°: Chicago NH Mus., M. Bernhauer collec-
tion. Figg. 39-41.

Paralectotypi: 1 maschio, Singapore, Coll. Baker, 12006, Orphnebius bakeri Brnh.,
Cotypus, Chicago Mus., 1 maschio, Sandakan, Borneo, Baker, Orphnebius bakeri Brnh.,
Cotypus, Chicago NH Mus., M. Bernhauer collection.

NOTA: Orphnebius bakeri Bernhauer è il genotypus del sottogenere
Mesocephalobius Bernhauer, 1929, designato da Blackwelder (1952). Unico carattere
distintivo è la presenza di capo largo quanto il pronoto o poco più stretto. Grazie
all'esame dell’edeago questa distinzione non ha alcun significato tassonomico. Una
specie con capo molto più largo del pronoto, O. borneanus sp. n., ha l’edeago con
caratteri simili a quelli dell’edeago di bakeri. Il medesimo grado di differenziazione
morfologica esterna si verifica nel genere Strabocephalium Bernhauer, 1911, che pre-
senta edeago simile a molte specie di Orphnebius. Il capo molto più largo del pronoto
anche in questo caso non è motivo sufficiente per mantenere ancora il genere, che
provvisoriamente va considerato sottogenere di Orphnebius a motivo della presenza di
occhi molto ridotti, mentre in Orphnebius gli occhi sono molto sviluppati.
Strabocephalium è inserito come sottogenere nella chiave data in seguito.

Va stabilita pertanto la seguente sinonimia:

Orphnebius Motschulsky, 1858: 253 (genotypus O. ventricosus Motschulsky,
1858, della Colombia)

Mesocephalobius Bernhauer, 1929: 146 (subgenotypus O. bakeri Bernhauer,
1929), syn. n.

ORPHNEBIUS NEL BORNEO 747

FIGG . 8-12

Spermateca, habitus e edeago in visione laterale e ventrale. (8) Orphnebius bigladiosus
(Bernhauer). (9-12) Orphnebius bakerianus Bernhauer, lectotypus femmina e paralectotypus
maschio.

748 R. PACE

Orphnebius bakerianus Bernhauer, 1915 Figg. 9-12

Orphnebius (Mesocephalobius) bakerianus Bernhauer, 1915: 148; Hammond, 1984: 211

Lectotypus femmina che porta i seguenti cartellini. I°: Sandakan, Borneo, Baker, II°:
Bakerianus Brnh Typ, HI°: Astilbus, IV: Orphnebius bakerianus Brnh Typus, IV°: Chicago NH
Mus., M. Bernhauer collection. Presente designazione, Fig. 12.

Paralectotypi: 2 maschi, stessa provenienza, cotypi, Chicago NH Mus., M. Bernhauer
collection.

Nora: Sia il maschio sia la femmina presentano il margine posteriore del quinto
urotergo libero trilobato.

Questa specie è stata attribuita al sottogenere Mesocephalobius, sinonimo di
Orphnebius.

Orphnebius quadricuspidatus Bernhauer, 1929

Orphnebius (Mesocephalobius) quadricuspidatus Bernhauer, 1929:149; Hammond, 1984: 211
Orphnebius quadrigladiosus Pace, 1986: 222, Figg. 247-249, syn. n.

Holotypus maschio, così etichettato. I° cartellino: Sandakan, Borneo, Baker. II° cartelli-
no: Orphnebius quadricuspidatus Brnh., TYPUS, II° cartellino: Chicago NH Mus., M.
Bernhauer collection.

NOTA: L’edeago dell’holotypus differisce di poco da quello di O. quadrigla-
diosus Pace, 1986. Presenta una lieve gibbosità della lama ventrale, assente in quadri-
gladiosus. Poiché l’armatura genitale interna dell’edeago di quadricuspidatus è estro-
flessa e quella di quadrigladiosus no, è difficile stabilire confronti. L’apice dell’edeago
di quadricuspidatus e di quadrigladiosus sono ugualmente assai stretti in visione
ventrale. Questo carattere mi ha permesso di proporre la sinonimia.

Orphnebius splendens Bernhauer, 1929 Figg. 19-20

Orphnebius (Mesocephalobius) splendens Bernhauer, 1929: 150; Hammond, 1984: 211

Holotypus femmina, etichettato come segue. I° Cartellino: Sandakan, Borneo, Baker. II°:
13453, IN°: Astilbus splendens B., IV°: Orphnebius splendens Brnh. TYPUS, V°: Chicago NH
Mus., M. Bernhauer collection.

NOTA: Nella descrizione Bernhauer si riferisce a un holotypus maschio, in base
agli sterniti liberi primo e secondo protratti all'indietro come spine, carattere normal-
mente riscontrabile nei maschi. Ho costatato che non è un maschio. Nonostante i
caratteri sessuali secondari maschili, la presenza della spermateca, Fig. 20, permette di
stabilire il corretto sesso dell’holotypus.

Orphnebius anguliceps Cameron, 1943 Figg. 24-25

Orphnebius (Megalocephalobius) anguliceps Cameron, 1943: 140; Hammond, 1984: 211
Holotypus femmina, Brunei, N. Borneo, Orphnebius anguliceps Cam. TYPE, (BMNH).

Nota: Nella sua descrizione Cameron non fa cenno del sesso dell’holotypus,
che da mio esame è una femmina, spermateca Fig. 25. Cameron attribuisce anguliceps
al sottogenere Megalocephalobius Bernhauer, 1929, per il capo più largo del pronoto.
La sua spermateca tuttavia ha forma simile a quella di specie che presentano capo più
stretto del pronoto come O. curticornis sp. n. e O. uniformis sp. n. Il subgenotypus di
Megalocephalobius è O. falagrioides Bernhauer, 1929, che non ho esaminato e di cui
Bernhauer non cita la località di provenienza. Cita e descrive questa specie nella

ORPHNEBIUS NEL BORNEO 749

FIGG. 13-18

Habitus e edeago in visione laterale e ventrale. (13-15) Orphnebius scalaris sp. n. (16-18)
Orphnebius directus sp. n.

750 R. PACE

chiave, facendola pero figurare descritta in altro lavoro, risultato inesistente anche per
Scheerpeltz (1934) e sconosciuta la sua localita tipica.

Orphnebius antennarius Bernhauer, 1929 Figg. 29-31

Orphnebius (Microcephalobius) antennarius Bernhauer, 1929: 153; Hammond, 1984: 211

Holotypus maschio, etichettato come segue. I° Cartellino: Sandakan, Borneo, Baker, II°:
Orphnebius antennarius Bernh., TYPUS UN. II°: Chicago NH Mus., M. Bernhauer collection.

NOTA: Il sottogenere Microcephalobius Bernhauer, 1929, ha come subgeno-
typus O. miricornis Bernhauer, 1926, delle Filippine. Non ho esaminato questa specie,
ma l’attribuzione di antennarius mi risulta ingiustificata se si confrontano gli edeagi.
Quello di antennarius ha struttura simile a quello di O. ideogramma sp. n. o di O.
penangensis sp. n. Queste specie presentano pronoto non trasverso come in anten-
narius. Di conseguenza dovrebbero essere collocati in sottogeneri differenti.

Nelle chiavi che seguono non è compresa la specie O. arachnoides Bernhauer,
1929, comb. n. ( da Deroleptus), affine a O. bigladiosus (Bernhauer, 1915), e della sua
stessa località tipica.

CHIAVE DEI MASCHI DELLE SPECIE DEL GENERE ORPHNEBIUS MOTSCHULSCKY, 1858, NEL
BORNEO

1 Apice dell’edeago, in visione ventrale, con una spina a ciascun lato,

Ei8728=Eunshez2a3,3:mmı(senzacapo)e rene O. biapicalis sp. n.
- Apice dell’edeago privo.di'spine laterality... ...:. lE 2
2 Capo pi largo del pronoto 7. i PIRRO LEE 3
- Capo. largo quanto il pronoto 0. pin stretto. 5... LE. „ee J
3 Occhi ridotti, molto più corti delle tempie; capo molto pit largo del pro-

noto Eunohezzas mme" ..... O. (Strabocephalium) mirabilis (Bernhauer)
- Occhi più lunghi delle tempie; capo poco più largo del pronoto ........... 4
4 Corpo rossiccio; antenne uniformemente giallo-rossicce; pronoto piü

sviluppato; edeago sinuato al lato ventrale. Lunghezza 3,7 mm

ER CO SOI, REVISION AO AS Gt O. platycephalus Pace
- Avancorpo rossiccio, addome giallo-rossiccio, antenne rossicce con base

e apice giallo-rossicci; pronoto meno sviluppato, Fig. 1; edeago arcuato

al lato:ventrale, Figs. 2-3. Lunghezza 5 mm” Serene O. borneanus sp. n.
5 Capo largo quanto il-pronoto®... ..............0 4. RR ROSEE 6
- Capo più stretto del pronoto . \....-. .... .... - 2. .2... foe ESTE 13
6 Antenne molto lunghe, con decimo antennomero più lungo che largo o

lungo quanto largo, Figs. 13 &16............. 2 m eee 7
- Antenne corte, con decimo antennomero fortemente trasverso, Figg. 13 e 16 . 8
7 Decimo antennomero più lungo che largo; capo e pronoto rossicci; gra-

nulosità delle elitre forte; margine posteriore del sesto urotergo libero

del maschio con due denti a ciascun lato del lobo mediano. Lunghezza

ASTM eR. | a OSANO re del RATER O. bigladiosus (Bernhauer)
- Decimo antennomero lungo quanto largo; capo e pronoto giallo-rossicci;

granulosità delle elitre superficiale; margine posteriore del sesto uro-

tergo libero del maschio senza denti laterali al lobo mediano. Lunghezza

Sil EI UE MMS Es TS EURE O. bakerianus Bernhauer

ORPHNEBIUS NEL BORNEO 751

1 mm

Fico. 19-24

Habitus, spermateca e edeago in visione laterale e ventrale. (19-20) Orphnebius splendens
Bernhauer, holotypus. (21-23) Orphnebius minor sp. n. (24) Orphnebius anguliceps Cameron,
holotypus.

8 Capo e pronoto neri; ciascun lato dell’addome con due spine; apice
dell’adeago strettissimo e lunghissimo, in visione ventrale. Lunghezza
NM. ISIN ZONE Er Le: O. quadricuspidatus Bernhauer

: Capo e pronoto bruni o giallo-rossicci, apice dell’edeago mai contem-
poraicamenterstrettoielun ORRORE CRE EC PRE 9

152

10

idl

12

13

14

IS

16

17

18

R. PACE

Addome bruno-rossiccio; undicesimo antennomero totalmente o par-
zialmentegiallo-rossiccio tè 21.2 Orto Rei 10
Addome giallo-rossiccio; undicesimo antennomero bruno ............... 11
Undicesimo antennomero giallo-rossiccio solo nella porzione distale;
quinto urotergo libero fortemente punteggiato; granulosità delle elitre
superficiale; edeago lungo e stretto, Figg. 14-15. Lunghezza 3,4 mm
te due ce à ou eos Pei oo. O. scalaris sp. n.
Undicesimo antennomero interamente giallo-rossiccio; quinto urotergo

libero senza punteggiatura, ma con tubercoli posteriori; granulosità delle

elitre saliente; edeago corto e largo, Figg. 17-18. Lunghezza 2,2 mm
ee RIOT O. directus sp. n.
Capo e pronoto giallo-rossicci; sutura delle elitre più corta del pronoto;

apice dell’edeago, in visione ventrale, stretto, Figg. 30-31. Lunghezza

HAE ER Re Ne lac i ea, O. antennarius Bernhauer
Capo bruno, pronoto giallo-rossiccio o bruno-rossiccio; sutura delle

elitre più lunga del pronoto; apice dell’edeago largo, in visione ventrale . . . 12
Pronoto bruno-rossiccio e poco trasverso; edeago Figg. 33-34:
Éungnezza 54 min, cs yc RIE O. ideogramma sp
Pronoto giallo-rossiccio e fortemente trasverso; edeago Figg. 22-23.

APPIA RE, i) e » oc Sivas Geb as Cy ae a ed oe O. minor sp. n.
Corpo nero-bruno con addome marginato di rossiccio; granulosita delle

elitre forte e fitta; edeago a lati paralleli, in visione ventrale. Lungh.

STI ees ee TRE RE GO RA OA PRE O. derougemonti Pace
Corpo bruno e giallo-rossiccio; granulosità delle elitre fine: edeago a lati

parallelisimivisione ventrale . ........... II 14
Addome;piall0-FOSSICCIO: Wii. RL eee 15
Addomebruno,o:bruno-Tossiceio. ERRE uh oe 18

Antenne giallo-rossicce; quarto urotergo libero del maschio sinuoso al
margine posteriore; edeago Figg. 54-55. Lunghezza 3,8 mm

PR Ns, APTE Ne | ae ie nee Se NE ee c O. perpenetrans Sp. n.
Antenne bruno-rossicce con base rossiccia o giallo-rossiccia; quarto uro-

tergo libero del maschio rettilineo al margine posteriore ................ 16
Capo e pronoto bruni, elitre rossicce; porzione posteriore del quinto uro-

tergo libero del maschio con quattro tubercoli salienti; edeago Figg.
40-41, Punshezza mm, PE ri RELA AE O. bakeri Bernhauer
Capo, pronoto ed elitre giallo-bruni o bruno-rossicci; porzione
posteriore del quinto urotergo libero del maschio senza tubercoli: al loro

posto. vi €.una rugosita longitudinale -..-..... ........ 2.202.120. ee 17
Pronoto più trasverso; edeago non sinuato ai lati, in visione ventrale,
610955859 Lunghezza;4 mme er EE O. krypticola sp. n.
Pronoto meno trasverso; edeago sinuato ai lati, in visione ventrale, Figg.

43 AA alsunshezz2a 37mm. este lee E BOSE O. bajauorum sp. n.
Undicesimo antennomero uniformemente bruno; pronoto lievemente
trasverso; pigidio giallo-rossiccio; edeago Figg. 37-38. Lunghezza
SPAM sut ee EES SRE er ee ee eee O. penangensis sp. n.

ORPHNEBIUS NEL BORNEO 753

DEREN

D SET DE WORT LIT ITR —

2

oD PAU QE Se “ BEN
DE D ÉTÉ Deposer ge y

2

23225359

Fico. 25-28

Spermateca, habitus e edeago in visione laterale e ventrale. (25) Orphnebius anguliceps
Cameron, holotypus (26-28) Orphnebius biapicalis sp. n.

754

R. PACE

Undicesimo antennomero bicolore bruno e giallo-rossiccio; pronoto
nettamente trasverso; pigidio bruno-rossiccio . . ........ 19
Corpo uniformemente rossiccio; pronoto poco trasverso; quinto urotergo
libero del maschio senza tubercoli lungo il margine posteriore; edeago
Bises 0 SIN EuUnLSheZzaS 2 mine. see eee eee O. uniformis sp. n.
Avancorpo bruno, addome bruno-rossiccio; pronoto molto trasverso;
quinto urotergo libero del maschio con tubercoli lungo il margine poste-
riore; edeago Figg. 46-47. Lunghezza 2,7 mm.......... O. curticornis Sp. n.

KEY TO MALES OF THE SPECIES OF THE GENUS ORPHNEBIUS MOTSCHULSCKY, 1858, FROM

BORNEO
1 Apex of the aedeagus, in ventral view, with a thorn to every lateral

margin, Fig. 28. Length 3.3 mm (without head) .......... O. biapicalis sp. n.
- Apex of the aedeagus deprived.of lateral thorns . . 22... we 2
2 Head wider. thanıthe pronotum ......... =: „acc I Se eee 5
- Head as wide'as'the pronotum or more narrow : 2... +. 2... 2 soe oe 5
3 Eyes reduced, very more courts of the temples; head very broader than

the pronotum. Length 5 mm ..... O. (Strabocephalium) mirabilis (Bernhauer)
- Eyes longer than the temples: head a little more breadth than the pronotum . . 4
4 Body reddish; antennae uniformly yellow-reddish; pronotum more

developed; aedeagus sinuous to the ventral margin. Length 3.7 mm

WERE ORT N tte ich aa ee O. platycephalus Pace
- Fore-parts reddish, abdomen yellow-reddish, antennae reddish with their

base and apex yellow-reddish; pronotum less developed, Fig. 1; aedea-

gus arched to the ventral side, Figs 2-3. Length 5 mm... . 0. borneanus sp. n.
5 Head. as: broad. asithe;pronotum.. -.......-.4.... . (43 eee 6
- Headinarrower thanithe pronotum!).......... «...: «Ae. COSROE 13
6 Antennae very long, with tenth antennomere longer than wide or as long

as wide, ‘Figs: 13) and Gr date ee RE 7
- Antennae short, with tenth antennomere strongly transverse, Figs 13 and 16.. 8
U Tenth antennomere longer than wide; head and pronotum reddish;

granularity of the elytra strong; posterior border of the sixth free uro-

tergum of the male with two teeth to every lateral margin of the median

lobes.Eensth,4.7: mm... ere Sr O. bigladiosus (Bernhauer)
- Tenth antennomere as wide as long; head and pronotum yellow-reddish;

granularity of the elytrae superficial; posterior border of the sixth free

urotergum of the male without teeth lateral to the median lobe. Length

SH LOL Lo SELLES I AR Ra hai O. bakerianus Bernhauer
8 Head and pronotum black; every lateral margin of the abdomen with two

thorns; apex of the aedeagus narrow and long, in ventral view. Length

PAIN: re ee cae O. quadricuspidatus Bernhauer
- Head and pronotum brown or yellow-reddish, apex of the aedeagus

never contemporarily narrowsand long 5.4 tue. CCE eee 9
9 Abdomen brown-reddish; eleventh antennomere totally or partially

yellow- Teddish........... 2c tags ter AR de EEE PEER ER 10

ORPHNEBIUS NEL BORNEO 755

33 34

FiGG. 29-34

Habitus e edeago in visione laterale e ventrale. (29-31) Orphnebius antennarius Bernhauer,
holotypus maschio. (32-34) Orphnebius ideogramma sp. n.

- Abdomen yellow-reddish; eleventh antennomere brown ................ al
10 Eleventh antennomere yellow-reddish only in the distal portion; fifth
free urotergum deeply punctured; granularity of the elytra superficial;

aedeagus long and narrow, Figs 14-15. Length 3.4 mm ..... O. scalaris sp. n.

756

ul

12

13

14

IS

16

19

R. PACE

Eleventh antennomere entirely yellow-reddish; fifth free urotergum with-
out punctuation, but with posterior tubercles; granularity of the elytra
salient; aedeagus short and wide, Figs 17-18. Length 2.2 mm . O. directus sp. n.
Head and pronotum yellow-reddish; suture of the elytra shorter than the
pronotum; apex of the aedeagus narrow, in ventral view, Figs 30-31.
LENIRE RR NEE SES O. antennarius Bernhauer
Head brown, pronotum yellow-reddish or brown-reddish; suture of the
elytra longer than the pronotum; apex of the aedeagus broad, in ventral

VO RSR RS N a a 3G) N CET TETTE WZ
Pronotum brown-reddish and few transverse; aedeagus Figs 33-34:
TEMERE AAA fas kas se ee O. ideogramma sp. n.
Pronotum yellow-reddish and strongly transverse; aedeagus Figs 22-23.
Rengih 2mm. VESTI Une IE a A er O. minor Sp. n.

Body black-brown with abdomen margined of reddish; granularity of the
elytra strongly sharp and close; lateral margins of the aedeagus parallel,

insventralsview: Eunsh3:mm Ar e O. derougemonti Pace
Body brown and yellow-reddish; granularity of the elytra fine: lateral

margins of the aedeagus parallel, in ventral view... 22.22.22 ee 14
Abdomen: yellow-reddish™ ......... 2.10 Saat EN PRE ET EEE 15
Abdomenibrown:or.brown-reddish.. .......... 2.2.2.2... ERP EEE 18

Antennae yellow-reddish; fourth free urotergum of the male sinuous to

the posterior border; aedeagus Figs 54-55. Length 3.8 mm

RE de ae Cel e A O. perpenetrans sp. n.
Antennae brown-reddish with reddish or yellow-reddish base; fourth

free urotergum of the male to the posterior border rectilinear ............ 16
Head and pronotum brown, elytra reddish; posterior portion of the fifth

free urotergum of the male with four salient tubercles; aedeagus Figs

AQ AT LensthA.mm. I... ERS ob Lula CNE PR bakeri Bernhauer
Head, pronotum and elytra yellow-brown or brown-reddish; posterior
portion of the fifth free urotergum of the male without tubercles: to their
place-there is alongitudimal roughness. ..... ... 2. PRE CE TT 17
Pronotum transverse; aedeagus not sinuous to the lateral margins, in
ventral view, Figs 58-59. Length 4 mm. Brunei.......... O. krypticola sp. n.
Pronotum less transverse; aedeagus sinuous to the lateral margins, in
ventral view, Figs 43-44. Length 3.7 mm.............. O. bajauorum sp. n.
Eleventh antennomere uniformly brown; pronotum slightly transverse;
pigidium yellow-reddish; aedeagus Figs 37-38. Length 3.2 mm

SME ONTO SIRIA O. penangensis Sp. n.
Eleventh antennomere bicolorous brown and yellow-reddish; pronotum
clearly transverse; pigidium brown-reddish' CR SARE 19
Body reddish uniformly; pronotum a little transverse; fifth free uro-
tergum of the male without tubercles along the posterior border;
aedeasus.Fios:S0-SiIM Length 2 mm are RESSE eee O. uniformis sp. n.
Foreparts brown, abdomen brown-reddish; pronotum very transverse;

fifth free urotergum of the male with tubercles along the posterior
border; aedeagus Figs 46-47. Length 2.7 mm........... O. curticornis sp. n.

ORPHNEBIUS NEL BORNEO 757

FıGG. 35-38

Spermateca, habitus e edeago in visione laterale e ventrale. (35) Orphnebius ideogramma sp. n.;
(36-38) Orphnebius penangensis sp. n.

CHIAVE DELLE FEMMINE DELLE SPECIE DEL GENERE ORPHNEBIUS MOTSCHULSCKY, 1858,
NEL BORNEO

1 Base delle mandibole sporgente di sotto agli occhi, Fig. 24; spermateca

Eie2 JaEunshezza2 0m 1 VR O. anguliceps Cameron
= Base delle mandibole non sporgente di sotto agli occhi................. 2
2 Capo più largo del pronoto; spermateca con bulbo prossimale sub-

SICHICOME19 94 NE unshezzaSmn EEE O. borneanus sp. n.

758

Wo!

PS

R. PACE

©aporlargo quanto il'pronoto o piùistretto ft. MERE RESA e 3
Capo largo quanto: il pronoto wie. RCE rr SNA OR 4
Capoypiùistretto delipronoto: 2. u... 2.2.2. LORO. RR RSA EE 8
Antenne molto lunghe, con decimo antennomero piü lungo che largo o lungo
quantollarsorFigg. I3e lo... ee. See 5
Antenne corte, con decimo antennomero fortemente trasverso ............ 6

Decimo antennomero più lungo che largo; capo e pronoto rossicci; gra-
nulositä delle elitre forte; spermateca piegata a Z, Fig. 8. Lunghezza
ARTRITE > Keen + stra dts WE Ge OR © aS O. bigladiosus (Bernhauer)
Decimo antennomero lungo quanto largo; capo e pronoto giallo-rossicci;
granulosita delle elitre superficiale; spermateca piegata a 7, Fig. 12.
Éunehezzas lemme" 2 a ee ee ae O. bakerianus Bernhauer
Addome rossiccio o bruno-rossiccio; bulbo distale della spermateca
foremente tlesso4lEunshezza 27mm 211 2 A Ei O. laetus Pace
Addome giallo-rossiccio; bulbo distale della spermateca non flesso ........ 7
Pronoto meno trasverso; base delle antenne rossiccia; bulbo distale della
spermateca con profonda introflessione apicale Fig. 35. Lunghezza
SD JUV ler isaac OR RS PE I O. ideogramma sp. n.
Pronoto più trasverso; base delle antenne giallo-rossiccia; bulbo distale
della spermateca con debole introflessione apicale, Fig. 20: Lunghezza

sesh TU NE arin aa ee Wa | eee RE O. splendens Bernhauer
Corpo giallo-rossiccio; bulbo distale della spermateca molto sviluppato,
Fig AGO Bunshezza2 mm" a ween En O. crassus Sp. n.
Corpo unicolore o bicolore nero-bruno, rossiccio o bruno-rossiccio;
bulbo'distale. della spermateca poco/sviluppato CCC 9
Antenne uniformemente giallo-rossicce; addome giallo-rossiccio; sper-
MateCa Figs yo, LUNPMEZZA S,6 MIM ar. Ur. O. perpenetrans sp. n.

