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Full text of "Revue suisse de zoologie."

QLl 
.R484 
v. 118 
no. 4 

2 811 1 
ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 
et du 

MUSÉUM D'HISTOIRE NATURELLE 
de la Ville de Genève 



tome 1 1 8 
fascicule 4 
2011 



t 

M GENÈVE DÉCEMBRE 



2011 



ISSN 0035 



- 418 X 



REVUE SUISSE DE ZOOLOGIE 



TOME 118- FASCICULE 4 

Publication subventionnée par: 
Académie suisse des Sciences naturelles (SCNAT) 
Ville de Genève 
Société suisse de Zoologie 



Comité de rédaction 

DANIELLE DECROUEZ 
Directrice du Muséum d'histoire naturelle de Genève 

ALICE CIBOIS, PETER SCHUCHERT 
Chargés de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), 

B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), 
M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger 
(arthropodes excl. insectes). 

Le comité soumet chaque manuscrit pour évaluation 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 

MUSÉUM 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-121 1 Genève 6, Suisse 



ANNALES 



de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 
et du 

MUSÉUM D'HISTOIRE NATURELLE 
de la Ville de Genève 



tome 1 1 8 
fascicule 4 
2011 



M 



GENÈVE DÉCEMBRE 201 1 ISSN 0035 - 41 8 X 



REVUE SUISSE DE ZOOLOGIE 



TOME 118 - FASCICULE 4 

Publication subventionnée par: 
Académie suisse des Sciences naturelles (SCNAT) 
Ville de Genève 
Société suisse de Zoologie 

Comité de rédaction 

DAM ELLE DECROUEZ 
Directrice du Muséum d'histoire naturelle de Genève 

ALICE CIBOIS. PETER SCHUCHERT 
Chargés de recherche au Muséum d'histoire naturelle de Genève 

Comité de lecture 

A. Cibois (oiseaux). G. Cuccodoro (coléoptères). S. Fisch-Muller (poissons), 

B. Merz (insectes, excl. coléoptères). J. Mari aux (invertébrés excl. arthropodes), 
M. Ruedi (mammifères). A. Schmitz (amphibiens. reptiles). P. Schwendinger 
(arthropodes excl. insectes). 

Le comité soumet chaque manuscrit pour évaluation à des experts d'institutions 
suisses ou étrangères selon le sujet étudié. 

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

Administration 

MUSÉUM 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-1 2 1 1 Genève 6. Suisse 



Revue suisse de Zoologie 118 (4): 599-637; décembre 201 1 



On two Heptathela species from southern Vietnam, vvith a 
discussion of copulatory organs and systematics of the Liphistiidae 
(Araneae: Mesothelae) 

Peter J. SCHWENDINGER 1 & Hirotsugu ONO 2 

1 Muséum d'histoire naturelle de la Ville de Genève, c. p. 6404, CH- 1211 Genève 6, 
Switzerland. E-mail: peter.schwendinger@ville-ge.ch 

2 Department of Zoology, National Muséum of Nature and Science, 4-1-1 
Amakubo, Tsukuba-shi, Ibaraki-ken, 305-005, Japan. E-mail: ono(« kahaku.go.jp 

On two Heptathela species from southern Vietnam, with a discussion of 
copulatory organs and systematics of the Liphistiidae (Araneae: 
Mesothelae). - Songthela australis Ono, 2002 is re-defined on the basis of 
new material from the type locality and from a second locality nearby. The 
maie of this species is described for the first time. A closely related species, 
Heptathela nui sp. n., is described from maies and females collected at two 
other localities in the same province. Variation in the copulatory organs of 
both species is illustrated and notes are given on their biology, particularly 
on their association with an ectoparasitic mite species of the genus Ljunghia 
Oudemans, 1932. "Twig-lining" is reported for one population of H. nui 
sp. n. Songthela australis and H. nui sp. n. are compared with Nanthela ton- 
kinensis (Bristowe in Bristowe & Millot, 1933) and H. tomokunii Ono, 
1997a from northern Vietnam, with H. kimurai (Kishida, 1920), H. kikuyai 
Ono, 1998, Ryuthela owadai Ono, 1997b and R. nishihirai (Haupt, 1979) 
from Japan, and with Liphistius ornatus Ono & Schwendinger, 1990 and L. 
thaleri Schwendinger, 2009 from Thailand. Additional taxonomic charac- 
ters are illustrated for most of thèse species. Judging from apomorphies in 
the maie copulatory organs, S. australis and H. nui sp. n. are sister species, 
but their female copulatory organs are very différent. This challenges the 
generic concepts of Songthela Ono, 2000 and Abcathela Ono, 2000 (the 
latter earlier placed in the synonymy of Heptathela). The genus Nanthela 
Haupt, 2003 has no clear apomorphies, and transitions of diagnostic 
characters between Nanthela, Songthela and Heptathela were found. Thus 
Nanthela and Songthela are here placed in the synonymy of Heptathela. 
Morphology and conflicting terminology of copulatory organs in Liphis - 
tiidae, and the current confusion in Heptathelinae systematics are discussed. 
The copulatory organs of maies and females of Liphistius Schiôdte, 1 849 
and Ryuthela Haupt, 1983 are here considered as more derived than those of 
Heptathela. Some observations on moulting of liphistiid spiders are given. 

Keywords: Taxonomy - new species - Liphistius - Nanthela - Ryuthela - 
Songthela - Ljunghia - Acari. 