Antenne bicolori brune o rossicce con base o apice distale giallo-rossicci . . 10
Undicesimo antennomero bruno, avancorpo nero-bruno; parte interme-

dia della spermateca molto lunga. Lunghezza 3 mm. . . O. derougemonti Pace
Undicesimo antennomero interamente o parzialmente giallo-rossicce;
parte intermedia della spermateca breve ORRORE SSA 11
Corpo uniformemente rossiccio; parte prossimale della spermateca des-
crivente numerosi meandri, Fig. 52. Lunghezza 3,2 mm. . . O. uniformis sp. n.
Corpo bicolore bruno e rossiccio; parte prossimale della spermateca
descrivente due sinuosita.. rl ITA 12

Solo l’apice dell’undicesimo antennomero é giallo-rossiccio; avancorpo
bruno; quinto urotergo libero della femmina con rugosità longitudinali;
bulbo distale della spermateca privo di introflessione apicale, Fig. 48.
unghezza 2 7. mm... erstere RE ES Tr O. curticornis Sp. n.
Undicesimo antennomero interamente giallo-rossiccio; capo e pronoto
bruni, elitre rossicce; quinto urotergo libero della femmina con forte
punteggiatura; bulbo distale della spermateca con profonda intro-
flessionetapicale- Lunghezza 2:S nm ke Re O. silvarum Pace

ORPHNEBIUS NEL BORNEO 759

Fısc. 39-41
Habitus e edeago in visione laterale e ventrale. (39-41) Orphnebius bakeri Bernhauer, holotypus.

KEY TO FEMALES OF THE SPECIES OF THE GENUS ORPHNEBIUS MOTSCHULSCKY, 1858, FROM

BORNEO
1 Base of the mandibles protruding under the eyes, Fig. 24; spermatheca

Rice allen sth 22 Omni wey ar Aer ete O. anguliceps Cameron
= Base of the mandibles non protruding under the eyes ................... 2
2 Pronotum wider than the head; spermatheca with proximal bulb sub-

Sphencal #19 AIN SO IMMERSIONE O. borneanus sp. n.
= Pronotwm) as wideras the head or more NATOWE see RECON ee 3
3 Headtasawidevassthe pronOtuinien = “ae eee en oe ee ote À
= Headmnarrowerthankthespronomumin peer sete IA I eee E 8
4 Antennae very long, with tenth antennomere longer than wide or as long

as wide Fos Span GE 6 pregio: dress ab res cule creas a ae ager yi 5

Antennae short, with tenth antennomere strongly transverse .............. 6

760 R. PACE

I Tenth antennomere longer than wide; head and pronotum reddish;
granularity of the elytra strong; spermatheca folded up as Z Fig. 8.
EIA] IRA, en A NN O. bigladiosus (Bernhauer)

- Tenth antennomere as wide as long; head and pronotum yellow-reddish;
granularity of the elytra superficial; spermatheca folded up as 7 Fig. 12.

Bensths.lkmmeear hate sale ur O. bakerianus Bernhauer
6 Abdomen reddish or brown-reddish; distal bulb of the spermatheca

stronsiyHexsEensth27mm PESCE O. laetus Pace
- Abdomen yellow-reddish; distal bulb of the spermatheca not flex......... 7
7 Pronotum less transverse; base of the antennae reddish; distal bulb of the

spermatheca with apical umbilicus deep Fig. 35. Length 3.4 mm

SUN RE VIELE 2 de ue A O. ideogramma sp. n.
- Pronotum transverse; base of the antennae yellow-reddish; distal bulb of

the spermatheca with apical umbilicus weak, Fig. 20: length 2.8 mm

TRA SO NE N Ins Fa tate sf lee Se GE O. splendens Bernhauer
8 Body yellow-reddish; distal bulb of the spermatheca very developed,

FisaG0MÉEnsth 2 amis ER lee ee eee O. crassus Sp. n.
- Body unicoloured or bicoloured black-brown, reddish or brown-reddish;

distal’ bulb of the spermatheca a little developed <>... 22. Sarg 9
9 Antennae uniformly yellow-reddish; abdomen yellow-reddish; sper-

mathecalbie. 56. Wenethy3:8 mm rar. O. perpenetrans Sp. n.
- Antennae bicoloured brown or reddish with base or distal apex yellow-

RE LS PER A PRE Root TTT OE 10
10 Eleventh antennomere brown, fore-parts black-brown; intermediary

portion of the spermatheca very long. Length 3mm.... 0. derougemonti Pace
- Eleventh antennomere entirely or partially yellow-reddish; intermediary

portion ofthe spennatheca shorts. 5......--.-..4.. So See 11
11 Body uniformly reddish; proximal portion of the spermatheca that

describes numerous meanders, Fig. 52. Length 3.2 mm .... ©. uniformis sp. n.
- Body bicoloured brown and reddish; proximal portion of the sper-

mathecaithat describes:two sinuosity... 27 24.732. 222 ES SE 12

12 Only the apex of the eleventh antennomere is yellow-reddish; fore-parts
brown; fifth free urotergum of the female with longitudinal roughness;
distal bulb of the spermatheca without apical umbilicus, Fig. 48. Length
Dimmi arie A SRE TE Se ee O. curticornis sp. n.

= Eleventh antennomere entirely yellow-reddish; head and pronotum
brown, elytra reddish; fifth free urotergum of the female with strong
punctuation; distal bulb of the spermatheca with deep apical umbilicus.
Ikeneth 2: Simm iu. gen, ER EEE O. silvarum Pace

Orphnebius borneanus sp. n. Figg. 1-4

Holotypus maschio, Borneo-Sabah, Crocker Ra., 1550-1650 m, 16.V.1987, leg.
Burckhardt & Löbl (IRSB).

Paratypi: 1 femmina, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ 1560 m, 30.IV.1987,
leg. A. Smetana; 1 femmina, Sabah, Mt. Kinabalu, 1450-1550 m, 23.IV.1987, leg. Burckhardt &
Löbl.

ORPHNEBIUS NEL BORNEO 761

TP

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i
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FIGG. 42-47

Habitus e edeago in visione laterale e ventrale. (42-44) Orphnebius bajauorum sp. n. (45-47)
Orphnebius curticornis sp. n.

DESCRIZIONE: Lunghezza 5 mm. Corpo lucido e rossiccio con addome giallo-
rossiccio; antenne rossicce con i due antennomeri basali e la base del terzo giallo-
rossicci e apice dell’undicesimo giallo; zampe rossicce con femori giallo-rossicci. Il
corpo non è coperto di reticolazione. Il capo e il pronoto non presentano punteggiatura,

762 R. PACE

né granulosità. La granulosità delle elitre & ben visibile. Gli uroterghi sono nudi, tranne
il quinto libero con punteggiatura evidente e profondo solco basale trasverso. Il capo è
molto più largo del pronoto, Fig. 1. Edeago Figg. 2-3, spermateca Fig. 4.

Orphnebius bigladiosus (Bernhauer, 1915) Figg. 5-7

Astilbus (Deroleptus) bigladiosus Bernhauer, 1915: 150
Deroleptus bigladiosus; Bernhauer, 1929: 143; Hammond, 1984: 209
Orphnebius bigladiosus; Pace, hoc opus

1 maschio e 1 femmina, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480-510 m,
30.VIII.1988, leg. A. Smetana.

Orphnebius scalaris sp. n. Figg. 13-15

Holotypus maschio, Sabah, Poring Hot Springs, 500 m, 8.V.1987, leg. Burckhardt &
Löbl (MHNG).

DESCRIZIONE: Lunghezza 3,4 mm. Corpo lucido e bruno-rossiccio con metà
posteriore delle elitre bruna; antenne brune con i tre antennomeri basali e la metà
apicale dell’undicesimo giallo-rossicci; zampe rossicce. Il corpo non è coperto di
reticolazione. La punteggiatura del capo e del pronoto e la granulosità delle elitre sono
superficiali. Gli uroterghi sono nudi, tranne il quinto libero con punteggiatura evidente
e profondo solco basale trasverso. Il capo è largo quanto il pronoto. Edeago Figg.
14-15.

DERIVATIO NOMINIS: Il nome delle nuova specie deriva dalle espansioni laterali
basali dell’addome disposte come 1 gradini di una scala.

Orphnebius directus sp. n. Figg. 16-18

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480 m,
10.V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 3,4 mm. Corpo lucido e bruno-rossiccio con elitre
giallo-brune; antenne brune con i tre antennomeri basali e l'undicesimo giallo-rossicci;
zampe giallo-rossicce. Il corpo è privo di reticolazione. La punteggiatura del capo è
fine e distinta. La granulosità del pronoto è molto saliente sul disco: sul resto dello
stesso pronoto è superficiale. La granulosità delle elitre è molto saliente e forte. Il
quinto urotergo libero del maschio presenta un tubercolo mediano posteriore. Il capo è
largo quanto il pronoto. Edeago Figg. 17-18.

DERIVATIO NOMINIS: Il nome della nuova specie deriva dalla parte distale
dell’edeago retta ventralmente, in visione laterale.

Orphnebius minor sp. n. Figg. 21-23

Holotypus maschio, Borneo-Sabah, Poring Hot Springs, 500 m, 13.V.1987, leg.
Burckhardt & Löbl (MHNG).

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido e giallo-rossiccio, con capo
bruno-rossiccio ed elitre rossicce; antenne rossicce con i due antennomeri basali e la
base del terzo giallo-rossicci; zampe rossicce con femori giallo-rossicci. Sul corpo non
è presente una reticolazione. Il capo e il pronoto non sono punteggiati, né granulosi. Il
capo è largo quanto il pronoto. Gli ommatidi sono grossolani. La granulosità delle

ORPHNEBIUS NEL BORNEO 763

Sa yy

0,1 mm

FIGG. 48-52
Spermateca, habitus e edeago in visione laterale e ventrale. (48) Orphnebius curticornis sp. n.
(49-52) Orphnebius uniformis sp. n.

elitre è distinta e moderatamente fitta. Il quinto urotergo libero del maschio è coperto
di forte punteggiatura, tranne che alla sua base dove è assente. Edeago Figg 22-23.

DERIVATIO NOMINIS: Il nome di «più piccola» della nuova specie deriva dalla
taglia corporea esigua a confronto con altre grandi specie del Borneo.

764 R. PACE

Orphnebius biapicalis sp. n. Figg. 26-28

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1540 m,
14.VIII-1.1X.1988, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 3,3 mm. Corpo lucido e rossiccio; zampe giallo-
rossicce. Il capo manca perché perduto nella raccolta. Il pronoto presenta punti isolati
superficiali disposti come da Fig. 26. Le elitre non presentano punteggiatura o granu-
losità. Il quinto urotergo libero del maschio mostra strie longitudinali, assai profonde
verso la base dello stesso urotergo libero. Edeago con due appendici apicali, Figg.
27-28.

DERIVATIO NOMINIS: Il nome di «due apici» della nuova specie deriva da quelli
dell’edeago, in visione ventrale.

Orphnebius ideogramma sp. n. Figg. 32-35

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,
12.V.1987, leg. A. Smetana (MHNG).

Paratypi: 1 femmina, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480-510 m,
30.V111.1988, leg. A. Smetana; 1 femmina, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs,
510 m, 13.V.1987, leg. A. Smetana; 1 femmina, Sabah, Poring Hot Springs, 500 m, 6.V.1987,
leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 3,3 mm. Corpo lucido e giallo-rossiccio con capo
bruno, pronoto bruno-rossiccio, elitre rossicce; antenne brune con i tre antennomeri
basali rossicci; zampe rossicce con femori giallo-rossicci. Il capo mostra tre punti, in
cui è inserita una setola, presso gli occhi. Il pronoto presenta due punti disposti come
da Fig. 32. La granulosità delle elitre è fine e distinta. Il quinto urotergo libero del
maschio è coperto di netta punteggiatura fitta solo sulla metà posteriore. Presenta una
profonda impressione basale trasversa. Il capo è largo quanto il pronoto che è poco
trasverso, Fig. 32. Edeago Figg. 33-34, spermateca Fig. 35.

DERIVATIO NOMINIS: L’armatura genitale interna dell’edeago, in visione laterale,
ha una forma che ricorda vagamente un ideogramma cinese, Fig. 33. Da esso la nuova
specie prende nome.

Orphnebius penangensis sp. n. Figg. 36-38

Holotypus maschio, Island of Penang, Baker, cotypus di Orphnebius bakeri Bernhauer
(FMNH). Designato il lectotypus di O. bakeri, grazie all'esame dell’edeago ho isolato la
presente specie.

DESCRIZIONE: Lunghezza 3,3 mm. Corpo lucido e bruno con addome bruno-
rossiccio e pigidio rossiccio; antenne brune con i tre antennomeri basali rossicci;
zampe bruno-rossicce con tarsi rossicci. Sul corpo la reticolazione è assente, tranne che
alla base degli uroterghi liberi. Il capo e il pronoto sono privi sia di punteggiatura che
di granulosità. La granulosità delle elitre è così fine che è poco visibile. Il quinto uro-
tergo libero del maschio è coperto di rugosità longitudinale tra cui si trovano dei punti.
Il capo è poco più stretto del pronoto che è lungo quanto largo. Edeago Figg. 37-38.

Orphnebius bajauorum sp. n. Figg. 42-44

Holotypus maschio, N. Borneo, Sabah, Bunsit Keningan, 31.VII.1985, leg.
K. Maruyama (DEI).

ORPHNEBIUS NEL BORNEO 765

56 de

S

0,1 mm

FIGG. 53-56

Habitus, edeago in visione laterale e ventrale e spermateca. (53-56) Orphnebius perpenetrans
sp. n.

DESCRIZIONE: Lunghezza 3,7 mm. Corpo lucido con l’avancorpo bruno-
rossiccio e l’addome giallo-rossiccio; antenne brune con i quattro antennomeri basali
giallo-rossicci; zampe gialle. Nessuna traccia di reticolazione sul corpo. Il capo e il
pronoto sono privi di punteggiatura. Solo il primo urotergo libero è coperto di granu-
losità molto superficiale, gli altri non hanno granulosità. La punteggiatura del quinto
urotergo libero è distinta, posteriormente confusa in una rugosità. Edeago Figg. 43-44.

DERIVATIO NOMINIS: La nuova specie prende nome dal gruppo etnico dei Bajau
del Borneo.

Orphnebius curticornis sp. n. Figg. 45-48

Holotypus maschio, Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg. Burckhardt &
Löbl (MHNG).

Paratypi: 2 maschi e 1 femmina, stessa provenienza dell’holotypus.

766 R. PACE

DESCRIZIONE: Lunghezza 2,7 mm. Corpo lucido e bruno con addome bruno-
rossiccio; antenne nero-brune con 1 tre antennomeri basali, base del quarto e apice
dell’undicesimo giallo-rossicci; zampe rossicce. Il corpo non ha reticolazione. Il capo
presenta un solo punto presso l’orbita di ciascun occhio. Il pronoto presenta due punti
post-discali molto superficiali. La granulositä delle elitre & distinta. Il quinto urotergo
libero del maschio é coperto di strie superficiali, assenti alla base dello stesso urotergo
libero, e mostra alcuni granuli forti lungo il margine posteriore. Edeago Figg. 46-47,
spermateca Fig. 48.

DERIVATIO NOMINIS: La nuova specie prende nome di «antenne corte» per averle
appunto corte rispetto quelle di altre specie del Borneo.

Orphnebius uniformis sp. n Figg. 49-52

Holotypus maschio, Borneo, Sabah, Crocker Range, 1600 m, 18.V.1987, leg. Burckhardt
& Löbl (MHNG).

Paratypi: 6 femmine, stessa provenienza dell’holotypus; 1 femmina, Sabah, Crocker
Range, 1600 m, km 51 road Kota Kinabalu- Tambunan, 18.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 3,2 mm. Corpo lucido e rossiccio; antenne rossicce
con i tre antennomeri basali giallo-rossicci e apice dell’undicesimo giallo; zampe
rossicce. Corpo senza reticolazione. Sul capo e sulle elitre la punteggiatura o la granu-
losità è assente. Il pronoto presenta solo due punti discali, Fig. 49. Il quinto urotergo
libero del maschio è coperto di strie longitudinali fino al solco trasverso basale. Il capo
è più stretto del pronoto. Edeago Figg. 50-51, spermateca Fig. 52.

DERIVATIO NOMINIS: La nuova specie prende nome dal colore uniforme del
corpo.

Orphnebius perpenetrans sp. n. Figg. 53-56

Holotypus maschio, Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1565 m,
3.VIIL.1988, leg. A. Smetana (MHNG).

Paratypi: 2 maschi, Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1540 m,
14.VIII-1.1X.1988, leg. A. Smetana; 2 maschi, Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ 1500
m, 8-16.1V.1987, leg. A. Smetana; 3 es., Sabah, Crocker Ra., 1550-1650 m, 16.V.1987, leg.
Burckhardt & Löbl; 6 es., Borneo, Sabah, Mt. Kinabalu N.P., 1550 m, 29.IV.1987, leg.
Burckhardt & Löbl; 6 es., Borneo, Sabah, Crocker Ra., 1600 m, Km 51 rte. Kinabalu-Tambunan,
18.V.1987, leg. Burckhardt & Löbl; 6 es., Sabah, Mt. Kinabalu, 1500 m, 21.V.1987, leg.
Burckhardt & Löbl; 3 es., 12 es., Sabah, Mt. Kinabalu, 1550 m, 28.IV.1987, leg. Burckhardt &
Löbl; 3 es., Sabah, Mt Kinabalu, 1500m, 21.V.1987, leg. Burckhardt-Löbl; 1 es., Sabah, Mt
Kinabalu, 1500m, 25.V.1987, leg. Burckhardt-Löbl.

DESCRIZIONE: Lunghezza 3,2 mm. Corpo lucido. Avancorpo bruno, addome
giallo-rossiccio; antenne e zampe giallo-rossicce. Sul corpo non vi € traccia di retico-
lazione. Il capo è senza punteggiatura. il pronoto presenta due deboli punti discali e due
con setola, Fig. 53. La punteggiatura delle elitre è fine. Il margine posteriore del quarto
urotergo libero del maschio è incavato a metà. Il quinto urotergo libero del maschio e
della femmina è coperto di strie longitudinali punteggiate che non raggiungono la base
dello stesso urotergo libero che ha un solco basale trasverso. Il capo è più stretto del
pronoto che è poco trasverso. Edeago Figg. 54-55, spermateca Fig. 56.

DERIVATIO NOMINIS: La nuova specie prende nome dalla punta acuta e ricurva

dell’edeago che ha la funzione di penetrare agevolmente nel dotto vaginale della
femmina.

ORPHNEBIUS NEL BORNEO 767

Fico. 57-61

Habitus, edeago in visione laterale e ventrale e spermateca. (57-59) Orphnebius krypticola sp. n.
(60-61) Orphnebius crassus sp. n.

Orphnebius kripticola sp. n. Figg. 57-59

Holotypus maschio, Borneo, Brunei, Temburong, Kuala Belalong (West), 14.11.1995,
leg. Borcherding (IRSN).

768 R. PACE

DESCRIZIONE: Lunghezza 4 mm. Corpo lucido senza reticolazione. Avancorpo
giallo-bruno, addome giallo-rossiccio; antenne nero pece con 1 tre antennomeri basali
giallo-rossicci; zampe gialle. Il capo e il pronoto non presentano punteggiatura o
granulosità, tranne quattro punti isolati sul pronoto distribuiti come da Fig. 57. La
granulosita delle elitre è rada e distinta. Il quinto urotergo libero del maschio presenta
punteggiatura anteriore e una rugosità posteriore mediane assenti ai lati dello stesso
urotergo libero e alla sua base che è profondamente incavata. Il capo è più stretto del
pronoto. Edeago Figg. 58-59.

DERIVATIO NOMINIS: La nuova specie prende nome di «abitatrice dei luoghi
nascosti».

Orphnebius crassus sp. n. Figg. 60-61

Holotypus femmina, Sabah, Poring Hot Springs, 600 m, nr. Bat Cave, 10.V.1987, leg.
Burckhardt & Löbl (MHNG).

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e giallo-rossiccio; antenne
giallo-rossicce con 1 tre antennomeri basali gialli; zampe gialle. Manca una retico-
lazione sulla superficie del corpo. Il capo non possiede punteggiatura. Il pronoto
presenta solo quattro punti distinti distribuiti come da fig. 61. La punteggiatura delle
elitre è molto superficiale. Il quinto urotergo libero della femmina è punteggiato
anteriormente e striato longitudinalmente nella porzione distale. Alla base di questo
urotergo libero manca la punteggiatura ed è presente un largo solco basale. Il capo è
molto più stretto del pronoto che è fortemente trasverso, Fig. 61. Spermateca Fig. 60.

DERIVATIO NOMINIS: La nuova specie prende nome di «pingue» a motivo del suo
corpo tozzo che simula pinguedine.

RINGRAZIAMENTI

Rivolgo i miei più cordiali ringraziamenti a coloro che mi hanno affidato in
studio il materiale oggetto del presente studio: il Dr. Ales Smetana di Ottawa, il Dr.
Ivan Löbl, già del Museo di Storia Naturale di Ginevra, il Dr Lothar Zerche del DEI di
Eberswalde (Berlino) e il Dr. Volker Assing di Hannover. Per il prestito di tipi ringrazio
il Dr. A.F. Newton del Field Museum of Natural History di Chicago e il Dr. P.M.
Hammond e il Dr. Brendell del Museo di Storia Naturale di Londra.

RÉSUMÉ

Les espèces du genre Orphnebius Motschulscky, 1858, de Bornéo
(Coleoptera, Staphylinidae). - La présente étude sur le genre Orphnebius de Bornéo
contient l'illustration et la révision des holotypes ou lectotypes des six espèces sui-
vantes de Bornéo: O. bigladiosus (Bernhauer), O. bakerianus Bernhauer, O. splendens
Bernhauer, O. anguliceps Cameron, O. antennarius Bernhauer et O. bakeri Bernhauer.
Le lectotype de O. bakeri Bernhauer et celui de O. bakerianus Bernhauer sont
désignés. Les sous-genres Deroleptus Bernhauer et Mesocephalobius Bernhauer ont
été mis en synonymie avec Orphnebius et Orphnebius quadrigladiosus Pace, 1986,
avec Orphnebius quadricuspidatus Bernhauer, 1929. Deroleptus bigladiosus
(Bernhauer) et Deroleptus arachnoides Bernhauer sont transférés dans le genre

ORPHNEBIUS NEL BORNEO 769

Orphnebius. Les 13 espèces suivantes sont décrites comme nouvelles: O. borneanus,
O. scalaris, O. directus, O. minor, O. biapicalis, O. ideogramma, O. penangensis,
O. bajauorum, O. curticornis, O. uniformis, O. perpenetrans, O. krypticola et O. cras-
sus. L’habitus et les organes génitaux masculins et féminins des nouvelles espèces sont
illustrés. Une clé des espèces est fournie.

BIBLIOGRAFIA

BERNHAUER, M. 1911. Zur Staphylinidenfauna Ostindiens und der Sundainseln (3. Beitrag.
Schluss). Entomologische Blätter für Biologie und Systematik der Käfer 7: 86-93.

BERNHAUER, M. 1915. Neue Staphyliniden der indomalayschen Fauna, insbesondere der Sunda-
Insel Borneo. Verhandlungen der zoologisch-botanischen Gesellschaft in Wien, 65:
134-158.

BERNHAUER, M. 1929. Zur Kenntnis der Gattungen Astilbus Steph., Orphnebius Motsch. und
Deroleptus Bernh. Zoologischer Anzeiger (1929): 142-155.

BLACKWELDER, R.E. 1952. The generic names of the beetle family Staphylinidae with an essay
on genotypy. Bulletin of the United States National Museum 200: 483 pp.

CAMERON, M. 1943. New species of Staphylinidae (Col.) from Borneo. The Entomologist’s
Monthly Magazine 79: 39-42.

MOTSCHULSKY, V. DE. 1858. Enumération des nouvelles espèces de Coléoptères rapportées de
ses voyages. Bulletin de la Société impériale des Naturalistes de Moscou 31: 204-264.

PACE, R. 1986. Aleocharinae dell’ Asia sudorientale raccolte da G. de Rougemont (Coleoptera,
Staphylinidae) (LXXII Contributo alla conoscenza delle Aleocharinae). Bollettino del
Museo civico di Storia naturale di Verona 23: 139-237, 291 figg.

SCHEERPELTZ, O. 1934. Staphylinidae VII. In: SCHENKLING, S. (ed.). (pp. 1501-1881).
Coleoperorum Catalogus, 6(130): Junk, Berlin.

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REVUE SUISSE DE ZOOLOGIE 114 (4): 771-815; décembre 2007

Le specie di Oligotini, Leucocraspedini, Hygronomini, Placusini,
Bolitocharini e Diestotini nel Borneo
(Coleoptera, Staphylinidae)*

Roberto PACE
Via Vittorio Veneto 13, 1-37032 Monteforte d’Alpone (Verona), Italia.
E-mail: pace.ent@tiscali.it

The species of Oligotini, Leucocraspedini, Hygronomini, Placusini,
Bolitocharini and Diestotini from Borneo (Coleoptera, Staphylinidae). -
Forty-five new species of the subfamily Aleocharinae collected in the Mt
Kinabalu National Park, Borneo, are described and illustrated. For the first
time from Borneo are reported genera and species of the following tribes:
Oligotini, Hygronomini, Bolitocharini and Diestotini. Three new species
belong to the genus Oligota (densa, kinabaluensis and borneensis), one to
the genus Cypha (sabahensis), nineteen to the genus Leucocraspedum
(sinuatum, pilosellum, obliquum, spirasferum, occultum, biguttae, mima-
naticula, divisum, anaticula, fugitivum, directum, hamifer, dilatatiapex,
lamelliferum, audax, cacuminum, anguineatheca, nechamifer and ventrio-
satheca), four to the genus Hygrochara (micropallida, microkinabaluicola,
spiniventris and kinabaluensis), six to the genus Placusa (robustipes,
superba, recensita, falcifera, evoluta and subspinigera), four to the genus
Erastriusa (masculina, borneensis, minima and lobifera), three to the genus
Pseudatheta (borneensis, kinabaluensis and seditiosa), one to the genus
Neoleptusa (kinabaluensis), two to the genus Chledophila (parallela and
borneensis) and two to the genus Diestota (plicae and pellita). A key of all
species known to Borneo of the aforesaid genera is provided. Lectotype is
designated for Placusa acuminata Kraatz, whose aedeagus and spermatheca
for the first time are illustrated.

Keywords: Coleoptera - Staphylinidae - Aleocharinae - taxonomy - Borneo.

INTRODUZIONE

Il presente lavoro ha lo scopo di esporre il risultato dell’esame degli
Staphylinidae della sottofamiglia Aleocharinae raccolti nel Parco Nazionale del Monte
Kinabalu e altrove nel Borneo, dal Dr. Ale$ Smetana di Ottawa, dal Dr. Burckhardt e
dal Dr. Ivan Löbl del Museo di Storia Naturale di Ginevra e dal Prof. Herbert Franz.
Le 46 specie appartengono alle tribù degli Oligotini, Leucocraspedini, Hygronomini,
Placusini, Bolitocharini e Diestotini. I generi e le specie di queste tribt nel Borneo sono

* 209° Contributo alla conoscenza delle Aleocharinae.

772 R. PACE

assai poco noti. Hammond (1984) nella sua checklist sugli Staphylinidae del Borneo
non elenca i seguenti generi, nel presente lavoro rappresentati da una o più specie:
Oligota Mannerheim, 1831, Cypha Leach, 1819, Hygrochara Cameron, 1939,
Erastriusa Pace, 1990, Pseudatheta Cameron, 1920, Neoleptusa Cameron, 1939,
Chledophila Cameron, 1920 e Diestota Mulsant & Rey, 1870.

MATERIALE E METODO

L’esame & basato sugli esemplari adulti raccolti prevalentemente nel Parco
Nazionale del Monte Kinabalu dal Dr. Aleò Smetana di Ottawa durante le sue spedi-
zioni nel 1987 e 1988, dalla spedizione Burckhardt & Löbl del Museo di Storia
Naturale di Ginevra del 1987 e Bright pure del 1987 e dal materiale raccolto dal Prof.
H. Franz (senza anno).

La tassonomia delle nuove specie del Borneo presenta serie difficolta, in molti
casi superate grazie all’esame dei caratteri dell’organo copulatore maschile, dei
segmenti genitali maschili e femminili e della spermateca. Prima della pubblicazione
del presente lavoro nessun esame a fini tassonomici di questi importanti organi e
strutture è stato compiuto dagli autori del lontano passato. Gli holotypi delle specie
note sono stati da me esaminati e disegnati, quando disponibili, e inseriti nelle chiavi
qui date per la prima volta. Le recenti restrizioni riguardo ai prestiti di materiale tipico
imposte dal Museo di Storia Naturale di Londra, tuttavia mi hanno impedito di esa-
minare alcuni esemplari tipici. L'etimologia delle nuove specie è omessa quando
evidente come borneensis 0 kinabaluensis.

Quasi tutti gli esemplari sono stati dissezionati per le serie di pochi individui.
Le strutture genitali sono state montate in balsamo del Canadà su piccoli rettangoli
trasparenti di materiale plastico, infilzati sullo spillo dell’esemplare. Le strutture geni-
tali sono state studiate usando un microscopio composto e disegnate mediante oculare
a reticolo. Gli habitus sono stati disegnati con l’uso di un oculare micrometrico di un
microscopio binoculare. Tutti i disegni sono dell’autore fino alla fase finale. Il sicuro
riconoscimento dei generi e delle specie è qui affidato soprattutto alla parte illustrativa
che ha linguaggio internazionale. Per questo motivo le descrizioni sono brevi, limitate
a porre in evidenza ciò che non è riproducibile graficamente come il colore, la retico-
lazione e la granulosità. D'altronde per le specie della sottofamiglia Aleocharinae la
sola descrizione anche molto accurata e lunga non dà quasi mai la certezza di un’esatta
identificazione delle varie specie. È l'osservazione del disegno dell’edeago e/o della
spermateca insieme a quello dell’habitus che aiuta molto a risolvere problemi inter-
pretativi dati dalla sola descrizione.

Gli holotypi delle nuove specie sono depositati nel Museo di Storia Naturale di
Ginevra (MHNG) e in collezione Franz al Naturhistorisches Museum di Vienna
(Austria) (NHMW). Paratypi sono conservati in collezione Smetana e nell’Institut
Royal des Sciences Naturelles de Belgique di Bruxelles.

ALEOCHARINAE NEL BORNEO 173

OLIGOTINI
Specie di questa tribù erano finora sconosciute per il Borneo (Hammond, 1984).

CHIAVE DELLE SPECIE DEL GENERE OLIGOTA MANNERHEIM, 1831, NEL BORNEO

I Tutti gli uroterghi liberi senza carene; corpo uniformemente bruno-

rossiccio, Fig. 1; edeago Figg. 2-3. Lunghezza 0,95 mm ...... O. densa sp. n.
- Alcuni urotershitliberi/conlearenezteorpo/bieolore .2...2......... 2.0... 2
2 Corpo tozzo, Fig. 4; sutura delle elitre più lunga della linea mediana del

pronoto; base delle antenne rossiccia; carene del secondo urotergo libero
lunghe; edeago Figg. 5-6; porzione prossimale della spermateca lunga
quanto la lunghezza del bulbo distale della stessa spermateca, Fig. 7.
lEunshezza4l 22mm er. RES Fr er O. kinabaluensis sp. n.
- Corpo snello Fig. 8; sutura delle elitre più corta della linea mediana del
pronoto; base delle antenne gialla; carene del secondo urotergo libero
corte; porzione prossimale della spermateca molto più corta della
lunghezza del bulbo distale della stessa spermateca, Fig. 9. Lunghezza
len AR ee ee a iN O. borneensis sp. n.

KEY TO SPECIES OF THE GENUS OLIGOTA MANNERHEIM, 1831, FROM BORNEO

1 All the free uroterga without carina; body uniformly brown-reddish,

Eis taedeagusiEigs 2-3. Bensth 0. II’ mma 2 ... O. densa sp. n.
- Some free uroterga with carinae; body bicolorous ..................... 2
2 Body stumpy, Fig. 4; suture of the elytra longer than the median line of

the pronotum; base of the antenna reddish; carinae of the second free

urotergum long; aedeagus Figs 5-6; proximal portion of the spermathe-

ca as long as the length of the distal bulb of the same spermatheca,

Eie@7akensthalt2zmmys Eee ee O. kinabaluensis sp. n.
- Body slender Fig. 8; suture of the elytra shorter than the median line of

the pronotum; base of the antennae yellow; carinae of the second free

urotergum court; proximal portion of the spermatheca shorter than the

length of the distal bulb of the same spermatheca, Fig. 9. Length

ISIS EHE TI RON RO ERDE A I TA CAES O. borneensis Sp. n.

Oligota densa sp. n. Figg. 1-3

Holotypus maschio, Sabah, Kibongol V., 7 Km N Tambunan, 700 m, 20.V.1987, leg.
Burckhardt & Löbl, (MHNG).

DESCRIZIONE: Lunghezza 0,95 mm. Corpo lucido e bruno-rossiccio; antenne
bruno-rossicce con i tre antennomeri basali gialli; zampe gialle. La reticolazione &
assente su tutto il corpo. La punteggiatura del capo e del pronoto è superficiale. La
granulosità delle elitre è poco saliente, quella dell’addome è netta. Il primo urotergo
libero presenta una debole scultura squamiforme. Edeago Figg. 2-3.

DERIVATIO NOMINIS: La nuova specie prende nome dal carattere dell’edeago
denso di pezzi copulatori dell’ armatura interna.

Oligota kinabaluensis sp. n. Figg. 4-7

Holotypus maschio, Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt
& Löbl (MHNG).

774 R. PACE

Paratypi: 3 maschi e 1 femmina Sabah, Kibongol V., 7 Km N Tambunan, 700 m,
20.V.1987, leg. Burckhardt & Löbl; 1 maschio, Sabah, Poring Hot Springs, 500 m, 13.V.1987,
leg. Burckhardt & Löbl; 1 femmina, Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg.
Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,2 mm. Corpo lucido, molto convesso e bruno con
addome bruno-rossiccio e pigidio giallo sporco; antenne e zampe rossicce. La retico-
lazione è assente su tutto il corpo. Capo e pronoto privi di granulositä e punteggiatura.
La punteggiatura delle elitre è ben visibile. Gli uroterghi liberi, tranne il quinto, pre-
sentano carene basali, Fig. 4. Edeago Figg. 5-6, spermateca Fig. 7.

NOTE: La nuova specie è simile a O. rougemontiana Pace, 1993, di Celebes, ma
questa specie presenta gli antennomeri quarto, quinto e sesto più lunghi che larghi
(trasversi nella nuova specie) e la spermateca ha la porzione mediana chiaramente più
lunga di quella della nuova specie. Non è noto il maschio di O. rougemontiana.

Oligota borneensis sp. n. Figg. 3-9

Holotypus femmina, Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt
& Lôbl (MHNG).

DESCRIZIONE: Lunghezza 1,1 mm. Corpo lucido e bruno, con pigidio bruno-
rossiccio; antenne rossicce con i tre antennomeri basali gialli e undicesimo bruno-
rossiccio. La reticolazione è assente su tutto il corpo. Capo e pronoto privi di granulo-
sità e punteggiatura. La granulosità delle elitre è saliente. Il secondo urotergo libero
presenta corte carene basali mentre O. kinabaluensis sp. n. le presenta lunghe.
Spermateca Fig 8.

Cypha sabahensis sp. n. Figg. 10-11

Holotypus femmina, Sabah, Poring Hot Springs, Langanan Falls, 900-950 m, 12.V.1987,
leg. Burckhardt & Löbl (MHNG).

Paratypus: 1 femmina, Borneo, Sabah, Crocker Ra., 1270 m, Km 60 rte. Kota Kinabalu-
Tambunan, 17.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,4 mm. Corpo lucido, molto convesso e bruno;
antenne e zampe giallo-rossicce. La parte anteriore del corpo è privo di punteggiatura,
di granulosità e di reticolazione. L’addome presenta una distinta reticolazione che sul
quinto urotergo libero & trasversa. Spermateca Fig. 11.

NOTE: Per la forma della spermateca la nuova specie si rivela tassonomicamente
affine a C. besuchetiella (Pace, 1985), del Bengala, ma questa specie presenta occhi
assai ridotti. La nuova specie per la forma della spermateca è tassonomicamente affine
anche a C. rougemonti (Pace, 1986) della Malaysia, ma questa specie presenta la clava
antennale composta di tre antennomeri, mentre nella nuova specie è composta di due.
C. rougemonti ha l’introflessione apicale del bulbo distale della spermateca meno
lunga di quella della nuova specie ed è divisa in quattro settori.

LEUCOCRASPEDINI

La sola specie Leucocraspedum robustum Cameron, 1928 era finora nota del

Borneo, non del Mt Kinabalu, ma del Mt Dulit. Questa specie è compresa nella
seguente chiave.

ALEOCHARINAE NEL BORNEO 775

EN
=
THERE is 2

nr da pi
(osi

Prada
LT ee

Fico. 1-9

Habitus, edeago in visione laterale e ventrale e spermateca. (1-3) Oligota densa sp. n. (4-7)
Oligota kinabaluensis sp. n. (8-9) Oligota borneensis sp. n.

CHIAVE DELLE SPECIE DEL GENERE LEUCOCRASPEDUM KRAATZ, 1859 NEL BORNEO
1 Corpo rossiccio con pigidio e margine posteriore degli uroterghi giallo-
COSsicciileunehezz 25 amma ER ERRO NS L. robustum Cameron

- Corpo diversamente colorato; margine posteriore degli uroterghi liberi
MATOS SUC CLO, on ONE II ERI RT RE Pi 2

776

ON

I '

13

R. PACE

Sutura delle elitre più corta della linea mediana del pronoto, Figg. 12 e 15 . . 3
Sutura delle elitre lunga quanto il pronoto, poco più corta o molto più

MMS MA a i PRE RO LIT RAI TE 5)
Corpo bicolore giallo-rossiccio e bruno; pronoto pit largo delle elitre,

Fig. 12; edeago largo e bisinuato, in visione ventrale, Fig. 14. Lunghezza

I No Stettino L. sinuatum sp. n.
Corpo unicolore giallo-rossiccio; pronoto poco piu largo delle elitre o
appena piu stretto, Figg. 15 e 18; edeago stretto e non sinuato in visione
Ventrale et. es ee ee nee AN © ROOT 4
Pronoto largo quanto le elitre e arcuato al margine anteriore, Fig. 15;

elitre piu ridotte, Fig. 15; armatura genitale interna dell’edeago con un
cespuglio di setole, Fig. 16. Lunghezza 1,7 mm......... L. pilosellum sp. n.
Pronoto poco più largo delle elitre, obliquamente ristretto al margine
anteriore, Fig. 18; elitre meno ridotte, Fig. 18; armatura genitale interna
dell’edeago senza un cespuglio di setole. Lunghezza 1,7 mm

MA SOR tO ST e ir eta RICA L. obliquum sp. n.

Sutura delle elitre poco più corta della linea mediana del pronoto ......... 6
Sutura delle elitre lunga quanto la linea mediana del pronoto, lievemente

più corta 0,molto più lung... at Ra ERRE 18
Pronoto;giallo-rossiccio:0 bruno... 0. ROLE TE 7
Pronoto rossiceio'0,bruno,. 2,150 PE MEET eee 16
Corpo unicolore giallo-rossiccio, talvolta il pigidio è giallo.............. 8
Corpo bicolore giallo-rossiccio e bruno o bruno-rossiccio ............... 11

Undicesimo antennomero lungo quanto i quattro antennomeri immedia-
tamente precedenti riuniti; spermateca Fig. 22. Lunghezza 2,2 mm

ARA RSR RE > à HS Stig deco à EEE or L. spirasferum sp. n.
Undicesimo antennomero lungo quanto 1 tre antennomeri immediata-

mente precedenti riuniti,0PoCO.menor... è... 2 Men. SEE 9
Corpo giallo-bruno; bulbo distale della spermateca a forma di due lobi

similia:socce Fig. 24. Tunshezzasl#8imm..e.. 22... RR L. biguttae sp. n.
Corporgialio=rossiccio..... RM RE ia DPI EEE 10
Pigidio giallo-rossiccio chiaro; pronoto regolarmente arcuato al margine

antenore,, Big) 25; edeago Bigg, 260770 23. nn ee L. occultum sp. n.
Pigidio giallo-rossiccio; pronoto obliquamente ristretto al margine ante-

More Fig 26 Spermateca Fe Den Sr ee L. mimanaticula sp. n.
Elitre giallo-rossicce o giallo-brune........ - .......... EE TE 12
Elitre,brune o TOSSICCE iii aL 13

Pronoto ed elitre giallo-rossicci, addome giallo-rossiccio con base degli
uroterghi liberi rossiccia o bruna; edeago Figg. 32-33. Lunghezza

ES: mm... > ees ages à ald ates RATTEN En L. divisum sp. n.
Pronoto ed elitre giallo-bruni; addome bruno con pigidio giallo-rossic-
Ci0:Spermateca Fo 59" Eunshezzatl: Immer er were L. anaticula sp.n.

Undicesimo antennomero lungo quanto i due antennomeri immediata-
mente precedenti riuniti che sono lunghi quanto larghi; edeago brusca-
mente ristretto presso la «crista apicalis», in visione ventrale, Figg. 37-
S82lEunshez73,2:6.mm.. ns er ee Dee L. fugitivum sp.n.

14

15

16

17

18

19

ALEOCHARINAE NEL BORNEO Tatil

Undicesimo antennomero lungo quanto I tre antennomeri immedia-
tamente precedenti riuniti che sono trasversi; edeago non come sopra..... 14
Addome bruno-rossiccio con base giallo-rossiccia; pronoto con quattro

punti disposti in rettangolo; edeago profondamente e ampiamente ar-

cuato al livello della «crista apicalis», Figg. 41-42. Lunghezza 1,9 mm
NEE Ro Re ROE RO LO ale L. directum sp. n.
Addome bruno-rossiccio, senza base giallo-rossiccia .................. 15
Pigidio giallo-rossiccio; pronoto obliquamente ristretto al margine
anteriore, Fig 45; apice dell’edeago a forma di piccolo gancio in visione
laterale, Figg. 43-44. Lunghezza 2,1 mm................. L. hamifer sp. n.
Pigidio bruno-rossiccio; pronoto arcuato al margine anteriore, Fig. 46;

apice dell’edeago terminante a paletta circolare, in visione ventrale,
Ei9s%47-48+Eunshezza lSmme LEE aep. ser L. dilatatiapex sp. n.
Margini laterali del pronotum rossicci; addome unicolore bruno; edeago

Figg. 51-52; spermateca Fig. 53. Lunghezza 1,9 mm ....L. lamelliferum sp. n.
Pronoto unicolore; addome bicolore bruno e rossiccio ................. 177,
Undicesimo antennomero lungo quanto i sette antennomeri immedia-
tamente precedenti riuniti, Fig. 54; edeago Figg. 55-56. Lunghezza
ZIMMER III DE RP LI L. audax sp. n.
Undicesimo antennomero lungo quanto i due antennomeri immediata-

mente precedenti riuniti, Fig. 58; spermateca Fig. 57. Lunghezza 2,7 mm

SVI PTE TR SESSI TAR ROERO OO RES BR CRE TO weiss L. cacuminum Sp. n.
Sutura delle elitre molto più lunga della linea mediana del pronoto; pro-

noto con quattro punti discali in quadrato; addome giallo-rossiccio con

una fascia bruna; spermateca Fig. 60. Lunghezza 2 mm

N e REED EEE L. anguineatheca sp. n.
Sutura delle elitre lunga quanto la linea mediana del pronoto o lieve-

mente più corta; pronoto mai con quattro punti discali in quadrato;
addome,senzattaSsC la DIUNA e O Re 19
Undicesimo antennomero lungo quanto i sette antennomeri precedenti
riuniti, Fig. 61; elitre giallo-rossicce; addome tricolore rossiccio con pi-

gidio giallo-rossiccio e base degli uroterghi liberi bruna; bulbo basale
dell’edeago stretto, in visione ventrale, Figg. 62-63. Lunghezza 2,7 mm
SEMI: A EI e EHESTEN L. nechamifer sp. n.
Undicesimo antennomero lungo quanto i tre antennomeri immediata-

mente precedenti riuniti, Fig. 67; elitre bruno-rossicce, addome uni-
colore bruno-rossiccio; bulbo basale dell’edeago assai dilatato, in
visione ventrale, Figg. 64-65; spermateca Fig. 66. Lunghezza 1,8 mm

A ae AA ah AE cnet ln ithe antares ri L. ventriosatheca sp. n.

KEY TO SPECIES OF THE GENUS LEUCOCRASPEDUM KRAATZ, 1859, FROM BORNEO

1

Body reddish with pigidium and basal border of the uroterga yellow-
reddish. Een ot 2 Skiers RE eee RO EE L. robustum Cameron
Body otherwise colored; basal border of the free uroterga never reddish... . 2
Suture of the elytra shorter than the median line of the pronotum, Figs
I yar IDEEN IE ONS POSTE QC PER AU PRET TE ee LPS LS ER QUÇSESS 3

778

©

I '

R. PACE

Suture of the elytra as long as the pronotum, a little shorter or very longer . . 5
Bicolorous body, yellow-reddish and brown; pronotum wider than the
elytra, Fig. 12; aedeagus wide and bisinuous, in ventral view, Fig. 14.
[Pen sthale7emmi: SERPENT LO RIA AE L. sinuatum sp. n.
Body yellow-reddish unicolorous; pronotum a little wider than the elytra
or scarcely more narrow, Figs 15 and 18; aedeagus narrow and not
sinuous'in-ventral view I. PISANO A O 4
Pronotum as wide as the elytra and arched to the anterior border, Fig. 15;
elytra more redoubts, Fig 15; inside genital armour of the aedeagus with
abush of bristles; Fig 16: Lensthl. «mm RR rer L. pilosellum sp. n.
Pronotum a little wider than the elytra, obliqually narrow to the anterior
border, Fig. 18; elytra less reduced, Fig. 18; inside genital armour of the

aedeagus without a bush of bristles. Length 1.7 mm....... L. obliquum sp. n.
Suture of the elytra a little shorter than the median line of the pronotum .... 6
Suture of the elytra as long as the median line of the pronotum, slightly

Shorterior very longer. neo seine PR RE 18
Pronotum yellow-reddish or brown =: . 2... e OE 7
Pronotum’reddishvormbrown AT EE EPL I SOC IR 16
Body yellow-reddish unicolorous, sometimes the pigidium is yellow....... 8
Body yellow-reddish and brown or brown-reddish bicolorous ............ 11

Eleventh antennomere as long as the four immediately precedents
reunited antennomere; spermatheca Fig. 22. Length 2.2 mm

CRI ac Rea aS A I III IGO II ma ER à de L. spirasferum Sp. n.
Eleventh antennomere as long as the three immediately precedents

reunited antennomere Or fewiless........ OE 9
Body yellow-brown; distal bulb of the spermatheca to form of two lobes
Similar todrops hie 24; en et les mim) E CCE L. biguttae sp. n.
Body, yellow-reddish mr 24.2. eh. Lu. han cae NG 10
Pigidium clear yellow-reddish; pronotum arched regularly to the anteri-
or border, Fig. 25: aedeagus Figs 26-27. Length 1.7 mm....L. occultum sp. n.

Pigidium yellow-reddish; pronotum obliqually narrow to the anterior
border, Fig. 28; spermatheca Fig 29. Length 1.7 mm... L. mimanaticula sp. n.
Elytra yellow-reddish or yellow-brown . .. ....... „>... . ..... io ee 12
Elytra,brown or reddish.. si. tI 13
Pronotum and elytra yellow-reddish, abdomen yellow-reddish with base
of the free uroterga reddish or brown; aedeagus Figs 32-33. Length

RON Peet ROS MR Ne MERC A EEE L. divisum sp. n.
Pronotum and elytra yellow-brown; abdomen brown with yellow-
reddish pigidium; spermatheca Fig. 35. Length 1.9 mm....L. anaticula sp. n.

Eleventh antennomere as long as the two immediately precedents
reunited antennomere that are as long as wide; aedeagus brusquely
narrow near the «crista apicalis», in ventral view, Figs 37-38. Length
PLO renti Oe wheels a RR es Nee dee eee L. fugitivum sp. n.
Eleventh antennomere as long as the three immediately precedents
reunited antennomere that are transverse; aedeagus not as above......... 14

16

17

ALEOCHARINAE NEL BORNEO 779

Abdomen brown-reddish with yellow-reddish base; pronotum with four

points set in rectangle; aedeagus deeply and broadly arched to the level

of the «crista apicalis», Figs 41-42. Length 1.9 mm ........ L. directum sp. n.
Abdomen brown-reddish, without yellow-reddish base ................. 15
Pigidium yellow-reddish; pronotum obliquely narrow to the anterior
border, Fig 45; apex of the aedeagus to form of small hook in lateral

Wiew iss 45-44 oBensth2.ll mimi are L. hamifer sp. n.
Pigidium brown-reddish; pronotum arched to the anterior border, Fig.

46; apex of the aedeagus finished like circular shovel, in ventral view,

iss 48s Wensth 1. 8 mimeo ann ee L. dilatatiapex sp. n.
Side borders of the pronotum reddish; abdomen brown unicoloured;
aedeagus Figs 51-52; spermatheca Fig. 53. Length 1.9 mm

So Bo Me ae PR CONS Eun Lite EC PRE ACRI L. lamelliferum sp. n.
Pronotum unicoloured; bicoloured abdomen brown and reddish .......... 11
Eleventh antennomere as long as the seven immediately precedents
reunited antennomere, Fig. 54; aedeagus Figs 55-56. Length 2.7 mm

à PRE RE OE Rr TEE RA I e SEND SR A ER L. audax sp. n.
Eleventh antennomere as long as the two immediately precedents
reunited antennomere, Fig. 58; spermatheca Fig. 57. Length 2.7 mm

20 (Blaue ZEAL à SIONS Wath TER. 5er 01 MONO APR ONE RE L. cacuminum sp. n.
Suture of the elytra longer than the median line of the pronotum; pro-

notum with four discal points in square; abdomen yellow-reddish with a

brown band; spermatheca Fig. 60. Length 2 mm ...... L. anguineatheca sp. n.
Suture of the elytra as long as the median line of the pronotum or slightly
shorter; pronotum never with four discal points in square; abdomen
Without brown bande 8. dare pa eh OE 19
Eleventh antennomere as long as the seven preceding reunited anten-
nomere, Fig. 61; elytra yellow-reddish; tricoloured abdomen reddish

with yellow-reddish pigidium and brown base of the free uroterga; basal

bulb of the aedeagus narrow, in ventral view, Figs 62-63. Length 2.7 mm

adh Scie RE MORE UE RSR a Vada RE RER APE RICO ES Tau. L. nechamifer sp. n.
Eleventh antennomere as long as the three immediately precedents
reunited antennomere, Fig. 67; elytra brown-reddish, unicoloured
abdomen brown-reddish; basal bulb of the aedeagus a great deal dilated,

in ventral view, Figs 64-65; spermatheca Fig. 66. Length 1.3 mm

EEE BEE RR IA ue CA L. ventriosatheca Sp. n.

Leucocraspedum sinuatum sp. n. Figg. 12-14

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,

12.V.1987, leg. A. Smetana, (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e bruno con pronoto e meta

posteriore del quinto urotergo libero del maschio giallo-rossicci; antenne e zampe
gialle. La reticolazione è assente su tutto il corpo. Il pronoto è coperto di granulosità
molto superficiale, le elitre e l’addome di granulosità distinta. Le elitre sono cortissime,
più corte e più strette del pronoto. Edeago Figg. 13-14.

780 R. PACE

Fico. 10-17

Habitus, spermateca e edeago in visione laterale e ventrale. (10-11) Cypha sabahensis sp. n.
(12-14) Leucocraspedum sinuatum sp. n. (15-17) Leucocraspedum pilosellum sp. n.

DERIVATIO NOMINIS: La nuova specie prende nome dai lati sinuosi del suo
edeago.

Leucocraspedum pilosellum sp. n. Figg. 15-17

Holotypus maschio, Borneo-Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,
13.V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e giallo-rossiccio, comprese
antenne e zampe. La reticolazione del pronoto e dell’addome è assente, quella delle

ALEOCHARINAE NEL BORNEO 781

elitre & superficiale. La granulosità del pronoto e dell’addome è fine, quella delle elitre
è distinta. Edeago Figg 15-16.

DERIVATIO NOMINIS: La nuova specie prende nome dal ciuffo di peli dell’arma-
tura genitale interna dell’edeago.

Leucocraspedum obliquum sp. n. Figg. 18-20

Holotypus maschio, Sabah, Poring Hot Springs, 500 m, 13.V.1987, leg. Burckhardt &
Lôbl (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e giallo-rossiccio, comprese
antenne e zampe. La reticolazione manca su tutto il corpo. La granulosita del pronoto
è molto superficiale, quella delle elitre è saliente e quella dell’addome è ben distin-
guibile. Edeago Figg. 19-20.

DERIVATIO NOMINIS: La nuova specie prende nome dai lati obliqui in avanti del
pronoto.

Leucocraspedum spirasferum sp. n. Figg. 21-22

Holotypus femmina, Borneo, Sabah, Poring Hot Springs, 500 m, 6.V.1987, leg.
Burckhardt & Löbl (MHNG).

Paratypus: 1 femmina, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 550 m,
7.V.1987, leg. Burckhardt & Löbl

DESCRIZIONE: Lunghezza 2,2 mm. Corpo lucido e giallo-rossiccio; antenne
rossicce con i tre antennomeri basali rossicci; zampe gialle. La reticolazione del capo
e dell’addome è assente, quella delle elitre è superficiale. La granulosità del pronoto è

superficiale, quella delle elitre è saliente e quella dell'addome è ben visibile.
Spermateca Fig. 22.

DERIVATIO NOMINIS: La nuova specie prende nome dalla porzione distale della
spermateca che descrive diverse spire, perciò è chiamata «portatrice di spire».

Leucocraspedum biguttae sp. n. Figg. 23-24

Holotypus femmina, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, area blw
Langanan Fall, 850-900 m, 14.V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 1,8 mm. Corpo lucido e giallo sporco; antenne brune
con i tre antennomeri basali gialli; zampe giallo-rossicce. La reticolazione è assente su
tutto il corpo. La granulosità del pronoto è molto superficiale, quella delle elitre e
dell’addome è superficiale. Spermateca Fig. 24.

DERIVATIO NOMINIS: La nuova specie prende nome dalla porzione distale della
spermateca che ha forma di due gocce.

Leucocraspedum occultum sp. n. Figg. 25-27

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,
12.V.1987, leg. A. Smetana, (MHNG).

Paratypus: 1 maschio, Sabah, Poring Hot Springs, 550-600m, 9.V.1987, leg. Burckhardt
& Löbl.

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e giallo sporco con pigidio
giallo-rossiccio chiaro; antenne giallo-rossicce con i cinque antennomeri basali gialli;

782 R. PACE

Fico. 18-24

Habitus, edeago in visione laterale e ventrale e spermateca. (18-20) Leucocraspedum obliquum
sp. n. (21-22) Leucocraspedum spirasferum sp. n. (23-24) Leucocraspedum biguttae sp. n.

zampe gialle. La reticolazione del pronoto e dell’addome è assente, quella delle elitre
è superficiale. La granulosita del pronoto è molto superficiale, quella delle elitre e
dell’addome è ben visibile. Edeago Figg. 26-27.

DERIVATIO NOMINIS: La nuova specie prende nome dal fatto che era occultata o
nascosta tra gli esemplari di L. sinuatum sp. n. e L. pilosellum sp. n.

ALEOCHARINAE NEL BORNEO 783

Leucocraspedum mimanaticula sp. n. Figg. 28-29

Holotypus femmina, Sabah, Crocker Ra., 1200 m, Km 63 r.te Kota Kinabalu-Tambunan,
19.V.1987, leg. Burckhardt & Löbl, (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e giallo-rossiccio; antenne
rossicce con i quattro antennomeri basali gialli; zampe giallo-rossicce. La reticolazione
del corpo è assente. La granulosità del pronoto è superficiale, quella delle elitre e
dell’addome è saliente. Spermateca Fig. 29.

DERIVATIO NOMINIS: La nuova specie prende nome di «imitatrice di anaticula»
poiché la sua spermateca è simile a quella di L. anaticula sp. n.

Leucocraspedum divisum sp. n. Figg. 30-33

Holotypus maschio, Sabah, Poring Hot Springs, 500 m, 8.V.1987, leg. Burckhardt &
Löbl.

Paratypi: 1 maschio e 1 femmina, Sabah, Poring Hot Springs, Langanan Falls, 900-950
m, 12.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,8 mm. Corpo lucido e giallo-rossiccio con base
degli uroterghi liberi secondo e terzo rossiccia e base degli uroterghi liberi quarto e
quinto bruna; antenne rossicce con i tre antennomeri basali gialli; zampe gialle. La
reticolazione del corpo è assente. La granulosità del pronoto e dell’addome è super-
ficiale, quella delle elitre è saliente. Edeago Figg. 32-33, spermateca Fig. 31.

DERIVATIO NOMINIS: La nuova specie prende nome di «divisa» poiché il suo
edeago ha la lamina ventrale separata dal tubulo che contiene l'armatura genitale
interna.

Leucocraspedum anaticula sp. n. Figg. 34-35

Holotypus femmina, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 485 m,
29.VII.1987, leg. A. Smetana, (MHNG).

DESCRIZIONE: Lunghezza 1,9 mm. Corpo lucido. Capo e pronoto gialli, elitre
giallo-brune e addome bruno con pigidio giallo-rossiccio; antenne giallo-rossicce;
zampe gialle. La reticolazione del pronoto e delle elitre è molto superficiale, quella
dell’addome è assente. La granulosità del pronoto è indistinta, quella delle elitre è
superficiale e quella dell’addome è ben visibile. Spermateca Fig. 35.

DERIVATIO NOMINIS: La nuova specie prende nome di «anatroccolo» perché la
sua spermateca ricorda il profilo di una piccola anitra.

Leucocraspedum fugitivum sp. n. Figg. 36-39

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m,
4.VIIL.1988, leg. A. Smetana (MHNG).

Paratypi: 10 esemplari, Borneo, Sabah, Mt Kinabalu Nat. Pk., HQ at Livagu Rv., 1500m,
17.V.1987, leg. A. Smetana; 8 es., Borneo, Sabah, Mt Kinabalu Nat. Pk., Silau-Silau Trail,
1558m, 2.1X.1988, leg. D.E. Bright; 1 es, Borneo-Sabah, Mt Kinabalu Nat. Pk. HQ, Silau-Silau
Trail, 1560m, 23.V.1987, leg. A. Smetana; 2 es., Borneo, Sabah, Mt Kinabalu Nat. Pk.,
Headquarters, 1550m, 3-12.VIII.1988, beating foliage, leg. D.E. Bright; 2 es., Borneo-Sabah, Mt
Kinabalu Nat. Pk., Headquarters, 1560m, 24.IV.1987, beating foliage, leg. D.E. Bright; 1 ma-
schio, Borneo, Sabah, Mt Kinabalu Nat. Pk., Poring Hot Springs, 486m, 14.VIIL.1988, leg. D.E.
Bright.

784 R. PACE

FIGG. 25-31
Habitus, edeago in visione laterale e ventrale e spermateca. (25-27) Leucocraspedum occultum
sp. n. (28-29) Leucocraspedum mimanaticula sp. n. (30-31) Leucocraspedum divisum sp. n.

DESCRIZIONE: Lunghezza 2,6 mm. Corpo lucido. Capo, pronoto e pigidio giallo-
rossicci, elitre e addome rossicci; antenne giallo-rossicce con i tre antennomeri basali
gialli; zampe giallo-rossicce. La reticolazione del corpo & assente. La granulositä del
pronoto manca, quella delle elitre è ben visibile e quella dell’addome è fine. Edeago
Figg. 37-38, spermateca Fig. 39.

ALEOCHARINAE NEL BORNEO

785

35

0,1 mm

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39

37 38
Fico. 32-39
Edeago in visione laterale e ventrale, habitus e spermateca. (32-33) Leucocraspedum divisum sp.
n. (34-35) Leucocraspedum anaticula sp. n. (36-39) Leucocraspedum fugitivum sp. n.
suggerito il nome di «fuggitiva».
Leucocraspedum directum sp. n.

Löbl.

DERIVATIO NOMINIS: La nuova specie poiché é presente in differenti località ha
12.V.1987, leg. A. Smetana, (MHNG).

Figg. 40-42
Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,

Paratypus: 1 maschio, Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg. Burckhardt &

786 R. PACE

DESCRIZIONE: Lunghezza 1,9 mm. Corpo lucido e giallo-rossiccio con elitre e
uroterghi liberi terzo, quarto, quinto e sesto bruno-rossicci; antenne rossicce con i tre
antennomeri basali gialli; zampe gialle. Sul pronoto e sull’addome la reticolazione
manca, sulle elitre è ben visibile. La granulosita del pronoto è fine e superficiale, quella
delle elitre e dell’addome è saliente. Il pronoto presenta sul disco quattro punti disposti
in quadrato. Edeago Figg. 41-42.

DERIVATIO NOMINIS: La nuova specie prende nome di «rettilinea» poiché la parte
distale dell’edeago è appunto rettilinea, in visione laterale.

Leucocraspedum hamifer sp. n. Figg. 43-45

Holotypus maschio, Borneo-Sabah, M. Kinabalu N.P., Summit Trail, 1890 m, leg. A.
Smetana, (MHNG).

DESCRIZIONE: Lunghezza 1,9 mm. Corpo lucido. Capo, pronoto e pigidio giallo-
rossicci, resto del corpo bruno-rossiccio; antenne giallo-rossicce con i tre antennomeri
basali gialli e l’undicesimo bruno-rossicci. La reticolazione del pronoto è assente,
quella delle elitre è superficiale e quella dell'addome è molto svanita. La granulosita
del pronoto è fine e molto superficiale, quella delle elitre è ben visibile e quella
dell’addome è saliente. Edeago Figg. 43-44.

DERIVATIO NOMINIS: La nuova specie prende nome di «portatrice di amo» perché
l’apice dell’edeago ha forma di amo.

Leucocraspedum dilatatiapex sp. n. Figg. 46-49

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m,
12.V.1987, leg. A. Smetana, (MHNG).

Paratypus: Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,8 mm. Corpo lucido. Capo e pronoto giallo-rossicci,
elitre e addome bruno-rossicci; antenne rossicce con i tre antennomeri basali gialli e
l’undicesimo bruno-rossiccio; zampe gialle. Solo le elitre presentano una reticolazione
che però è molto superficiale, sul resto del corpo non è visibile alcun tipo di retico-
lazione. La granulosità del pronoto è fitta, fine e superficiale, quella delle elitre e
dell’addome è saliente. Apice dell’edeago a paletta circolare Figg. 47-48, spermateca
Fig. 49.

DERIVATIO NOMINIS: La nuova specie prende nome di «apice dilatato». L’apice
dilatato è quello dell’edeago.

Leucocraspedum lamelliferum sp. n. Figg. 50-53
Holotypus maschio, Borneo-Sabah, Mt. Kinabalu, Poring Hot Springs, 500 m,
13.V.1987, leg. A. Smetana (MHNG).
Paratypi: 1 femmina, Borneo-Sabah, Mt. Kinabalu, Poring Hot Springs, 490 m,
16.VIIL.1988, leg. A. Smetana; 1 femmina, Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg.
Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,9 mm. Corpo lucido. Capo e pronoto bruno-rossicci
con margini laterali rossicci, resto del corpo bruno; antenne gialle; zampe giallo-ros-
sicce. La reticolazione del corpo manca. La granulosità del pronoto è finissima e fitta,
quella delle elitre e dell’addome è saliente. Edeago Figg. 51-52, spermateca Fig. 53.

ALEOCHARINAE NEL BORNEO 787

FıGG. 40-45

Habitus e edeago in visione laterale e ventrale. (40-42) Leucocraspedum directum sp. n. (43-45)
Leucocraspedum hamifer sp. n.

DERIVATIO NOMINIS: Il nome della nuova specie significa «portatore di lamella».
La lamella è quella dell’armatura genitale interna dell’edeago.

788 R. PACE

RDA
INNEN I

; 3
TATRA
Hs € "4
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VERSA y
1
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46 49

53

0,1 mm
,l mm

52

FIGG. 46-53
Habitus, edeago in visione laterale e ventrale e spermateca. (46-49) Leucocraspedum dilatatia-

pex sp. n. (50-53) Leucocraspedum lamelliferum sp. n.

Leucocraspedum audax sp. n. Figg. 54-56

Holotypus maschio, Borneo Sabah, Mt. Kinabalu Nat Pk., HQ 1560 m, 24.IV.1987,
Beating foliage, D.E Bright collector (MHNG).

ALEOCHARINAE NEL BORNEO 789

DESCRIZIONE: Lunghezza 2,7 mm. Corpo lucido e bruno-rossiccio con pigidio
rossiccio; antenne gialle; zampe giallo-rossicce. La reticolazione è presente solo sulle
elitre dove è superficiale. La granulosità del pronoto è fine e distinta, quella delle elitre
è saliente. Gli uroterghi liberi sono coperti di scultura squamiforme. Edeago Figg.
55-56.

DERIVATIO NOMINIS: Il nome della nuova specie di «audace» è stato suggerito
dall’aggressiva armatura genitale interna dell’edeago a forma di gancio.

Leucocraspedum cacuminum sp. n. Figg. 57-58

Holotypus femmina, Borneo, Sabah, Mt. Kinabalu N.P., summit trail Pondok-Ubah,
2050 m, 26.1V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 2,7 mm. Corpo lucido e rossiccio con gli uroterghi
liberi secondo, terzo e quarto bruni; antenne brune con 1 tre antennomeri basali gialli;
zampe giallo-rossicce. La reticolazione del corpo è assente. La granulosità del pronoto
è fine e distinta, quella delle elitre è molto saliente e quella dell’addome è squami-
forme. Spermateca Fig. 57.

DERIVATIO NOMINIS: Il nome della nuova specie significa «delle vette». È stata
infatti raccolta ad alta quota.

Leucocraspedum anguineatheca sp. n. Figg. 59-60

Holotypus femmina, Borneo, Sabah, M. Kinabalu, 2600 m, 2.V.1987, leg. A. Smetana
(MHNG).

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido e giallo-rossiccio con elitre e base
degli uroterghi liberi primo, secondo e terzo rossicci, quarto e base del quinto bruni;
antenne rossicce con 1 quattro antennomeri basali giallo-rossicci; zampe giallo-
rossicce. La reticolazione del pronoto è assente, quella delle elitre è superficiale. La
granulosita del pronoto è molto superficiale, quella delle elitre è saliente. Spermateca
Fig. 60.

DERIVATIO NOMINIS: Il nome della nuova specie significa «spermateca a forma
di serpente». La parte distale di quest’organo, infatti, ha la figura di serpente.

Leucocraspedum nechamifer sp. n. Figg. 61-63

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., Liwagu River tr., 1495-1550 m,
12.V111.1988, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 2,7 mm. Corpo lucido. Capo e pronoto bruni, margine
posteriore del pronoto giallo-rossiccio, elitre giallo-rossicce, addome rossiccio con
base di ciascun urotergo libero bruna, pigidio giallo-rossiccio; antenne gialle con i tre
antennomeri distali giallo-rossicci; zampe brune con femori giallo-rossicci. La retico-
lazione è assente su tutto il corpo. La granulosità del pronoto è fitta e finissima, quella
delle elitre e dell’addome è saliente. Edeago Figg. 62-63.

DERIVATIO NOMINIS: Il nome della nuova specie significa «non portatrice di
amo». Prima di dissezionare l'esemplare, secondo mia ipotesi provvisoria l’avevo
determinato come L. hamifer sp. n., sopra descritto.

790 R. PACE

n
MAM)
Acer

a se ern

Fico. 54-60

Habitus, edeago in visione laterale e ventrale e spermateca. (54-56) Leucocraspedum audax
sp. n. (57-58) Leucocraspedum cacuminum sp. n. (59-60) Leucocraspedum anguineatheca sp. n.

Leucocraspedum ventriosatheca sp. n. Figg. 64-67

Holotypus maschio, Borneo-Sabah, Mt. Kinabalu, Poring Hot Springs, 500 m,
13.V.1987, leg. Burckhardt & Löbl (MHNG).

Paratypus: 1 femmina, Sabah, Poring Hot Springs, Langanan Falls, 900-950 m,
12.V.1987, leg. Burckhardt & Löbl.

ALEOCHARINAE NEL BORNEO 791

ì

Lan
A

FIGG. 61-67

Habitus, edeago in visione laterale e ventrale e spermateca. (61-63) Leucocraspedum nechamifer
sp. n. (64-67) Leucocraspedum ventriosatheca sp. n.

DESCRIZIONE: Lunghezza 1,8 mm. Corpo lucido. Capo e pronoto giallo-rossicci,
elitre e addome bruno-rossicci; antenne bruno-rossicce con i tre antennomeri basali
gialli; zampe gialle. La reticolazione è assente su tutto il corpo. La granulosità del
pronoto e dell’addome è ben visibile, quella delle elitre è saliente. Edeago Figg. 64-65,
spermateca Fig. 66.

DERIVATIO NOMINIS: Il nome della nuova specie significa «spermateca panciuta».

792

R. PACE

HYGRONOMINI

Nessuna specie di questa tribù era finora nota per il Borneo (Hammond, 1984).

Per i caratteri della spermateca e a motivo della taglia corporea e dell’habitus, le specie
del genere Hygrochara Cameron, 1939, si rivelano tassonomicamente affıni al genere
Apimela Mulsant & Rey, 1874. I due generi sono collocati in due tribü differenti a
motivo della formula tarsale: 4-4-4 per Hygrochara e 5-5-5 per Apimela (Oxypodini).
Pertanto l’attribuzione di Hygrochara alla tribù Hygronomini è qui del tutto provvisoria
in attesa di una revisione delle specie di altri generi affini appartenenti a questo gruppo.

CHIAVE DELLE SPECIE DEL GENERE HYGROCHARA CAMERON, 1939, NEL BORNEO

1

Occhi molto ridotti; sutura delle elitre più corta della linea mediana del

PIONOLOLEI NA ie ee ele. OP EEE 2
Occhi grandi; sutura delle elitre lunga quanto la linea mediana del pro-
notoo piulunga E e N RER EEE 3

Corpo snello, Fig. 68; undicesimo antennomero giallo; elitre e quarto
urotergo libero rossicci; edeago poco arcuato al lato ventrale, Figg.
69-70; spermateca minore, Fig. 71. Lunghezza 1,3 mm . A. micropallida sp. n.
Corpo tozzo, Fig. 72; undicesimo antennomero bruno; elitre e quarto
urotergo libero giallo-rossicci; edeago molto arcuato al lato ventrale,

Figg. 73-74; spermateca maggiore, Fig. 75. Lunghezza 1,28 mm
RÉ I a a, H. microkinabaluicola sp. n.
Quarto antennomero trasverso; sutura delle elitre lunga quanto la linea
mediana del pronoto; pigidio giallo-rossiccio; edeago con due spine ven-

trali, Figg. 77-78; spermateca Fig. 79. Lunghezza 2 mm. . H. spiniventris sp. n.
Quarto antennomero più lungo che largo; sutura delle elitre più lunga

della linea mediana del pronoto; pigidio bruno-rossiccio; edeago senza

spine ventrali, Figg. 73-74; spermateca Fig. 75. Lunghezza 2,1 mm

BE SISTER x che fe sre Late N CONTRE H. kinabaluensis sp. n.

KEY TO SPECIES OF THE GENUS HYGROCHARA CAMERON, 1939, FROM BORNEO

l

i)

Eyes distinctly reduced; suture of the elytra shorter than the median line

of-the,ProNOLUM.. .. . 1... Ye cae hee IE, 2
Eyes large; suture of the elytra as long as the median line of the pro-
DOMWMIORIONSET i... n OE 3)

Body slender, Fig. 68; eleventh antennomere yellow; elytra and fourth

free uroterga reddish; aedeagus a little arched to the ventral side, Figs
69-70; spermatheca smaller, Fig. 71. Length 1.3 mm... H. micropallida sp. n.
Body stumpy, Fig. 72; eleventh antennomere brown; elytra and fourth

free uroterga yellow-reddish; aedeagus very arched to the ventral side,

Figs 73-74; spermatheca larger, Fig. 75. Length 1.28 mm
Seren. See RENE RER asa H. microkinabaluicola sp. n.
Fourth antennomere transverse; suture of the elytra as long as the
median line of the pronotum; pigidium yellow-reddish; aedeagus with

two ventral thorns, Figs 77-78; spermatheca Fig. 79. Length 2 mm
EIER ET PERS II BODEN CARRE H. spiniventris Sp. n.

ALEOCHARINAE NEL BORNEO 793

- Fourth antennomere longer than wide; suture of the elytra longer than
the median line of the pronotum; pigidium brown-reddish; aedeagus
without ventral thorns, Figs 73-74; spermatheca Fig. 75. Length 2.1 mm
RE AA LR nee One rile Ce RAR AL lee: H. kinabaluensis sp. n.

Hygrochara micropallida sp. n. Figg. 68-71

Holotypus maschio, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 485 m,
29.VIIL.1988, leg. A. Smetana (MHNG).

Paratypi: 4 esemplari, stessa provenienza dell’holotpus; 1 maschio e 1 femmina, Sabah,
Poring Hot Springs, 600 m, 10.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,3 mm. Corpo lucido e giallo-rossiccio con elitre e
uroterghi liberi quarto e base del quinto rossicci; antenne bruno-rossicce con i tre
antennomeri basali e l’undicesimo gialli; zampe gialle. La reticolazione & assente su
tutto il corpo. La punteggiatura del capo è fine, fitta e distinta, quella del pronoto è
superficiale. La granulosità delle elitre è superficiale e quella dell’addome è ben
visibile. Edeago Figg. 69-70, spermateca Fig. 71.

DERIVATIO NOMINIS. Il nome della nuova specie significa «piccolissima e
pallida».

Hygrochara microkinabaluicola sp. n. Figg. 72-75

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N. P., Poring Hot Springs, area
Eastern Ridge tr., 850 m, 28.VIII.1988, leg. A. Smetana, (MHNG).

Paratypi: 1 maschio e 1 femmina, stessa provenienza dell’holotypus.

DESCRIZIONE: Lunghezza 1,28 mm. Corpo lucido e giallo-rossiccio con pigidio
giallo; antenne brune con i tre antennomeri basali gialli; zampe gialle. La reticolazione
del capo e del pronoto è superficiale, quella delle elitre e dell’addome manca. La pun-
teggiatura del capo e del pronoto è quasi indistinta. La granulosità delle elitre è fitta e
superficiale, quella dell’addome è svanita e meno fitta di quella delle elitre. Edeago
Figg. 73-74, spermateca Fig. 75.

DERIVATIO NOMINIS: Il nome della nuova specie significa «piccolissima abita-
trice del Kinabalu».

Hygrochara spiniventris sp. n. Figg. 76-79

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Rv. tr., 1655 m,
11.VIII.1988, leg. A. Smetana (MHNG).

Paratypi: 9 esemplari, stessa provenienza dell’holotypus; 2 es., Borneo, Sabah, Mt.
Kinabalu N.P., HQ Liwagu Rv. tr., 1485 m, 10.VIIL.1988, leg. A. Smetana; 1 maschio, Borneo,
Sabah, Mt. Kinabalu N.P., HQ Liwagu Rv. tr., 1495 m, 22.V.1987, leg. A. Smetana; 1 maschio,
Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu River, 1505 m, 14.VIIL.-1.IX.1988, leg. A.
Smetana

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido e bruno-rossiccio con gli uro-
terghi liberi quarto e quinto bruni; antenne bruno-rossicce con i due antennomeri basali
gialli e l’undicesimo rossiccio; zampe gialle. L’avancorpo è privo di reticolazione,
quella dell’addome è a maglie trasverse fini e ben visibili. La punteggiatura del capo è
fine e distinta, quella del pronoto è superficiale e quella delle elitre è finissima, fit-

794 R. PACE

FIGG. 68-76

Habitus, edeago in visione laterale e ventrale e spermateca. (68-71) Hygrochara micropallida
sp. n. (70-75) Hygrochara microkinabaluicola sp. n. (76) Hygrochara spiniventris sp. n.

tissima e nettamente visibile. L’addome è coperto di granulositä fine, più fitta sui tre
uroterghi liberi basali. Il disco del capo è impresso. Edeago Figg. 77-78, spermateca
Fig. 79.

ALEOCHARINAE NEL BORNEO 795

82

Ficc. 77-82

Edeago in visione laterale e ventrale, spermateca e habitus. (77-79) Hygrochara spiniventris
sp. n. (80-82) Hygrochara kinabaluensis sp. n.

DERIVATIO NOMINIS: Il nome della nuova specie significa «spine ventrali», quelle
dell’ edeago.

Hygrochara kinabaluensis sp. n. Figg. 80-83

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Rv. tr., 1655 m,
11.VIIL.1988, leg. A. Smetana (MHNG).

796 R. PACE

Paratypi: 19 esemplari, stessa provenienza.

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e bruno-rossiccio con capo e
uroterghi liberi quarto e base del quinto bruni, pigidio giallo-rossiccio; antenne brune
con i due antennomeri basali gialli il terzo e l’undicesimo bruno-rossicci; zampe gialle.
L’avancorpo è privo di reticolazione, l’addome è coperto di reticolazione trasversa ben
visibile. La punteggiatura del capo e del pronoto é fitta, fine e distinta. La granulositä
delle elitre è così fitta da simulare una reticolazione. La granulosita dell’addome è più
fitta sugli uroterghi liberi basali che sui posteriori. Il capo presenta una debole impres-
sione discale e il pronoto una debole fossetta mediana posteriore. Edeago Figg. 81-82,
spermateca Fig. 83.

PLACUSINI

Del Borneo è citata (Hammond, 1984) una specie considerata largamente
diffusa nel Sudest Asiatico, Placusa acuminata Kraatz, 1859. Nel materiale in esame
non ho riconosciuto questa specie. Ho esaminato 8 esemplari della serie tipica di
Placusa acuminata così etichettati: Ceylon, J. Nietner, Placusa acuminata Kr. (DEI).

Ho scelto come lectotipo un maschio il cui edeago è illustrato alle Figg. 88-89.
Presente designazione. La spermateca di un paralectotipo è quella di Fig. 90.

CHIAVE DELLE SPECIE DEL GENERE PLACUSA ERICHSON, 1837, NEL BORNEO

1 Pronoto con due fossette discali; elitre con reticolazione obliqua molto
trasversa; femori robusti; edeago Figg. 85-86, spermateca Fig. 87.
Fun shezza de) mn eee u ee eee P. robustipes sp. n.
- Pronoto senza fossette discali; elitre senza reticolazione o con retico-
lazione superficiale, ma non obliqua o trasversa; femori esili............. 2
2 Pronoto notevolmente più ristretto in avanti che all’indietro, Fig. 91 ....... 3
- Pronoto poco ristretto In avant. Ce CM a STRETTE +
3 Antennomeri nono e decimo molto trasversi; occhi più corti delle

tempie; sutura delle elitre lunga quanto la linea mediana del pronoto;
margine posteriore del sesto urotergo libero del maschio con quattro
corti denti a ciascun lato dell’incavatura mediana; edeago con un
lunghissimo tubulo dell’armatura genitale interna, Figg. 88-89; sperma-
tecahis 190! Lunshe72a25 mm... ... RR P. acuminata Kraatz
- Antennomeri nono e decimo lunghi quanto larghi; occhi più lunghi delle
tempie; sutura delle elitre più corta della linea mediana del pronoto;
margine posteriore del sesto urotergo libero del maschio rettilineo;
edeago con corto tubulo dell’armatura genitale interna, Figg. 92-93.
Eunshezza 22 mm... akon se eee P. superba sp. n.
4 Antennomeri nono e decimo lunghi quanto larghi; sesto urotergo libero
del maschio con margine posteriore plurilobato tra due spine, Fig. 97;
edeasoFigs 195 06%Eunchezza2 imme eee ene P. recensita sp. n.
- Antennomeri nono e decimo molto trasversi; sesto urotergo libero del
maschio con margine posteriore rettilineo o monolobato tra due spine,
Eigs#105 4.2.8 2.24.22 4.222080 LEI 5
3 Corpo unicolore nero; edeago Figg. 99-100. Lunghezza 2,1 mm
O ee DR SR) SIR P. falcifera sp. n.

ALEOCHARINAE NEL BORNEO 797

- Gorpoibicolore DEUNOICTOSSICCLON PET NM ee See 6

6 Pronoto con punti isolati allineati a ciascun lato della linea mediana;
edeago Figg. 103-104; spermateca Fig 102. Lunghezza 2 mm . P evoluta sp. n.

- Pronoto senza punti come sopra; spermateca Fig. 107. Lunghezza
TEEN LR o SCR OR A RP Fr P. subspinigera sp. n.

KEY TO SPECIES OF THE GENUS PLACUSA ERICHSON, 1837, FROM BORNEO

l Pronotum with two impressions on the disc; elytra with very transverse
oblique reticulation; femurs strong; aedeagus Figs 85-86, spermatheca
FOSTE noth 9 ME Sree LA Ln etek EN P. robustipes sp. n.

- Pronotum without impressions on the disc; elytra without reticulation or
with superficial reticulation, but not oblique or transverse; femurs slender . . . 2

2 Bronotummere retractedinsfrontakieH 9 mr. ae 3
- Pronotum:a little retracted insfron@Eig 94 nen. ls sce 4
3 Ninth and tenth antennomeres very transverse; eyes shorter than the tem-

ples; suture of the elytra as long as the median line of the pronotum;

distal border of the sixth free urotergum of the male with four short teeth

to every side of the median hollow; aedeagus with a long tubule of the

inside genital armour, Figs 88-89; spermatheca Fig. 90. Length 2.5 mm

IR naar ie AFFARI PAW A ARR ROSE HA CHORI DA oR EHS P. acuminata Kraatz
- Ninth and tenth antennomeres as long as wide; eyes longer than the

temples; suture of the elytra shorter than of the median line of the pro-

notum; distal border of the sixth free urotergum of the male rectilinear;

aedeagus with short tubule of the inside genital armour, Figs 92-93.

DR rn esting, mme ERE ao aes Re ect Pie P. superba sp. n.
4 Ninth and tenth antennomeres as long as wide; sixth free urotergum of

the male with distal border plurilobed among two thorns, Fig. 97;

aedeaous Fos 95 96 "Éenoth2lmmi re 2 202 P. recensita Sp. n.
- Ninth and tenth antennomeres very transverse; sixth free urotergum of

the male with rectilinear distal border or monolobed between two thorns,

ENG NYS) PRE CR ER: AR RI, NE REN AY A PRE I
5 Body unicolourous black; aedeagus Figs 99-100. Length 2.1 mm

5 BL! SR E E I I ORRORI MC a nla be Ce dite AR ro ae ame P. falcifera sp.
- Bodyibieolourousibrownandzzeddish... 22 cesar ae ee 6
6 Pronotum with isolated points lined up to every side of the median line;

aedeagus Figs 103-104; spermatheca Fig 102. Length 2 mm .. P evoluta sp. n.
= Pronotum without points as above; spermatheca Fig. 107. Length

IE TETTI RS Re ae RE AO P. subspinigera sp. n.

>

Placusa robustipes sp. n. Figg. 84-87

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ at Liwagu Rv., 1500 m,
21.V.1987, D.E Bright collector (MHNG).

Paratypi: 1 maschio e 2 femmine, stessa provenienza dell’holotypus; 4 es., Borneo
Sabah, Mt. Kinabalu Nat Pk., HQ 1560 m, 24.1V.1987, D.E Bright collector; 1 maschio, Borneo,
Sabah, Mt Kinabalu Nat. Pk., HQ at Livagu Rv., 1500m, 21.V.1987, beating foliage, leg. D.E.
Bright.

798 R. PACE

Fico. 83-90

Spermateca, habitus e edeago in visione laterale e ventrale. (83) Hygrochara kinabaluensis
sp. n. (84-87) Placusa robustipes sp. n. (88-90) Placusa acuminata Kraatz, lectotypus maschio
e paralectotypus femmina.

DESCRIZIONE: Lunghezza 1,9 mm. Corpo bruno e lucidissimo con addome
debolmente opaco: antenne brune con i due antennomeri basali bruno-rossicci; zampe
giallo-rossicce. La reticolazione del capo e dell’addome manca, quella del pronoto è
obliqua, ondulata e superficiale, quella delle elitre & pure obliqua, molto trasversa e

ALEOCHARINAE NEL BORNEO 799

netta. La punteggiatura del capo & superficiale e rada, quella del pronoto manca tranne
per forti punti isolati distribuiti come da Fig. 84, quella delle elitre & netta. L’addome
è coperto di granulosità fitta. Edeago Figg. 85-86, spermateca Fig. 87.

DERIVATIO NOMINIS: Il nome della nuova specie significa «zampe robuste».

Placusa superba sp. n. Figg. 91-93
Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat.Pk., Poring Hot Springs, 485 m,
24.VII.1988, leg. A. Smetana (MHNG).
DESCRIZIONE: Lunghezza 2,2 mm. Corpo bruno e opaco con addome debol-
mente lucido. Antenne brune con antennomero basale giallo-rossiccio; zampe giallo-
rossicce. La reticolazione del capo é superficiale, quella del resto del corpo manca. La

granulosita dell’avancorpo è saliente, quella dell’addome è superficiale. Edeago
Figg. 91.

Placusa recensita sp. n. Figg. 94-97

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480 m,
15.V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e bruno con pronoto e addome
giallo-bruni; antenne brune con i due antennomeri basali rossicci; zampe giallo-
rossicce. La reticolazione del capo è ben visibile, quella del pronoto manca, quella
delle elitre e dell'addome è superficiale. La granulosita del capo e delle elitre è distinta,
quella del pronoto e dell'addome è superficiale. Edeago Figg. 95-96, sesto urotergo
libero del maschio Fig. 97.

DERIVATIO NOMINIS: Il nome della nuova specie significa «passata in rassegna».

Placusa falcifera sp. n. Figg. 98-100

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m,
6.V.1987, leg. Burckhardt & Löbl (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e nero pece; antenne nere con 1
tre antennomeri basali gialli e i tre seguenti rossicci; zampe gialle. La reticolazione del
capo e dell’addome è ben visibile, quella del pronoto manca e quella delle elitre è
superficiale. La granulosità del capo è poco distinta, quella del pronoto e dell’addome
è saliente, quella delle elitre è superficiale. Edeago Figg. 99-100.

DERIVATIO NOMINIS: Il nome della nuova specie significa «portatrice di falce».
La lama ventrale dell’edeago, infatti, ha forma di falce.

Placusa evoluta sp. n. Figg. 101-105

Holotypus maschio, Sabah, Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, leg.
Burckhardt & Löbl (MHNG).

Paratypus: 1 femmina, Borneo, Sabah, Poring Hot Springs, 500 m, 6.V.1987, leg.
Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido e bruno con pronoto rossiccio e
addome nero-bruno con pigidio rossiccio; antenne rossicce con i tre antennomeri basali
gialli e l’undicesimo bruno-rossiccio; zampe brune con tibie e tarsi giallo-bruni.

800 R. PACE

Ed}
1 mm

ZAR -:
1 a ANAL
\

paul

SO =

0,1 mm

95 96

Fico. 91-97

Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. (91-93)
Placusa superba sp. n. (94-97) Placusa recensita sp. n.

L’avancorpo non è reticolato, l’addome presenta una reticolazione molto superficiale.
La punteggiatura del capo è fitta e ben visibile, quella del pronoto è superficiale tranne
che per i punti forti distribuiti come da Fig. 101. La granulosità delle elitre è fitta e
distinta, quella dell’addome è ben visibile, ma assente sulla linea mediana di ciascun

urotergo libero. Edeago Figg. 103-104, spermateca Fig. 102, sesto urotergo libero del
maschio Fig. 105.

ALEOCHARINAE NEL BORNEO 801

Fica. 98-105
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
(98-100) Placusa falcifera sp. n. (101-105) Placusa evoluta sp. n.

DERIVATIO NOMINIS. La nuova specie prende nome di evoluta perché il suo
edeago si presenta complesso rispetto alla semplicita dell’edeago di altre specie di
Placusa.

802 R. PACE

Placusa subspinigera sp. n. Figg. 106-107

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Riv. trail, 1500-1550
m, 27.1V.1987, leg. A. Smetana (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e bruno con pronoto bruno-
rossiccio; antenne brune con i due antennomeri basali giallo-bruni; zampe gialle. Il
corpo è privo di reticolazione. La granulosità dell’avancorpo è finissima, fitta ed
evidente, quella dell’addome è ben visibile. Spermateca Fig. 107.

DERIVATIO NOMINIS: La nuova specie prende nome di «vicina a spinigera»
perché dal confronto con la spermateca degli esemplari della serie tipica da me esami-
nati (8 maschi e 2 femmine etichettati Ceylon, J. Nietner, Placusa spinigera Kr), la
nuova specie si mostra sicuramente e tassonomicamente Vicina.

BOLITOCHARINI

I generi Erastriusa, Pseudatheta e Neoleptusa erano finora sconosciuti per il
Borneo (Hammond, 1984).

CHIAVE DELLE SPECIE DEL GENERE ERASTRIUSA PACE, 1990, NEL BORNEO

l Pronoto con depressioni laterali; sutura delle elitre lunga quanto la linea
mediana del pronoto; spermateca Fig. 109; sesto urotergo libero della
femmina Fie, 110 /ÆEunehezza2 moe eee E. masculina sp. n.

- Pronoto senza depressioni laterali; sutura delle elitre più lunga della
linea.mediana del'pronotor n.9) DR see, oe nae 2

2 Capo largo quanto 1l pronoto; corpo bruno con addome nero-bruno;
edeago Figg. 112-113; sesto urotergo libero del maschio Fig. 114.
lEunehezza2 2 mm en... eh. A RUE ae E. borneensis sp. n.

- Capo più stretto del pronoto; corpo prevalentemente giallo-rossiccio . ...... 3

3 Corpo interamente giallo-rossiccio; pronoto con una plica posteriore a
ciascun lato del solco mediano; undicesimo antennomero giallo; edeago
senza lamella ventrale Figg. 116-117; spermateca Fig. 118. Lunghezza
132,000 Ree u N ee Re SU ty giles ayo as E. minima sp. n.

- Corpo tricolore giallo-rossiccio, rossiccio e bruno; pronoto senza plica;
undicesimo antennomero bruno-rossiccio; edeago con lamella ventrale
Figg. 120-121; spermateca Fig. 122. Lunghezza 2 mm ...... E. lobifera sp.

=

KEY TO SPECIES OF THE GENUS ERASTRIUSA PACE, 1990, FROM BORNEO

1 Pronotum with lateral depressions; suture of the elytra as long as the
median line of the pronotum; spermatheca Fig. 109; sixth free urotergum
ofsthe female Ris 110. TÉencth 2 mm ee eee E. masculina sp.

- Pronotum without lateral depressions; suture of the elytra longer than the
median line‘of the pronotum VU NE 21.22.02 1:02 SSR 2

2 Head as wide as the pronotum; body brown with black-brown abdomen;
aedeagus Figs 112-113; sixth free urotergum of the male Fig. 114.
kensth; 22mm: 22:5 Sbst ey Ree E. borneensis Sp. n.

- Head narrower than the pronotum; body predominantly yellow-reddish...... 3

>

ALEOCHARINAE NEL BORNEO 803

3 Body entirely yellow-reddish; pronotum with a back fold to every side
of the median furrow; eleventh antennomere yellow; aedeagus without
ventral thin plate Figs 116-117; spermatheca Fig. 118. Length 1.25 mm
RD RE ER de Se OR NOOR e LER. E. minima sp. n.
- Body tricolorous yellow-reddish, reddish and brown; pronotum without
fold; eleventh antennomere brown-reddish; aedeagus with ventral thin
plate Figs 120-121; spermatheca Fig. 122. Length 2 mm. ... E. lobifera sp. n.

Erastriusa masculina sp. n. Figg. 108-110

Holotypus femmina, Borneo-Sabah, Mt. Kinabalu N. P., Headquarters, 1558 m,
2.1X.1988, D.E. Bright collector (MHNG).

Paratypus: 1 femmina, stessa provenienza dell’holotypus.

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido e rossiccio con pronoto e pigidio
giallo-rossicci, elitre bruno-rossicce; antenne bruno-rossicce con i due antennomeri
basali giallo-rossicci e i tre successivi rossicci; zampe gialle. Corpo senza retico-
lazione. La punteggiatura del capo è evidente e quella del pronoto è superficiale. La
granulosità delle elitre è superficiale quella dell’addome è saliente. Il capo presenta un
solco longitudinale discale, il pronoto mostra una depressione a ciascun lato del solco
mediano. Spermateca Fig. 109, sesto urotergo libero della femmina Fig. 110.

DERIVATIO NOMINIS: La nuova specie presenta il sesto urotergo libero della
femmina con caratteri del margine posteriore che si trovano normalmente nei maschi.
Per questo prende nome di «dotata dei caratteri del maschio».

Erastriusa borneensis sp. n. Figg. 111-114

Holotypus maschio, Borneo-Sabah, Mt. Kinabalu N. P., Layang Layang, 2610 m,
2.V.1987, D.E. Bright collector (MHNG).

Paratypus: 1 maschio, stessa provenienza.

DESCRIZIONE: Lunghezza 2,2 mm. Corpo lucido e bruno con addome nero-
bruno; antenne bruno-rossicce con i due antennomeri basali rossicci e apice dell’undi-
cesimo giallo-rossiccio; zampe giallo-rossicce. La reticolazione del capo e delle elitre
è netta, quella del pronoto è vigorosa e quella dell’addome è distinta. La punteggiatura
del capo è ombelicata, superficiale e assente sulla fronte. La granulosità del pronoto e
delle elitre è assai poco visibile, quella dell’addome è saliente sui tre uroterghi liberi
basali e superficiale sui restanti. Edeago Figg. 112-113, sesto urotergo libero del
maschio Fig. 114.

Erastriusa minima sp. n. Figg. 115-118

Holotypus maschio, Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt
& Löbl (MHNG).

Paratypi: 2 femmine, Sabah, Mt. Kinabalu, 1550 m, 28.1V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,25 mm. Corpo lucido e giallo-rossiccio; antenne
giallo-rossicce con i due antennomeri basali e l’undicesimo gialli; zampe gialle. Corpo
senza reticolazione. La granulosità del corpo è superficiale. La fronte è solcata longi-
tudinalmente. Il pronoto presenta una plica posteriore a ciascun lato del solco media-
no. Edeago Figg. 116-117, spermateca Fig 118.

DERIVATIO NOMINIS. Il nome della nuova specie significa «piccolissima» a
motivo della taglia corporea assai ridotta.

804 R. PACE

111

12 113

Ficc. 106-114
Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero della femmina
(110) e del maschio (114). (106-107) Placusa subspinigera sp. n. (108-110) Erastriusa mascu-
lina sp. n. (111-114) Erastriusa borneensis sp. n.

Erastriusa lobifera sp. n. Figg. 119-123

Holotypus maschio, Sabah, Mt. Kinabalu, 1750 m, 21.1V.1987, leg. Burckhardt & Löbl
(MHNG).

Paratypi: 1 maschio e 1 femmina, stessa provenienza; 1 maschio, Sabah, Mt. Kinabalu,
1500 m, 21.V.1987, leg. Burckhardt & Löbl; 1 femmina, Borneo, Sabah, Mt. Kinabalu N. P.,
Headquarters, 1558 m, 9.VIIL.1988, D.E. Bright collector.

ALEOCHARINAE NEL BORNEO 805

118

0,1 mm

122

120 121

Fısc. 115-123
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
(115-118) Erastriusa minima sp. n. (119-123) Erastriusa lobifera sp. n.

DESCRIZIONE: Lunghezza 2 mm. Corpo lucido. Capo rossiccio, pronoto giallo-
rossiccio, elitre brune con base giallo-bruna, addome giallo-rossiccio con quarto
urotergo libero bruno; antenne bruno-rossicce con i due antennomeri basali giallo-
rossicci; zampe giallo-rossicce. La reticolazione del capo è superficiale, quella del pro-
noto è netta e quella delle elitre e dell’addome è ben visibile. La punteggiatura del capo
è distinta. La granulosità del pronoto è confusa nell’evidente reticolazione, quella delle
elitre è ben visibile e quella dell’addome è saliente. Edeago Figg. 120-121, spermateca
Fig. 122, sesto urotergo libero del maschio Fig. 123.

806 R. PACE

CHIAVE DELLE SPECIE DEL GENERE PSEUDATHETA CAMERON, 1920, NEL BORNEO

1 Corpo prevalentemente nero: solo gli omeri, il pigidio e l’undicesimo
antennomero giallo-rossicci. Edeago Figg. 125-126, spermateca
He EunmEhezza2 MN ieee eee ee P. borneensis sp. n.
- Nessuna parte del corpo é nera; undicesimo antennomero bruno.......... 2
2 Pronoto rossiccio, addome unicolore giallo-bruno. Edeago Figg. 129-

130, spermateca sinuosa, Fig. 127. Lunghezza 1,7 mm. . P. kinabaluensis sp. n.
- Pronoto giallo-rossiccio, addome giallo-rossiccio con una fascia giallo-
bruna. Spermateca non sinuosa Fig.133. Lunghezza 1,8 mm . P. seditiosa sp. n.

KEY TO SPECIES OF THE GENUS PSEUDATHETA CAMERON, 1920, FROM BORNEO

| Body predominantly black: only the humeruses, the pigydium and the
eleventh antennomere yellow-reddish. Aedeagus Figs 125-126, sper-
mathecawica /aLenoth 221 mm. een eee ee P. borneensis sp. n.
- No portion of the body is black; eleventh antennomere brown ............ 2
2 Pronotum reddish, abdomen unicolorous yellow-brown. Aedeagus

Figs 129-130, spermatheca sinuous, Fig. 127. Length 1.7 mm

SE ge ae RE A PR ER ME II P. kinabaluensis sp. n.
- Pronotum yellow-reddish, abdomen yellow-reddish with a yellow-

brown band. Spermateca not sinuous Fig. 133. Length 1.8 mm

SMI ce ia RS SL DE 0 A AIRES LA US à P seditiosa sp. n.

Pseudatheta borneensis sp. n. Figg. 124-127

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ at Liwagu Rv., 1500 m,
25.1V.1987, leg. A. Smetana (MHNG).

Paratypi: 3 es., stessa provenienza; 3 esemplari, Borneo, Sabah, Mt Kinabalu Nat. PK.,
HQ at Livagu Rv., 1500m, 30.V.1987, leg. A. Smetana; 17 es., Borneo, Sabah, Mt Kinabalu Nat.
Pk., Silau-Silau Trail, 1560 m, 3.VIIL.1988, leg. A. Smetana; 4 es., Borneo, Sabah, Mt Kinabalu
Nat. Pk., Silau-Silau Trail, 1550m, 2.VIIL.1988, leg. A. Smetana; 4 es., Sabah, Mt Kinabalu,
1750 m, 27.1V.1987, leg. Burckhardt & Löbl; 1 es., Sabah, Mt Kinabalu, 1750 m, 21.1V.1987,
leg. Burckhardt & Löbl; 56 es., Sabah, Mt Kinabalu, 1500 m, 25.IV.1987, leg. Burckhardt &
Löbl; 1 es., Sabah, E Mt Kinabalu, road Ranau-Kota Kinabalu, 1150 m, 24.V.1987, leg.
Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e nero con omeri e pigidio
giallo-rossicci; antenne bruno-rossicce con i tre antennomeri basali e l’undicesimo
giallo-rossicci; zampe gialle. L’avancorpo è privo di reticolazione, l’addome è coperto
di reticolazione trasversa e superficiale. La punteggiatura del capo è ben visibile ed è
assente sulla fascia longitudinale mediana e sulla fronte. La granulosità del pronoto e
delle elitre è ben visibile, quella dell’addome è superficiale. Edeago Figg. 125-126,
spermateca Fig. 127.

Pseudatheta kinabaluensis sp. n. Figg. 128-131

Holotypus maschio, Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt
& Löbl (MHNG).

Paratypi: 19 es., Sabah, Poring Hot Springs, 550-600 m, V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e rossiccio con elitre brune a
base giallo-bruna, addome giallo-bruno; antenne rossicce con i due antennomeri basali

ALEOCHARINAE NEL BORNEO 807

>=
CAS

0,1 mm

Fico. 124-131

Habitus, edeago in visione laterale e ventrale e spermateca. (124-127) Pseudatheta borneensis
sp. n. (128-131) Pseudatheta kinabaluensis sp. n.

gialli; zampe gialle. L’avancorpo è privo di reticolazione, l'addome è coperto di reti-
colazione superficiale. La punteggiatura del capo è fitta e superficiale. La granulosità
del pronoto e delle elitre è fine, fitta e ben visibile e quella dell'addome è superficiale
e fitta. Le elitre del maschio presentano un tubercolo allungato molto saliente presso

808 R. PACE

l’angolo posteriore interno. Dietro di esso la sutura è saliente. Il quinto urotergo libero
del maschio mostra un tubercolo allungato mediano basale. Edeago Figg. 129-130,
spermateca Fig. 131.

Pseudatheta seditiosa sp. n. Figg. 132-133

Holotypus femmina, Borneo, Sabah, Mt. Kinabalu N.P., Por. H. S., area Kipungit Crk.,
1530 m, 26.VIIL.1988, leg. A. Smetana (MHNG).

Paratypus: 1 femmina, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 550-600
m, 9.V.1987, leg. Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 1,8 mm. Corpo lucido e giallo-rossiccio con capo
giallo-bruno, elitre brune con uroterghi liberi quarto e base del quinto giallo-bruni;
zampe gialle. L’avancorpo è privo di reticolazione, l’addome è coperto di reticolazione
molto superficiale. La punteggiatura del capo è distinta. La granulosità del pronoto è
saliente, quella delle elitre è ben visibile e quella dell'addome è fine e superficiale.
Spermateca Fig. 133.

DERIVATIO NOMINIS: La spermateca della nuova specie ha forma unica nel genere
Pseudatheta, pertanto la nuova specie prende nome di sediziosa o ribelle.

Neoleptusa (Physetotoma) kinabaluensis sp. n. Figg. 134-137

Holotypus femmina, Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu River, 1505 m,
14.VIH-1.KX.1988, leg. A. Smetana (MHNG).

Paratypus: 1 maschio, Sabah, Mt. Kinabalu, 3300 m, Panar Laban, 4.V.1987, leg.
Burckhardt & Löbl.

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e giallo-rossiccio con meta
posteriore delle elitre e il quarto urotergo libero bruni; antenne giallo-rossicce; zampe
gialle. La fronte è nettamente reticolata. La reticolazione del pronoto, delle elitre e dei
tre uroterghi liberi basali è assente, quella del quarto urotergo libero è superficiale e
quella del quinto urotergo libero è netta, tranne alla sua base. La punteggiatura del capo
è fittissima, ben visibile e assente sulla fronte, quella del pronoto è simile a quella del
capo. La granulosità dell'addome è superficiale. Spermateca Fig. 135, edeago Figg.
136-137.

NOTA: Il genere Neoleptusa era finora sconosciuto per il Borneo (Hammond,
1984). L’habitus e la forma generale della spermateca della nuova specie sono simili a
quelli di N. luzonensis Pace, 1990, delle Filippine. La nuova specie ne è distinta per il
differente colore del corpo e delle antenne, per il pronoto meno trasverso e distinta-
mente punteggiato e per la spermateca priva di lunghe docce alla base del bulbo distale.
Di N. luzonensis non è noto l’edeago.

DIESTOTINI

I generi Chledophila e Diestota erano finora sconosciuti per il Borneo
(Hammond, 1984).

CHIAVE DELLE SPECIE DEL GENERE CHLEDOPHILA CAMERON, 1920, NEL BORNEO

1 Corpo unicolore; pronoto reticolato; antenne bicolori rossicce con base
gialla; spermateca Fig. 139. Lunghezza 1,7 mm......... C. borneensis sp. n.
- Corpo bicolore; pronoto non reticolato; antenne unicolori giallo-brune;
Spermateca.Rig:-141 Lunghezza 41] Immun RR C. parallela sp. n.

ALEOCHARINAE NEL BORNEO 809

0,1 mm

rey ARI È
agg nile

lmm

133

0,1 mm

135

137

Fico. 132-139
Habitus, spermateca e edeago in visione laterale e ventrale. (132-133) Pseudatheta seditiosa

sp. n. (134-137) Neoleptusa (Physetotoma) kinabaluensis sp. n. (138-139) Chledophila
borneensis sp. n.

KEY TO SPECIES OF THE GENUS CHLEDOPHILA CAMERON, 1920, FROM BORNEO

1 Body unicolorous; pronotum reticuled; antennae bicolorous reddish with
yellow base; spermatheca Fig. 139. Length 1.7 mm ...... C. borneensis Sp. n.
- Body bicolorous; pronotum not reticuled; antennae unicolorous yellow-

brown; spermatheca Fig. 141. Length 1.8 mm........... C. parallela sp. n.

810 R. PACE

Chledophila borneensis sp. n. Figg. 138-139

Holotypus femmina, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ Liwagu River tr., 1495
m, 12.V111.1988, leg. A. Smetana (MHNG).

Paratypus: 1 femmina, Borneo-Sabah, Mt. Kinabalu N.P., blw Layang Layang, 2600 m,
2-8.V.1987, leg. A. Smetana.

DESCRIZIONE: Lunghezza 1,7 mm. Corpo lucido e bruno-rossiccio; antenne
rossicce con 1 tre antennomeri basali giallo-rossicci; zampe gialle. La reticolazione del
capo e delle elitre è assente, quella del pronoto è distinta e quella dell’addome è
superficiale. La punteggiatura del capo è fittissima. La granulosità del pronoto e
dell'addome è poco saliente, quella delle elitre è ben visibile. La granulosità degli
uroterghi liberi quarto e quinto è più rada di quella degli uroterghi liberi basali.
Spermateca Fig. 139.

Chledophila parallela sp. n. Figg. 140-141

Holotypus femmina, Borneo-Sabah, Crocker Ra., 1550-1650 m, 16.V.1987, leg.
Burckhardt & Löbl (MHNG).

DESCRIZIONE: Lunghezza 1,7 mm. Corpo debolmente lucido e giallo-bruno, con
capo ed elitre bruno-rossicce; antenne bruno-rossicce; zampe gialle. La reticolazione
del corpo è indistinta. La punteggiatura del capo è fine, fitta e profonda. La granulositä
del pronoto e delle elitre è saliente, quella dell’addome è fitta sui quattro uroterghi
liberi basali, sul quinto è rada alla base e assente sul resto dell’urotergo libero.
Spermateca Fig. 141.

DERIVATIO NOMINIS: Il nome della nuova specie deriva dai lati del suo corpo che
sono paralleli.

CHIAVE DELLE SPECIE DEL GENERE DIESTOTA MULSANT & REY, 1870, DEL BORNEO

l Pronoto con due impressioni mediane posteriori; elitre con forti punti
assai radi; armatura genitale interna dell’edeago robusta Figg. 147-148;
spermateca,kie. 149 Enunohezza /Îllimm PP man Ener D. plicae sp. n.

- Pronoto con una fossetta mediana posteriore; elitre con punteggiatura
forte e fitta o assai superficiale e fitta; armatura genitale interna
dell'edeago;esile, Figg! 143-144... bll LIE 2
2 Corpo unicolore giallo-rossiccio; punteggiatura delle elitre superficiale;
quarto antennomero trasverso; margine posteriore del quinto urotergo
libero del maschio con quattro lunghi denti smussati tra due spine
laterali; edeago a lati paralleli, in visione ventrale. Lunghezza 2,5 mm
SA ES dai I ER RAR EE CR NET GER N D. testacea (Kraatz)
= Corpo bicolore rossiccio e bruno; punteggiatura delle elitre forte; quarto
antennomero lungo quanto largo; margine posteriore del quinto urotergo
libero del maschio rettilineo; edeago a lati fortemente convergenti verso
l’apice, in visione ventrale. Lunghezza 2,4 mm ............ D. pellita sp. n.

KEY TO SPECIES OF THE GENUS DIESTOTA MULSANT & REY, 1870, FROM BORNEO

1 Pronotum on the disc with two medians impressions; elytra strongly and
sparingly punctured; internal genital armour of the aedeagus strong, Figs
147-148; spermateca Fig. 149. Length 2.1mm............. D. plicae sp. n.

ALEOCHARINAE NEL BORNEO 811

- Pronotum with a posterior median fovea; elytra strongly and closely
punctured or superficially and closely punctured; internal genital armour
Of the acdeacus:slendemsFigs 14314 zn. NEED RE ET To, 2
2 Body yellow-reddish unicolorous; punctuations of the elytra superficial;
fourth antennomere transverse; posterior border of the fifth free uro-
tergum of the male with four long teeth bevelled among two side thorns;
sides of the aedeagus parallel, in ventral view. Length 2.5 mm
ee ro OR AR MN er RE ER: D. testacea (Kraatz)
- Body bicolorous reddish and brown; punctuation of the elytra strong;
fourth antennomere as wide as long; posterior border of the fifth free
urotergum of the male rectilinear; sides of the aedeagus strongly
convergent toward the apex, in ventral view. Length 2.4 mm . . D. pellita sp. n.

Diestota testacea (Kraatz, 1859)
Bolitochara testacea Kraatz, 1859: 7
Diestota testacea; Fauvel, 1905: 86; Cameron, 1939: 164; Pace, 1984: 15

MATERIALE DA AGGIUNGERE: 15 es., Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs,
485 m, 29. VIII.1988, leg. A. Smetana; 1 es., Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs,
495 m, 30.VIIL.1988, leg. A. Smetana; 3 es., Borneo-Sabah, Mt. Kinabalu, Poring Hot Springs,
500 m, 13.V.1987, leg. Burckhardt & Löbl; 1 es., Sabah, Mt. Kinabalu Nat. Pk., Poring Hot
Springs, 495 m, 25.VIIL.1988, leg. A. Smetana.

DISTRIBUZIONE: Specie distribuita dalle Mascarene alla Regione Orientale, Cina

e Papua Nuova Guinea comprese. Nuova per il Borneo.

Diestota pellita sp. n. Figg. 142-145

Holotypus maschio, Sarawak, Borneo, Mulu Nat. Park, (senza data), leg H. Franz
(NHMW).

Paratypi: 11 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 510 m, 30.VI-
11.1988, leg. A. Smetana; 3 es., Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 495 m, 27.VI-
11.1983, leg. A. Smetana; 13 es., Borneo, Sabah, Crocker Rge. N.P., Hwy. A 3, Km 48 cca, 1000
m, 5.1X.1988, leg. A. Smetana; 7 es., Borneo, Sabah, Poring Hot Springs, 500 m, 6.V.1987, leg.
Burckhardt & Löbl; 4 es., Sabah, Crocker Range, 1270 m, Km 60 r.te Kota Kinabalu-Tambunan,
17.V.1987, leg. Burckhardt & Löbl; 15 es., Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg.
Burckhardt & Löbl; 3 es., Sabah, E Mt. Kinabalu, 1150 m, rte. Ranau-Kota Kinabalu, 24.V.1987,
leg. Burckhardt & Löbl; 5 es., Sabah, Poring Hot Springs, 500 m, 13.V.1987, leg. Burckhardt &
Löbl; 1 maschio e 1 femmina, Sabah, Mt. Kinabalu, Poring Hot Springs, 480 m, 10.V.1987, leg.
A. Smetana.

DESCRIZIONE: Lunghezza 2,4 mm. Corpo lucido e rossiccio con elitre brune
tranne la base rossiccia, quarto urotergo libero bruno; antenne bruno-rossicce con i due
antennomeri basali e l’undicesimo giallo-rossiccio; zampe giallo-rossicce. La punteg-
giatura del capo e del pronoto é fitta e distinta, quella delle elitre & profonda e irrego-
larmente distribuita. L’addome presenta una granulosita saliente. Il pronoto mostra una
fossetta mediana posteriore e una laterale. La reticolazione delle elitre & assai superfi-
ciale. Edeago Figg. 143-144, spermateca Fig. 145.

DERIVATIO NOMINIS. I] nome della nuova specie significa «coperta di pelliccia»,
a motivo della pubescenza del corpo.

Diestota plicae sp. n. Figg. 146-150

Holotypus maschio, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Riv. trail, 1500-1550
m, 27.1V.1987, leg. A. Smetana (MHNG).
Paratypi: 7 es., stessa provenienza.

812 R. PACE

0,1 mm

0,1 mm

Ficc. 140-145
Habitus, spermateca e edeago in visione laterale e ventrale. (140-141) Chledophila parallela
sp. n. (142-145) Diestota pellita sp. n.

DESCRIZIONE: Lunghezza 2,1 mm. Corpo lucido e giallo-rossiccio; antenne
rossicce con i tre antennomeri basali e l’undicesimo giallo-rossicci; zampe giallo-
rossicce. La punteggiatura del capo è profonda solo sul disco, sul resto del capo è fine
e superficiale. La punteggiatura del pronoto è ben visibile e quella delle elitre è
composta di punti forti isolati e irregolarmente distribuiti. La reticolazione delle elitre

ALEOCHARINAE NEL BORNEO 813

FIGG. 146-150

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
(146-150) Diestota plicae sp. n.

è molto superficiale. Edeago con una plica ventrale a ciascun lato, Figg. 147-148, sper-
mateca Fig. 149, sesto urotergo libero del maschio Fig. 150.

DERIVATIO NOMINIS. Il nome della nuova specie significa «della plica», quella
ventrale dell’ edeago.

RINGRAZIAMENTI

Rivolgo i miei più cordiali ringraziamenti a coloro che mi hanno affidato in
studio il materiale oggetto del presente lavoro: il Dr. Ale’ Smetana di Ottawa, il Dr.
Ivan Löbl, gia del Museo di Storia Naturale di Ginevra e il defunto Prof. Dr. H. Franz
di Mödling (Austria). Per il prestito di tipi ringrazio il Dr. PM. Hammond e il Dr.
Brendell del Museo di Storia Naturale di Londra, il Dr. L. Zerche del DEI di
Eberswalde (Berlino) e il Dr. A.F. Newton del Field Museum of Natural History di
Chicago.

814 R. PACE

RÉSUMÉ

Les espèces d’Oligotini, Leucocraspedini, Hygronomini, Placusini,
Bolitocharini et Diestotini de Bornéo (Coleoptera, Staphylinidae). - Quarante-cinq
nouvelles espèces de la sous-famille Aleocharinae recueillies dans le Parc National du
Mt Kinabalu, Bornéo, sont décrites et illustrées. Les espèces décrites appartiennent aux
tribus suivantes: Oligotini, Hygronomini, Bolitocharini et Diestotini. Trois nouvelles
espèces appartiennent au genre Oligota (densa, kinabaluensis et borneensis), une au
genre Cypha (sabahensis), dix-neuf au genre Leucocraspedum (sinuatum, pilosellum,
obliquum, spirasferum, occultum biguttae, mimanaticula, divisum, anaticula, fugiti-
vum, directum, hamifer, dilatatiapex, lamelliferum, audax, cacuminum, anguineatheca,
nechamifer, ventriosatheca) quatre au genre Hygrochara (micropallida, microkinaba-
luicola, spiniventris, kinabaluensis), six au genre Placusa, (robustipes, superba,
recensita, falcifera, evoluta et subspinigera), quatre au genre Erastriusa (masculina,
borneensis, minima, lobifera), trois au genre Pseudatheta (borneensis, kinabaluensis,
seditiosa), une au genre Neoleptusa (kinabaluensis), deux au genre Chledophila,
(parallela et borneensis) et deux au genre Diestota (plicae et pellita). Une clé de toutes
les espèces connues des genres susmentionnés est fournie. Le lectotype est désigné
pour Placusa acuminata Kraatz dont l’édéage et la spermathèque sont illustrés pour la
premiere fois.

BIBLIOGRAFIA

CAMERON, M. 1920. New Species of Staphylinidae from Singapore. Transactions of the
Entomological Society of London 1920: 212-284.

CAMERON, M. 1928. XXIX. New species of Staphylinidae from Borneo. Sarawak Museum
Journal 3: 413-451.

CAMERON, M. 1939. The Fauna of British India, including Ceylon and Burma. Coleoptera,
Staphylinidae vol. 4, part 1: i-xviii + 1410. Taylor & Francis Ed., London.

ERICHSON, G. F 1839. Genera et species staphylinorum, insectorum coleopterorum familiae 1.
Berlin, pp. 400.

FAUVEL, A. 1905. Staphylinides de Java, recueillis par M. le Dr. Kraepelin et M. le Dr.
Koningsberger en 1904. Mitteilungen aus dem Naturhistorischen Museum in Hamburg
22: 77-86.

HAMMOND, P. M. 1984. An annotated Check-List of Staphylinidae (Insecta: Coleoptera) recor-
ded from Borneo. Sarawak Museum Journal 33: 187-218.

KRAATZ, G. 1859. Die Staphyliniden-Fauna von Ostindien, insbesondere der Insel Ceylan.
Archiv für Naturgeschichte 25: 1-45.

LEACH, W. 1819. New Genera. In: SAMOUELLE, G. (Ed.). The Entomologist’s useful
Compendium, London: 1-496.

MANNERHEIM, C. G. 1831. Précis d’un nouvel arrangement de la Famille des Brachelytres de

l’ordre des Insectes Coléoptères. Mémoires de l’Académie impériale des Sciences de St

Pétersbourg 1: 415-501.

MULSANT, E. & REY, CI. 1870. Description d’un genre nouveau de l’ordre des Coléoptères.

Opuscules entomologiques, quatorzième cahier: 194-199.

MULSANT, E. & REY, CI. 1874. Tribu des Brévipennes: Famille des Aléochariens: Septième
Branche: Myrmédoniaires. Annales de la Société d'Agriculture de Lyon (4) 6 (1873):
33-738.

PACE, R. 1984. Aleocharinae delle Mascarene, parte I: tribù Myllaenini, Pronomaeini, Oligotini
e Bolitocharini (Coleoptera Staphylinidae) (XLV Contributo alla conoscenza delle
Aleocharinae). Revue suisse de Zoologie 91: 3-36, 159 figg.

ALEOCHARINAE NEL BORNEO 815

PACE, R. 1985. Hypocyphtini dell’India, Ceylon e Nepal del Museo di Ginevra (Coleoptera
Staphylinidae) (LXV Contributo alla conoscenza delle Aleocharinae). Revue suisse de
Zoologie 92: 77-87, 30 figg.

PACE, R. 1986. Aleocharinae dell’ Asia sudorientale raccolte da G. de Rougemont (Coleoptera,
Staphylinidae) (LXXII Contributo alla conoscenza delle Aleocharinae). Bollettino del
Museo civico di Storia naturale di Verona 23: 139-237, 291 figg.

PACE, R. 1990. Aleocharinae delle Filippine (82° contributo alla conoscenza delle Aleocharinae)
(Coleoptera Staphylinidae). /n: BERTI, N. (Ed.). Miscellanées sur les Staphylins.
Memoires du Museum National d’Histoire Naturelle (A) 147: 57-113, 273 figg.

PACE, R. 1993. Nuove Aleocharinae orientali (Coleoptera Staphylinidae) (XCVI Contributo alla
conoscenza delle Aleocharinae). Bollettino del Museo civico di Storia naturale di Verona
17: 127-180.

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REVUE SUISSE DE ZOOLOGIE 114 (4): 817-824; décembre 2007

Imparfinis mishky (Siluriformes, Heptapteridae) a new species
from the rios Paranä and Uruguay basins in Argentina

Adriana ALMIRON!, Jorge CASCIOTTA!, José BECHARA?, Federico RUIZ DIAZ?,

Cecilia BRUNO, Sabina D’AMBROSIO?, Patricio SOLIMANO? & Paula SONEIRA?

lDivisiön Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo, UNLP,
Paseo del Bosque s/n, 1900 La Plata, Argentina.
E-mail: aalmiron@fcnym.unlp.edu.ar

2 CONICET and Instituto de Ictiologfa del Nordeste, Facultad de Ciencias Veterinarias,
UNNE, Sargento Cabral 2139, 3400 Corrientes, Argentina.

3 Facultad de Ciencias Naturales y Museo, UNLP, Paseo del Bosque s/n, 1900 La Plata,
Argentina.

Imparfinis mishky (Siluriformes, Heptapteridae) a new species from the
rios Parana and Uruguay basins in Argentina. - Imparfinis mishky sp. n.
is described from the rio Parana and rio Uruguay basins in Argentina.
Imparfinis mishky sp. n. is distinguished from other species of the genus by
the following combination of characters: pectoral-fin spine with smooth
anterior and posterior margins; caudal fin deeply forked with dorsal lobe
longer than ventral lobe; adipose fin not reaching caudal fin; body with six
dark saddles across the dorsum, and without a mid-lateral band. It was
found inhabiting well oxygenated, neutral pH, flowing waters, with sandy
or cobble covered bottom.

Keywords: Freshwaters - Siluriformes - Heptapteridae - Imparfinis - new
species - systematics - habitat.

INTRODUCTION

Some years ago several specimens similar to Imparfinis cochabambae (Fowler,
1940) were collected in the rios Parana and Uruguay basins. The study of these
specimens showed that they belong to a new species described below.

The genus /mparfinis includes eighteen species (Bockmann & Guazzelli, 2003).
Two of these species have been recorded from La Plata basin in Argentina: /mparfinis
cochabambae from a single specimen taken from the rio Paraguay basin, and J. hol-
landi Haseman, 1911, which was described from and is apparently restricted to the rio
Iguazu basin.

The genus /mparfinis Eigenmann & Norris, 1900 was erected for a group of
South American catfishes that possess, among other characters, a tooth patch on vomer,
a head that is longer than wide, a short supraoccipital process, and a long cranial
fontanel that extends posteriorly to the base of the supraoccipital process and bears a
bridge behind the eyes. Later on, Mees (1974) re-diagnosed /mparfinis by having “the

Manuscript accepted 09.07.2007

818 A. ALMIRON ET AL.

posterior border of skull as if cut off straight, with but a rudimentary postoccipital
process; fontanel a long slit, reaching to the occiput; dorsal and pectoral spines present
but very inconspicuous, continued as soft rays, the soft part usually longer than the
bony part; maxillary barbells short to moderate in length, varying from scarcely
reaching pectoral base, to reaching to the end of the anal fin; no pectoral pore; adipose
fin comparatively short; eye-rim free”. The presence of a free eye-rim was later
considered of limited generic value by Mees & Cala (1989).

In the last years the genus /mparfinis was not revised, and a phylogenetic
diagnosis of the genus is still pendant. For this reason, there is no consensus about the
species included in this genus (Mees, 1974; Mees & Cala, 1989).

The aim of this paper is to describe a new species collected in the rios Parana
and Uruguay basins and placing it in the genus /mparfinis following the generic
diagnoses given by Eigenmann & Norris (1900) and Mees (1974).

MATERIAL AND METHODS

Measurements were taken using digital calliper to the nearest 0.1 mm. Counts
include holotype, and 13 paratypes (1 ex. cleared and stained). Values of the holotype
are indicated by an asterisk. Vertebral counts were taken from specimens that were
cleared and counterstained following Taylor & Van Dyke (1985); counts excludes
vertebrae corresponding to the Weberian apparatus and the caudal complex centrum.
Institutional abbreviations are as listed in Leviton et al. (1985) with the addition of
Asociaciön Ictiolégica, La Plata, Argentina (AI).

In each cite some of the water quality variables were recorded with calibrated
electrodes, including temperature, pH, conductivity (Hanna, Italy), and dissolved
oxygen (YSI, USA). Water velocity was registered using a digital flowmeter (Global
Water, USA). Transparency was measured using a Secchi disk. Alcalinity was
measured in-situ by titration and colorimetric methods (Hach kit Model FF-2, USA).

Comparative material examined (SL in mm): /mparfinis hollandi Haseman,
1911: AI 211, 3 (1 C&S) ex., 69.4-135.2, Argentina, rio Iguazi basin, arroyo Deseado
(25°47°8.1°S - 54°02°21.1”W). Imparfinis cochabambae (Fowler, 1940): ANSP
69066, holotype, 59.5, Bolivia, Cochabamba, Boca Chapare (figured in
http://acsi.acnatsci.org/). Imparfinis cf. cochabambae: MACN-ict 6971, 1 ex., 83.4,
Argentina, Salta, rio Paraguay basin, rio Bermejo.

RESULTS
Imparfinis mishky sp. n. Figs 1-3, Tables 1-2

HoLoryPE: MACN-ict 8973, 56.6 mm SL, Esteros del Iberä, rio Corriente, Capita Mini
(28°53’15.3”S - 58°22°02.7”W), Rio Parana basin, Argentina, coll: F Ruiz Diaz, April 2006.

PARATYPES: All from Argentina. MHNG 2690.011, 2 ex., 40.5-43.7 mm SL, rio Uruguay
basin, rio Timboy (30°14°20.7”S - 57°47’05.3”W), coll: P. Garcia Tartalo, S. D’ Ambrosio, C.
Bruno & P. Solimano, July 2003. AI 207, 3 ex., 40.6-42.5 mm SL, rio Uruguay basin, rio Timboy
(30°14’20.7”S - 57°47’05.3”W), coll: P. Garcia Tartalo, S. D’Ambrosio, C. Bruno & P.
Solimano, December 2003. AI 208, 1 ex., 47.0 mm SL, rio Parana at Ituzaingò (27°29°54.57S -
56°42’47.0”W), coll: F Ruiz Diaz, March 2003. AI 209, 1 ex., 41.2 mm SL, rio Parana at
Yahapé (27°22°12.1”S - 57°39’ 14.6” W), coll: F. Ruiz Diaz, February 2006. AI 210, 17 (1 C&S)
ex., 21.5-65.6 mm SL, same data as holotype.

NEW IMPARFINIS FROM ARGENTINA 819

Fic. 1-2

Imparfinis mishky sp. n., (holotype) 56.6 mm SL, Esteros del Iberä, rio Corriente, Capita Mini.
(1) lateral view. (2) dorsal view.

DIAGNOSIS: Imparfinis mishky sp. n. is diagnosed by the following combination
of characters: Pectoral-fin spine with smooth anterior and posterior margins; caudal fin
deeply forked with dorsal lobe longer than ventral lobe; adipose fin not reaching cau-
dal fin; body with six dark saddles across the dorsum, and without a mid-lateral band.

DESCRIPTION: Morphometrics of holotype and 13 paratypes are presented in
Table 1. Body elongated, posterior part of the body behind the adipose fin,
compressed. Greatest body depth at adipose-fin origin (Fig. 1). Head 4.2-4.7 times in
standard length. Supraoccipital process short, fontanel long and narrow, continued to
the base of the occipital process, interrupted by a narrow bridge at the rear part of
orbits. Snout tip rounded in dorsal view, upper jaw slightly longer than lower jaw.
Teeth of both jaws in narrow bands, upper tooth band rounded, without posterior pro-
jection; lower tooth band interrupted at symphysis and laterally curved posteriorly.
Nostrils on snout almost form corners of square, posterior nostrils closer to eye rim
than anterior ones. Barbels long and slender; maxillary barbels surpass pelvic-fin
origin; outer mental barbels reach middle of pectoral fin; inner mental barbels inserted
only slightly in advance of outer barbels and extend past pectoral-fin origin. Eyes
dorsolateral, with free rim only along dorsal-rostral part of orbit (Fig. 2). Lateral line
complete and straight, without branches, continued onto base of caudal fin. Dorsal fin
with one weak, smooth spine, extended distally as filament, filamentous portion longer
than spinous part, and 6 branched rays; first branched ray longest. Adipose fin well
developed, its base 6.6-7.1 times in SL. Adipose-fin origin anterior of anal-fin origin;
adipose-fin base extends to posterior of anal-fin base. Anal-fin margin rounded, with

820 A. ALMIRÖN ET AL.

TABLE 1. Morphometric data of the holotype and 13 paratypes of Imparfinis mishky sp. n. SD:
standard deviation.

Holotype Range Mean SD
Standard length (mm) 56.6 32357
Percents of SL
Predorsal distance 3951 32.0-34.7 33.6 0.80
Preventral distance 39.8 39.4-45.7 42.4 155
Preanal distance 66.2 63.8-69.0 67.3 1.45
Prepectoral distance 21.6 20.0-25.9 233 1.76
Body depth 12.7 12.6-15.1 13.8 1.06
Dorsal-fin base 15.6 12.8-15.6 14.0 0.86
Anal-fin base 15.0 13.1-16.3 14.6 0.86
Pectoral-fin length 19.7 16.9-20.1 18.6 1202
Pelvic-fin length 193 15.8-20.2 18.1 1.26
Distance between dorsal
and adipose fins 18.5 16.6-20.4 18.4 0.96
Adipose-fin base 26.5 23.4-26.5 24.7 0.90
Caudal peduncle depth 6.9 6.4-7.8 UD 0.44
Caudal peduncle length 20.0 17.7-20.1 18.8 0.75
Head length 213 21.0-23.6 22.4 0.77
Snout length 8.0 7.7-9.0 8.4 0.36
Horizontal eye diameter 3.9 3.9-4.9 4.5 0.31
Interorbital width 4.4 4.3-5.1 4.7 0.27
Percents of HL
Snout length 31-3 34.9-40.4 37.4 1.86
Horizontal eye diameter 18.5 18.5-22.1 20.2 1.14
Interorbital width 20.6 18.5-23.1 20.9 1922

iv-v, 7-8 (1V,8*) rays. Pectoral fin not reaching ventral-fin origin; fin composed of well
developed, smooth spine continued as slender filament, with filamentous portion
longer than spinous portion; 1,8 rays. Ventral fin with 1,5 rays, rounded in outline,
ventral-fin origin at level of fourth branched dorsal-fin ray. Caudal fin long and deeply
forked; dorsal lobe longer than ventral lobe; i+7/9+1i principal rays. Vertebrae 36. Gill
rakers on first branchial arch, 9.

Coloration of specimens upon capture very similar to those preserved in alco-
hol. Ground color of dorsolateral body surface pale gray, sprinkled all over with minute
dark dots; ventrolateral and ventral surface of body whitish. Dorsal surface of head,
cheek, and maxillary barbel with scattered minute dark spots. Six dark saddles over
dorsum, four anterior saddles larger than others. Anterior most saddle placed just
behind occipital process; second saddle at dorsal-fin origin; third saddle at end of
dorsal-fin base; fourth saddle midway between end of dorsal-fin base and adipose-fin
origin; fifth saddle at adipose-fin origin, and the posterior most at end of adipose-fin
base. Dorsal, pectoral, and adipose fins with many minute spots on surface. Pelvic and
anal fin with scattered chromatophores on rays. Caudal fin with scattered minute dark
spots, more concentrated on ventral lobe.

ETYMOLOGY: The specific epithet mishky is a Quichua word that means sweet.
The epithet was dedicated to Patricia Garcia Tartalo, our friend and student who died
tragically in February, 2006.

NEW IMPARFINIS FROM ARGENTINA 821

Fic. 3

Geographical distribution of /mparfinis mishky sp. n. (1) rio Parana at Yahapé. (2) rio Parana at
Ituzaingé. (3) rio Corriente at Capita Mini (type locality). (4) rio Timboy, rio Uruguay basin.

DISTRIBUTION: Imparfinis mishky sp. n. is known from the rio Parana above the
confluence with the rio Paraguay; rio Corriente, rio Parana basin below the confluence
with the rio Paraguay; and rio Timboy, rio Uruguay basin, all localities are within
Corrientes Province, Argentina (Fig. 3).

HABITAT: Imparfinis mishky sp. n. was only collected in marginal areas of lotic
habitats with high current speeds (Table 2). In the portions of the rios Parana and
Corriente where the species was taken, the bottom was generally sandy with variable
content of gravel or silt, whereas cobble predominated in rio Timboy. The pH fluc-
tuated little around the neutral value. Conductivity was generally low, also although
exceptionally high values were found in rio Corriente, a condition that is not common
in this river and occurs only during pronounced droughts in summer (Casciotta et al.,
2005). Dissolved oxygen concentration was always high, slightly below saturation or
supersaturated (Table 2).

A. ALMIRON ET AL.

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NEW IMPARFINIS FROM ARGENTINA 823

REMARKS: Only two species belonging to the genus /mparfinis were previously
recorded from freshwater in Argentina: /mparfinis hollandi and I. cochabambae (cited
by Castello er al., 1978 as Pimelodella cochabambae).

Imparfinis hollandi is quite different from /. mishky in several characters; I. hol-
landi has an asymmetrical caudal fin with an oblique posterior margin and a long
adipose fin that reaches the caudal fin. In contrast, /. mishky has a nearly symmetrical
forked caudal fin with the upper lobe longer than the lower one, and a short adipose fin
that does not reach the caudal fin. /mparfinis hollandi is restricted to the rio Iguazu
basin (Gömez & Somay, 1989, as Pariolius hollandi; and personal observations)
whereas /. mishky was found in the rio Parana and rio Uruguay basins.

The only other species recorded from Argentina was /mparfinis cochabambae.
Based on information in the original description (Fowler, 1940), this species differs
from J. mishky in having the outer edge of the first pectoral-fin ray serrated (vs.
smooth) and the interorbital distance 4 times in the head length (vs. 4.5-5.1 times). In
addition, /. cochabambae has chromatophores concentrated along lateral line forming
a lateral band that is absent in /. mishky.

The single specimen examined by Castello et al. (1978) that was identified as
Pimelodella cochabambae (MACN-ict 6971) appears to be neither /. mishky nor
I. cochabambae. This specimen differs from /. mishky in having the outer edge of the
pectoral-fin spine serrated (vs. smooth) a deeper caudal peduncle (9.6 vs. 6.4-7.8% of
SL); and a shorter adipose-fin base (20.0 vs. 23.4-26.5% SL). This specimen also
differs from Imparfinis cochabambae in having a greater body depth (6.4 vs. 7.2 in
SL), a smaller eye (6.0 vs. 5.0 in head length, 2.3 vs. 1.8 in snout length, and 1.6 vs.
1.2 in interorbital distance). Thus, although we were unable to assign this specimen to
any named species of /mparfinis, it appears not to be /. cochabambae. Therefore,
following Bockmann & Guazzelli (2003), the presence of /. cochabambae in fresh-
waters environments of Argentina has not been established.

ACKNOWLEDGEMENTS

We greatly benefited from comments on the manuscript by C. Ferraris (USA)
Also we would like to express our gratitude to C. Tremouilles (UNLP) for help with
figures. M. Sabaj (ANSP), allowed us to see the holotype figures of /mparfinis
cochabambae in http://acsi.acnatsci.org/. M. Batistön, gave us lodging in Capita Mint.
J. Casciotta is a researcher of CIC, Buenos Aires Province. This work was partially
financed by Entidad Binacional Yacyretä (EBY) through an agreement with the
Facultad de Ciencias Veterinarias, UNNE (Complementary Acts 16 and 17) and the
Agencia Nacional de Promociön Cientifica y Tecnolögica (ANPCyT) from Argentina,
through the project PICT 2003 N°1-13307.

REFERENCES

BOCKMANN, F A. & GUAZZELLI, G. M. 2003. Family Heptapteridae. (pp. 406-431). In: REIS, R.
E., KULLANDER, S. O. & FERRARIS, C. J., Jr. (eds). Check list of the freshwater fishes of
South and Central America. Edipucrs, Porto Alegre. 729 p.

CASCIOTTA, J., ALMIRON, A. & BECHARA, J. 2005. Peces del Iberä. Habitat y Diversidad.
Grafikar, Sociedad de impresores, La Plata, Argentina, 244 pp.

824 A. ALMIRON ET AL.

CASTELLO, H. P., EHRLICH, M. D., Wats, I. R. & PUIG, A. 1978. Adiciones a la fauna de peces de
los rios Paranä medio y Bermejo. Revista del Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia“, zoologia, XII(9): 119-135.

EIGENMANN, C. H. & NORRIS, A. A. 1900. Sobre alguns peixes de S. Paulo, Brasil. Revista do
Museu Paulista 4: 349-362.

GOMEZ, S. E. & Somay, D. E. 1989. La Ictiofauna del Parque Nacional Iguazu (Argentina). I.
Pariolius hollandi e Hypostomus albopunctatus, primeras citas para Argentina (Pisces
Siluriformes). Limnobios 2(10): 725-728.

LEVITON, A. E., GIBBS Jr., R. H., HEAL, E. & DAWSON, C. E. 1985. Standards in herpetology and
ichthyology: Part I. Standard symbolic codes for institutional resource collections in
herpetology and ichthyology. Copeia 1985: 802-832.

MEES, G. F. 1974. The Auchenipteridae and Pimelodidae of Suriname (Pisces, Nematognathi).
Zoologische Verhandelingen 132: 1-256.

MEES, G. F. & CALA, P. 1989. Two new species of /mparfinis from northern South America
(Pisces, Nematognathi, Pimelodidae). Proceedings C 92(3): 379-394.

TAYLOR, W. R. & VAN DYKE, G. C. 1985. Revised procedures for staining and clearing small
fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119.

REVUE SUISSE DE ZOOLOGIE 114 (4): 825-837; décembre 2007

Larval morphology of reed frogs, Hyperolius kivuensis and
H. viridiflavus, from western Kenya (Amphibia, Hyperoliidae)

Bruno VIERTEL!, Stefan LOTTERS2*, Andrea BAUMGART!, Marc OBERST],
Gerhard EISENBEIS! & Michael VEITH2*
! Zoology Department, Mainz University, D-55099 Mainz, Germany.
2 Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam,
P.O. Box 94062, NL-1090 GB Amsterdam, The Netherlands.
E-mail: loetters@uni-trier.de
* New address: Trier University, Biogeography Dpt., D-54286 Trier, Germany.

Larval morphology of reed frogs, Hyperolius kivuensis and H. viridi-
flavus, from western Kenya (Amphibia, Hyperoliidae).- We describe
external characters, oral disc and oral cavity structures of larval Hyperolius
kivuensis and A. viridiflavus, i.e. exotrophous lentic benthic type IV tad-
poles. External morphology of both is comparable and matches descriptions
of other Hyperolius species. Differences among the two taxa were noted in
the keratinized labial teeth. In H. kivuensis, the spoon-shaped cusped tooth
type is frequent and represented by two different subtypes: (1) “cusped” with
four cusps, the two distal of which only slightly larger than the lateral cusps;
(11) “pointed” with the two distal cusps larger than the two lateral cusps
forming a bifurcation. The latter subtype is rare in A. kivuensis, while most
frequent in H. viridiflavus. In this species, also another type occurs, charac-
terized by two long distal cusps, forming a bifurcation, and two small lateral
cusps; the cusped type with cusps of the same size as in A. kivuensis is rare.
Both species show similar oral cavity structures except for the choanae
position, i.e. arranged in a V that is opened posteriorly in H. kivuensis versus
a V opened anteriorly in H. viridiflavus. Choanae position and keratinized
labial teeth structure may provide diagnostic features for these and other
species of Hyperolius.

Keywords: Anura - labial teeth - oral cavity - tadpoles.

INTRODUCTION

There are approximately 120 described species of Afrotropical reed frogs, genus
Hyperolius (Schigtz, 1999). However, detailed descriptions of the larvae exist for less
than 20 of them (Rödel, 1999; Schigtz, 1999; Channing, 2001; Channing & Howell,
2006). During field surveys in western Kenya, adults and larvae of five syntopic reed
frog species were collected (Lötters er al., 2004). The larvae of Hyperolius kivuensis
Ahl, 1931 and H. viridiflavus (Duméril & Bibron, 1841), for which a large series of
positive species allocation could be obtained (see below), are described here. The
former species is known from eastern and central Africa, while the latter is distributed
in eastern Africa. (Schigtz, 1999; Lötters et al., 2004).

Manuscript accepted 05.07.2007

826 B. VIERTEL ET AL.

CLS TPS
a «| ia Wu o
AZ
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= Sen LE (LS
amare on™

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Fıc. 1

Views of lateral body and oral disc of a Hyperolius kivuensis larva from western Kenya, stage
36 (MHNG 2661.31). Lines equal 1 cm and 1 mm, respectively. Illustration by A. and J.
Channing.

Larval morphology may provide useful characters for species diagnoses and for
assessment of phylogenetic relationships (Orton, 1953, 1957; Starrett, 1973). However,
the known larvae of Hyperolius are similar in external morphology. Differences in size,
body proportions, and colors between the species are apparently too minor for
unambiguous identification or systematic purposes (e.g. Channing, 2001).

More than 100 years ago Heron-Royer & van Bambeke (1889) and Boulenger
(1891) introduced the oral disc morphology including the oral teeth as diagnostic
tadpole characters. The morphology of the oral cavity was also used by Wassersug
(1980, 1984), Viertel (1982), and Wassersug & Heyer (1988) for the diagnosis of
anuran larvae. Oral disc and oral cavity morphology has yet not been studied in
Hyperolius larvae.

The goal of the present study is (1) to describe and to discuss the external, oral
disc, and oral cavity morphology of A. kivuensis and H. viridiflavus larvae, and (ii) to
discuss their utility for Hyperolius taxonomy.

MATERIAL AND METHODS

Egg clutches were obtained from amplexed pairs collected in the field between
16 March and 5 April 2003 in the Kakamega Forest (0°21’ N, 34°51’ E; 1650 m above
sea level), Western Province, Kenya. The adults were diagnosed according to Lötters
et al. (2004). Their tadpoles were reared in captivity, and different developmental
stages were fixed and stored in 4% formaldehyde. Voucher specimens have been
deposited at the Muséum d’histoire naturelle, Geneva (Hyperolius kivuensis: MHNG
2661.31, N = 23; A. viridiflavus: MHNG 2661.42, N = 50).

Larval stages were diagnosed according to Gosner (1960). Body measurements
were taken using the primary landmarks of Altig & McDiarmid (1999a). A W-PL
10x/23 ocular attached to a Zeiss Stemi 2000-C stereomicroscope was applied. The
labial tooth row formula (LTRF) was defined by Altig & Johnston (1989). Photographs
of live larvae were taken with a Contax Aria digital camera to obtain life color infor-
mation.

FIG. 2
The labial teeth of Hyperolius kivuensis in stage 36: (a) cusped teeth of the anterior (upper)
labium (posterior view). (b) non-cusped (pointed) teeth and cusped (bifurcated) teeth (at the
bottom of the micrograph) of the posterior (lower) labium. LDC = long distal bifurcated cusps.

LARVAL MORPHOLOGY OF KENYAN REED FROGS

827

828 B. VIERTEL ET AL.

FIG. 3

The oral cavity of Hyperolius kivuensis in stage 36: the roof of the mouth. PAP = prenarial arena
pustulations, POP = postnarial papillae, PRP = prenarial papillae, IP = infralabial papillae, CH =
choanae, internal nares, NVP = narial valve projection, LRP = lateral ridge papillae, MR =
median ridge, BRAP = buccal roof arena papillae or pustulations (smaller than the height twice
the diameter of the base), VP = prevelar pustulations, GZ = glandular zone, DV = dorsal velum
(terminology according to Wassersug, 1980; Viertel, 1982).

The heads of 25 tadpoles (stage 36) from each species were dissected under the
binocular microscope (carcasses were not maintained as vouchers) to describe the oral
cavities. For scanning electron microscopy (SEM) twelve specimens from each species
were dehydrated in 70%, 80%, 96%, and 100% acetone. After desiccation with liquid
CO; in a critical-point drying apparatus (BAL-TEC CPD 030) the specimens were
coated with gold (sputter coater BAL-TEC SPD 005). A Philips ESEM XL 30 scanning
electron microscope was used.

LARVAL MORPHOLOGY OF KENYAN REED FROGS 829

— 7

Fic. 4

The oral cavity of Hyperolius kivuensis in stage 36: the floor of the mouth. OP = pustulations in
the oral orifice, MO = median papilla in the oral orifice, TA = tongue anlage, LP = lingual papilla,
IP = infralabial papillae, BFAP = buccal floor arena papillae or pustulations (smaller than the
height twice the diameter of the base), BP = buccal pocket, VV = ventral velum, VVP = velar
papilla, FP = filter plate, G = glottis (terminology according to Wassersug, 1980; Viertel, 1982)

RESULTS

THE TADPOLE OF HYPEROLIUS KIVUENSIS AHL, 1931

External morphology: Measurements are as follows (mm): stage 36 (N = 14),
total length (TL) = 34.9 (range 28.8-40.7), body length (BL) = 11.6 (range 10.6-13.1),
BL/TL = 0.33 (range 0.31-0.38). The following description 1s based on a larva stage 36
as shown in Figure 1 (MHNG 2661.31, individual number W19). In dorsal view the
body is elongated and ovoid and is widest at mid-body. The snout is rounded both in
lateral and dorsal views. The eyes are relatively large, somewhat bulging, and not
visible in ventral view. The interorbital distance is equal to the shortest distance to the
tip of the snout. The eyes are positioned laterally and are directed dorsolaterally. The
external nares are ovoid (horizontally elongated), small, and positioned laterally. They

830 B. VIERTEL ET AL.

FIG. 5

Views of lateral body and oral disc of a Hyperolius viridiflavus larva from western Kenya, stage
36 (MHNG 2661.42). Lines equal 1 cm and 1 mm, respectively. Illustration by A. and J.
Channing.

are more closely positioned to the snout than to the eyes. In lateral view the body is
highest at the posterior half. The height is about one half of the body length. The
spiracle is single, sinistral, and attached to the body wall. Its shape is cylindrical and
in length it equals half eye diameter to eye diameter. The spiracle opening is rounded,
directed posteriorly, and located slightly anterior to the mid-axis of eye and vent tube.
The length of the tail represents approximately two thirds of the total length and is
highest at about mid-tail. The total tail height is about three quarters of the body length.
The tail fin height is greater than that of the body. The upper tail fin is larger in height
than the lower. The dorsal fin is highest from mid tail to the posterior quarter of the tail.
The ventral fin is almost as high just before the rounded tip of the tail. The height of
the tail musculature is about half of the maximum tail height. The dorsal fin does not
extend onto the body. The vent tube is short, dextral, posteriorly directed, and linked
to the tail musculature. The oral disc is anteroventral, not emarginated, and entirely
bordered by a row of short round papillae. Few submarginal papillae are present. The
LTRE is 1/3(1, 2). The tooth rows are equal in length, occupying nearly the entire width
of the oral disc except the shorter, most posterior one. Jaw sheaths are finely serrated.
The upper jaw sheath is inversely U-shaped and the lower V-shaped and shorter.

The variation in external morphology of 23 larvae between stages 32 and 40 is
limited to size (Table 1) and LTRF including 1/3, 1/3(1), 1/3(1, 2), 1/3(2), 1/3(1, 3),
and 1/3(3). LTRF observed in specimens in stages 25 and 26 (tadpoles not preserved)
was 1/2, 1/2(1), 1/2(1, 2), and 1/2(2). Gaps occurred in the centers or more to the
periphery of tooth rows.

Two different types of keratinized labial tooth are found. The spoon-shaped
cusped type is often seen and is represented in two different morphological subtypes.
One has four cusps and the two distal cusps were only slightly larger than the lateral
cusps (Fig. 2a). In the other subtype, the two distal cusps are clearly larger than the one
or two lateral cusps (Fig. 2b) and form a bifurcated fork. The non-cusped (pointed)
type is rarely found.

FIG. 6
The labial teeth of Hyperolius viridiflavus in stage 36: (a) pointed teeth with small lateral cusps
in the anterior (upper) labium. (b) bifurcated type of the anterior (upper) labium. LDC = long
distal bifurcated cusps, SLC = small lateral cusp.

LARVAL MORPHOLOGY OF KENYAN REED FROGS 831

832 B. VIERTEL ET AL.

FIG. 7

The oral cavity of Hyperolius viridiflavus in stage 36: the roof of the mouth. For abbreviations
see Figure 3.

In preservative, the larvae are entirely light gray to tan. They are darker on the
dorsal body from dense scattered melanophores and are translucent on the posterior
venter. Melanophores on the tail musculature and the fins are almost uniformly
arranged. The margins of the fins are dark gray and melanophores are observed on the
spiracle. The coloration in life of the body was dorsally tan and ventrally translucent
whitish. The tail musculature was tan and the fins were translucent tan with irregular
dark marbling. A dark line occurred on the tail musculature along one third of tail
length beginning at the base of the tail. The pupil coloration was black and the iris was
bronze with a bright ring around the pupil.

Oral cavity: The description of the roof and floor of the mouth is based on
Figures 3 and 4, respectively. The buccal roof is divided into three areas. The prenarial
arena is positioned in the anterior region of the upper beak. The small prenarial arena
postulations (PAP) are arranged in a horseshoe shape, sometimes in a V. The central
PAP are fused at their base, the lateral PAP are not. The prenarial arena is bordered
caudally by the choanae (CH) or internal nares. They are arranged in a posteriorly open

LARVAL MORPHOLOGY OF KENYAN REED FROGS 833

Fic. 8

The oral cavity of Hyperolius viridiflavus in stage 36: the floor of the mouth. For abbreviations
see Figure 4.

V forming an obtuse angle. The CH is surrounded anteriorly by the PAP and posteriorly
by the narial valve projection (NVP). There is one large prenarial papillae (PRP) per
CH which is flanked by one or two small pustulations and also one small and lobed
NVP on the median margin of each CH. The CH are joined caudally to the postnarial
arena with the single large postnarial papilla (POP) positioned dorsolaterally on each
side. Close to the POP but more laterally are two large lateral ridge papillae (LRP)
equal in size with a separated base, found on each side. The largest of the arenas is the
buccal roof arena which is bordered anteriorly by the flat and broadly based median
ridge (MR) with a serrated margin, and laterally by two buccal roof arena papillae
(BRAP). Around 50 small pustulations are on the buccal roof arena. The buccal roof
arena is bordered caudally by the glandular zone (GZ) and the dorsal velum (DV) with
15-20 small pustulations (VP).

The buccal floor arena is the largest region of the buccal floor. It is bordered
laterally by the buccal pockets (BP) and anteriorly by the tongue anlage (TA). Three
large buccal floor arena papillae (BFAP) on each side are arranged in the form of an

834 B. VIERTEL ET AL.

anteriorly open horseshoe and around 60 pustulations cover the buccal floor arena. Two
large lingual papillae (LP) are positioned on the TA. The ten small pustulations of the
oral orifice (OP) are close to the ventral oral beak anterior to the TA. A single large
blunt or pointed lancet-shaped median papilla of the oral orifice (MO) with a broad
base and sometimes serrated apex is seen between the OP and the TA. One flat and
shovel-like infralabial papilla (IP) with large surface and small pustulations on the
margin is positioned laterally on each side in the level of the MO. The buccal floor
arena is bordered caudally by the ventral velum (VV) of the pitted type with two broad-
based velar papillae (VVP) on each side. Spot-tests with larval stage 32 demonstrated
the same arrangement of oral cavity structures as described for stage 36.

Remarks: Wild-caught larvae attributed to A. kivuensis from the same pond at
the Kakamega Forest (MHNG 2661.32-41, N = 60) well coincide with the external
description given above.

THE TADPOLE OF HYPEROLIUS VIRIDIFLAVUS (DUMERIL & BIBRON, 1841)

External morphology: Measurements are as follows (mm): stage 36 (N = 38),
total length (TL) = 35.4 (range 30.0-39.6), body length (BL) = 11.5 (range 10.3-12.9),
BL/TL = 0.33 (range 0.31-0.36). The following description is based on a larva stage 36
as shown in Figure 5 (MHNG 2661.42, individual number D14): In dorsal view the
body is elongated, ovoid, and widest behind the eyes. The snout is rounded in dorsal
and ventral view. The eyes are relatively large and somewhat bulging. The interorbital
distance is larger than the shortest distance to the tip of the snout. The eyes are posi-
tioned laterally, directed dorsolaterally, and not visible in ventral view. The nares are
slit-like (horizontally), small, and positioned dorsolaterally. They are more closely
positioned to the snout than to the eyes. In lateral view the body is highest at the
posterior half. The height is about one half of the body length. The spiracle is single,
sinistral, and not attached to the body wall. Its shape is cylindrical and the length equals
the eye diameter. The spiracle opening is rounded, directed posteriorly, and located on
the mid-axis of eye and vent tube. The length of the tail represents about two thirds of
the total length and is highest at the anterior half. The total tail height is about two
thirds of the body length. The tail fin height is greater than that of the body. The upper
tail fin is taller than the lower. The dorsal fin is highest from mid tail to the posterior
quarter of the tail. The ventral fin is almost as high just before the rounded tip of the
tail. The height of the tail musculature is less than half of the maximum tail height. The
dorsal fin does not extend onto the body. The vent tube is short, dextral, posteriorly
directed, and linked to the tail musculature. The oral disc is anteroventral, not
emarginated, and entirely bordered by a row of short round papillae. Submarginal
papillae are absent. The LTRF is 1/3(1). The tooth rows are equal in length occupying
nearly the entire width of the oral disc except for the shorter, most posterior one. Jaw
sheaths are finely serrated. The upper jaw sheath has an arched, inverted U-shape, the
lower is V-shaped and is shorter.

The variation in external morphology of 50 larvae between stages 30 and 40 is
limited to size (Table 1) and LTRF including 1/3, 1/3(1), 1/3(1, 2), 1/3 (1, 2, 3), 1/3(1,
3), 1/3(2), and 1/3(3). LTRF observed in specimens in stages 25 and 26 (tadpoles not
preserved) is 1/2, 1/2(1), 1/2 (1, 2), and 1/2(2). Gaps are observed in the centers or
more to the periphery of tooth rows.

LARVAL MORPHOLOGY OF KENYAN REED FROGS 835

TABLE 1. Measurements (mm) and ratios of 23 larvae of Hyperolius kivuensis and 50 larvae of
H. viridiflavus from western Kenya in different stages. In N > 2, the mean is given, followed by
one standard deviation, and the range in parentheses.

stage N total length body length body length/total length
Hyperolius kivuensis

32 1 325 1181 0.34

34 À) 31.0, 32.4 10.6, 11.2 0.34, 0.35

35 1 335 11.4 0.34

36 14 34.9 + 3.1 (28.8-40.7) 11.6 + 0.6 (10.6-13.1) 0.33 + 0.1 (0.31-0.38)
SH 2 36.1, 36.6 201287 0251035

38 1 38.2 122 0.32

39 1 38.6 137 0.36

40 1 40.4 13.4 0.33

Hyperolius viridiflavus

30 1 ZIA 9.4 0.35

33 1 32.9 10.2 0.31

34 1 30.8 11.1 0.36

35 2 35.4, 36.8 10.5, 11.5 0.30, 0.31

36 38 35.4 + 2.2 (30-39.6) 11.5 + 0.6 (10.3-12.9) 0.33 + 0.01 (0.31-0.36)
37 3 36.2 + 2.5 (33.5-38.4) 11.7 + 0.3 (11.5-12.0) 0.30 + 0.1 (0.30-0.34)
38 2 33.4, 40.0 12.7138 0.33,088

39 1 40.6 12.8 0.31

Two different types of keratinized labial tooth are found. A pointed type with
small lateral cusps (Figs 6a, b) is the most common type. The second type is charac-
terized by two long distal cusps forming a bifurcated fork and two small lateral cusps
(Fig. 6b). The cusped type with cusps of the same size as seen in A. kivuensis is rare.
All these types are seen in one individual tadpole. There is no difference between
stages 32 and 36.

In preservative, larvae are entirely translucent gray. They are darker on the
dorsal body from dense scattered melanophores. Melanophores on the tail musculature
are almost uniformly arranged while on the fins they are arranged in patches. On the
spiracle the melanophores are distally missing. The coloration in life of the body was
dorsally brownish with tiny dark brown and occasionally cream spots and ventrally
translucent cream. The tail musculature was tan and the fins were translucent tan with
dark blotches. The pupil coloration was black and the iris was bronze with a bright ring
around the pupil.

Oral cavity: The description of the roof and floor of the mouth is based on
Figures 7 and 8, respectively. The structures and their number are the same as in
H. kivuensis with the exception of the choanae. These are arranged in an anteriorly
open V forming an obtuse angle, in contrast to the latter species with a posteriorly open
V. There is no difference between stages 32 and 36.

Remarks: Wild-caught larvae attributed to A. viridiflavus from the same pond at
the Kakamega Forest (MHNG 2661.43-51, N = 54) well coincide with the external
description given above.

836 B. VIERTEL ET AL.

DISCUSSION

External features in the two species are similar and both are comparable with
other Hyperolius (Rödel, 1999; Schigtz, 1999; Channing, 2001; Channing & Howell,
2006). Hyperolius kivuensis and A. viridiflavus differ in the frequency of occurrence of
the two morphological types of teeth. The main type in the former is the cusped type
with four cusps of nearly the same size. This type is rare in A. viridiflavus, while in
larvae of this species the pointed type with or without small lateral cusps is the main
type. Oral cavity structures of the two species are similar. Only the position of the
choanae shows a clear difference. In H. kivuensis, they are arranged in a V which is
opened posteriorly forming an obtuse angle, while in A. viridiflavus the V is open
anteriorly in an obtuse angle. In some larvae of the latter species, the choanae do not
form an angle.

The external morphology, including the oral disc, the keratinized labial teeth,
and the oral cavity structures characterize both H. kivuensis and H. viridiflavus as
exotrophous lentic benthic generalized tadpoles (Altig & McDiarmid, 1999a, b). The
ventral velum of the pitted type and the oral disc are in connection with the sinistral
position of the spiracle criteria of the larval type IV (Starrett, 1973). This is corro-
borated by the repertoire of structures in the oral cavity that are comparable to those
from European larvae of Bufo bufo, Epidalea calamita, and Rana temporaria of the
same larval type (Viertel, 1985, 1987). These tadpoles are known to filter suspended
particles such as phytoplankton and detritus from the water (Viertel, 1990, 1992). They
also scrape food off the substrate with the keratinized labial teeth in connection with
the oral disc, suspend it in the water current and filter it. We expect larvae of both
Hyperolius species to share this generalized feeding behavior.

Our analyses clearly show that morphological differences between tadpoles of
theses two syntopic Hyperolius species are restricted to a few characters only and at
best visible when applying REM technology. We expect many ecologically equivalent
Hyperolius species to have ecologically — and hence morphologically — similar tad-
poles. This casts doubts that external larval morphology will enable field researchers to
discriminate between all ca. 120 Hyperolius species, even though only a few always
occur sympatrically (e.g. Schigtz, 1999; Channing, 2001; Channing & Howell, 2006).
DNA barcoding may therefore become the future method of choice to identify
Hyperolius tadpoles, although inevitably after fieldwork.

ACKNOWLEDGEMENTS

We are grateful to staff at the Herpetology Department of the National
Museums of Kenya, Nairobi, for logistic support. Fieldwork was sponsored by the
Federal Ministry of Education and Research, Germany (BMB+F) through BIOLOG-
BIOTA to MV and SL (FZ 01LC0025). The Kenya Wildlife Service (KWS) kindly
granted permissions. We are indebted to E. Sehn (Mainz) for her kind help preparing
the larvae for scanning electron microscopy. A. and J. Channing, Stellenbosch, kindly
contributed the drawings.

LARVAL MORPHOLOGY OF KENYAN REED FROGS 837

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i se

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REVUE SUISSE DE ZOOLOGIE

Tome 114 — Fascicule 4

SCHÄFFER, Sylvia & KRISPER, Günther. Morphological analysis of the adult
and juvenile instars of Scutovertex minutus (Acari, Oribatida, Scuto-
VEItICIdae) ns Etat ne nen here

CHAMI-KRANON, Thanaphum, WONGSAWAD, Chalobol & DANKITTIPAKUL,
Pakawin. Description of the male of Hersilia vicina Baehr & Baehr,
1993 from northeastern Thailand, with notes on the albomaculata-
eroup (Araneae, Hersilirdae). Toi LETI RAI

DE CHAMBRIER, Alain, SCHOLZ, Tomas, BELETEW, Moges & MAHMOUD,
Zuheir N. Redescription of Proteocephalus sulcatus (Klaptocz, 1906)
(Cestoda: Proteocephalidea), a poorly known parasite of Clarotes
laticeps (Pisces: Silurtformes) in the Sudan a. m lu ee:

Kim, Byung-Woo. Description of Ambanus jaegeri Sp. n. and of the male of
A. euini (Paik) from Korea (Arachnida: Araneae: Amaurobiidae) . . . .

HONG, Yong. Some new earthworms of the genus Amynthas (Oligochaeta:
Megascolecidae) with male discs from Bogildo Island, Korea......

SCHWENDINGER, Peter J. A taxonomic revision of the family Oncopodidae
VII. A new Oncopus species (Opiliones, Laniatores) from eastern
KT ANT AN pb e cect de RE RO

BOYADZHIEV, Peter S. & TRIAPITSYN, Serguei V. Description of a new
species of Entedonomphale (Hymenoptera: Eulophidae) from
Bulsariawithinotes(en#\\carbonania Nas. dI Ah

PACE, Roberto. Le specie del genere Orphnebius Motschulscky, 1858, nel
Bomeoi(Coleoptera, Staphylinidae) ner an. sn Tee

PACE, Roberto. Le specie di Oligotini, Leucocraspedini, Hygronomini,
Placusini, Bolitocharini e Diestotini nel Borneo (Coleoptera, Staphy-
IEEE TEE EEE e Si RR n)

ALMIRON, Adriana, CASCIOTTA, Jorge, BECHARA, José, Ruiz Diaz, Federico,
BRUNO, Cecilia, D’ AMBROSIO, Sabina, SOLIMANO, Patricio & SONEIRA,
Paula. Imparfinis mishky (Siluriformes, Heptapteridae) a new species
from the rios Parana and Uruguay basins in Argentina............

VIERTEL, Bruno, LOTTERS, Stefan, BAUMGART, Andrea, OBERST, Marc,
EISENBEIS, Gerhard & VEITH, Michael. Larval morphology of reed
frogs, Hyperolius kivuensis and H. viridiflavus, from western Kenya
CAmphibiasily peroliidac) e Re A Re

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Volume 114 — Number 4

SCHÄFFER, Sylvia & KRISPER, Günther. Morphological analysis of the adult
and juvenile instars of Scutovertex minutus (Acari, Oribatida, Scuto-
VECI ANALI TIRANO VIE

CHAMI-KRANON, Thanaphum, WONGSAWAD, Chalobol & DANKITTIPAKUL,
Pakawin. Description of the male of Hersilia vicina Baehr & Baehr,
1993 from northeastern Thailand, with notes on the albomaculata-
croup (Araneae; Hersilidae)....-....-- ste LESLIE

DE CHAMBRIER, Alain, SCHOLZ, Tomä$, BELETEW, Moges & MAHMOUD,
Zuheir N. Redescription of Proteocephalus sulcatus (Klaptocz, 1906)
(Cestoda: Proteocephalidea), a poorly known parasite of Clarotes
laticepsi(Pisces: Siluriformes) in the Sudan? Me PAIE seen

Kim, Byung-Woo. Description of Ambanus jaegeri sp. n. and of the male of
A. euini (Paik) from Korea (Arachnida: Araneae: Amaurobiidae).....

HONG, Yong. Some new earthworms of the genus Amynthas (Oligochaeta:
Megascolecidae) with male discs from Bogildo Island, Korea......

SCHWENDINGER, Peter J. A taxonomic revision of the family Oncopodidae
VII. A new Oncopus species (Opiliones, Laniatores) from eastern
RAT AMAR RE re ER EE CR TL LEE

BOYADZHIEV, Peter S. & TRIAPITSYN, Serguei V. Description of a new
species of Entedonomphale (Hymenoptera: Eulophidae) from
Bulsariaywith notes on E carbonara VASEN LE

PACE, Roberto. The species of the genus Orphnebius Motschulscky, 1858,
fromBorneo (Coleoptera; Staphylinidae) 4 wa. Re

PACE, Roberto. The species of Oligotini, Leucocraspedini, Hygronomini,
Placusini, Bolitocharini and Diestotini from Borneo (Coleoptera,
Staphylinidae)e nad wesen are Cantal ee ee RITI

ALMIRON, Adriana, CASCIOTTA, Jorge, BECHARA, José, RUIZ DIAZ, Federico,
BRUNO, Cecilia, D’AMBROSIO, Sabina, SOLIMANO, Patricio & SONEIRA,
Paula. /mparfinis mishky (Siluriformes, Heptapteridae) a new species
from the rios Parana and Uruguay basins in Argentina ............

VIERTEL, Bruno, LÖTTERS, Stefan, BAUMGART, Andrea, OBERST, Marc,
EISENBEIS, Gerhard & VEITH, Michael. Larval morphology of reed
frogs, Hyperolius kivuensis and A. viridiflavus, from western Kenya
(AmphibiasHyperolüdae)....u.... AE

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Revue suisse de Zoologie: Instructions to Authors

The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific
results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies
that it has been approved by all named authors, that it reports their unpublished work and that it is not being
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of
colour plates and large manuscripts. All papers are refereed by experts.

In order to facilitate publication and avoid delays authors should follow the Instructions to Authors and refer to a
current number of R.S.Z. for acceptable style and format. Papers may be written in French, German, Italian and
English. Authors not writing in their native language should pay particular attention to the linguistic quality of the text.

Manuscripts must be typed or printed, on one side only and double-spaced, on A4 (210 x 297 mm) or equi-
valent paper and all pages should be numbered. All margins must be at least 25 mm wide. Authors must submit
three paper copies (print-outs), including tables and figures, in final fully corrected form, and are expected to retain
another copy. Original artwork should only be submitted with the revised version of the accepted manuscript.

We encourage authors to submit the revised final text on a CD-R, using MS-WORD or a similar software. The
text should be in roman (standard) type face throughout, except for genus and species names which should be for-
matted in italics (bold italics in taxa headings) and authors’ names in the list of references (not in other parts of the
text!), which should be formatted in SMALL CAPITALS. LARGE CAPITALS may be used for main chapter headings
and SMALL CAPITALS for subordinate headings. Footnotes and cross-references to specific pages should be avoided.
Papers should conform to the following general layout:

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, full
name(s) and surname(s) of author(s), and full address(es) including e-mail address(es) if possible.

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should sum-
marise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed by up
to 10 keywords, separated by hyphens, which are suitable for indexing. Some of the terms used in the title may be
omitted from the list of keywords in favour of significant terms not mentioned in the title.

Introduction. A short introduction to the background and the reasons for the work.

Material and methods. Sufficient experimental details must be given to enable other workers to repeat the work.
The full binominal name should be given for all organisms. The International Code of Zoological Nomenclature
must be strictly followed. Cite the authors of species on their first mention.

Results. These should be concise and should not include methods or discussion. Text, tables and figures should
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp.
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by
synonyms, material examined, description, distribution, and comments. All material examined should be listed in
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex sym-
bols should be used rather than “male” and “female” (text file: $= d,£=?).

Discussion. This should not be excessive and should not repeat results nor contain new information, but should
emphasize the significance and relevance of the results reported.

References. The author-date system (name-year system) must be used for the citation of references in the text,
e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown
et al. (1995) or (Brown et al., 1995; White et al., 1996) should be used. In the text authors’ names have to be
written in standard type face. However, in the list of references they should be formatted in SMALL CAPITALS (see
below). The list of references must include all publications cited in the text and only these. References must be
listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated
for each reference. The title of the paper and the name of the journal must be given in full in the following style:
PENARD, E. 1888. Recherches sur le Ceratium macroceros. These, Geneve, 43 pp.

PENARD, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61.
MERTENS, R. & WERMUTH, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp.
HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: WENZEL, R. L. & Tipton, V. J.

(eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XI + 861 pp.

Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit
122 x 180 mm, each table on a separate sheet and numbered consecutively.

Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered
consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering
should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain
from mixing drawings and half tones. Originals of figures (ink drawings, photographs, slides) should be submitted
together with the revised version of the accepted manuscript. Original drawings will not be returned automatically.
The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If scanned figures
are submitted on CD, this should be clearly indicated on the print-out. Scanned line drawings must be saved as TIF files
in bitmap mode with a resolution of at least 600 dpi. Half tone illustrations and photos must have at least 300 dpi reso-
lution.

Legends to figures. These should be typed in numerical order on a separate sheet.

Proofs. Only page proofs are supplied, and authors may be charged for alterations (other than printer’s errors) if
they are numerous.

Offprints. The authors receive a total of 25 offprints free of charge; more copies may be ordered at current prices
when proofs are returned.

Correspondence. All correspondence should be addressed to

Revue suisse de Zoologie, Muséum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland.
Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: danielle.decrouez@ville-ge.ch
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

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