Manuscript accepted 14.08.201 1 



600 



P. J. SCHWENDINGER & H. ONO 



INTRODUCTION 

The Heptathelinae (considérée! as a family by Haupt, 2003: 69) is a group of 41 
species (including the new one described in here), which are very distinct in their 
génital morphology from the 49 currently known species of Liphistiinae (ail in the 
genus Liphistius) and which are also geographically separated. Heptathelinae occur in 
eastern Asia, Liphistiinae in southeastern Asia, both ranges roughly divided (but not 
everywhere; Schwendinger, in préparation) by the river Mékong. Originally only 
comprising the genus Heptathela Kishida, 1923, other gênera were added to the 
Heptathelinae as the number of species increased: Ryuthela Haupt, 1983, Abcathela 
Ono, 2000, Songthela Ono, 2000, Vinathela Ono, 2000, Nanthela Haupt, 2003 and 
Sinothela Haupt, 2003. Unfortunately many of the recently added gênera were based 
on characters of a single sex. 

The discovery of heptatheline spiders at four localities in Lam Dong Province, 
including the type locality of Songthela australis, by one of us (PJS) in 2003 yielded 
the previously unknown maie of that species and a new, closely related one. With this 
also came a surprise: the maie copulatory organs of both species are very similar, but 
the female copulatory organs so strongly différent that, according to the current System, 
they would need to be placed in différent gênera. Thus an examination of additional 
heptatheline species had to be carried out to re-evaluate the existing generic concepts 
in this group, and this further revealed serious confusion in the terminology used to 
describe copulatory organs of liphistiid spiders in the literature. 

MATERIAL AND METHODS 

External morphology was studied and drawn with a Zeiss S Vil stereomicro- 
scope, the vulvae (tissue removed with forceps) with a Nikon Optiphot compound 
microscope (both with a drawing tube). SEM micrographs were taken with a Zeiss 
DSM-940A scanning électron microscope; prosoma photos were taken at several focal 
planes with a digital caméra on a Leica MZ APO stereomicroscope and assembled with 
the AutoMontage® System. Hairs were partly or completely omitted in the drawings 
of the maie palp. Body measurements were taken with a stereomicroscope and are 
given in mm. The total body length includes the chelicerae. The carapace length was 
measured in a slightly forward-inclined position, with the anterior and the posterior 
margins at the same focal plane. Lengths of leg articles and palpai articles were mea- 
sured on the dorsal side, from midpoint of anterior margin to midpoint of posterior 
margin, and are given in the following order: total (fémur + patella + tibia + metatarsus 
+ tarsus). 

Terminology of the copulatory organs in Heptathelinae largely follows Song & 
Haupt (1984), but what they called "réceptacles" (other authors called them bursae) are 
here called "receptacular clusters" (except for the H. tomokunii females examined). 
The term "génital plate" is inappropriate for Heptathelinae because that refers to the 
ventral wall of the génital atrium plus the entire bursa copulatrix with its dorsal and 
ventral walls, ail of which are unsclerotised. "Slerotised" refers to areas in the cuticle 
that are hard, stiff and strongly pigmented; "unsclerotised" refers to cuticular structures 
that are thin and unpigmented or thick and moderately pigmented (as is the bursa 
copulatrix) but that are always flexible. We use "cymbial projection" as a shorter form 
of Haupt's (2003: 69) "distal projection on the ventral side (of the cymbium)", which 



HEPTATHELA FROM VIETNAM 



601 



can also be described as the "elongate proventral-distal lobe of the cymbium". 
"Carapace" is used instead of "dorsal plate of prosoma". The informai appellation 
"allotype" refers to the paratype on which the description of the female of the new 
species is based. Positioning (dorsal, ventral, etc.) on the palpai organ refers to an 
unexpanded, outstretched palp. In the following text generic names, which are 
currently in synonymy or which are here placed in synonymy, are given between 
inverted commas. 

Abbreviations not explained in the figure legends are: AME, ALE, PME, PLE 
= anterior (posterior) médian (latéral) eyes; IEBR = Institute of Ecology and Biological 
Resources of the Vietnamese Academy of Science and Technology, Hanoi. Vietnam; 
MHNG = Muséum d'histoire naturelle de la Ville de Genève, Switzerland; MOQ = 
médian ocular quadrangle; MNHN = Muséum national d'Histoire naturelle, Paris, 
France; NSMT = National Muséum of Nature and Science (formerly National Science 
Muséum), Tokyo, Japan. 

TAXONOMY 

Liphistiidae Thorell, 1869 
Heptathelinae Kishida, 1923 
Heptathela Kishida, 1923 

Abcathela Ono, 2000; type species Heptathela abca Ono, 1999; placed in the synonymy of 

Heptathela by Haupt, 2003: 71, 79. 
Nanthela Haupt, 2003; type species Liphistius tonkinensis Bristowe in Bristowe & Millot. 1933; 

syn. n. 

Sinothela Haupt, 2003; type species Heptathela sinensis Bishop & Crosby, 1932; placed in the 
synonymy of Songthela by Platnick, 201 1 (in an earlier version of that online catalogue); 
syn. n. 

Songthela Ono, 2000; type species Heptathela hangzhouensis Chen, Zhang & Zhu. 1981; placed 
in the synonymy of Sinothela by Haupt, 2003: 71; syn. n. 

Vinathela Ono, 2000; type species Heptathela cucphuongensis Ono, 1999; placed in the syno- 
nymy of Heptathela by Haupt, 2003: 71, 79. 

TYPE SPECIES: Liphistius kimurai Kishida, 1920. 

Emended DIAGNOSIS: Distinguished from Ryuthela Haupt. 1983 by maies pos- 
sessing a conductor on sclerite III of the palpai organ and lacking an enlarged denticle 
(= contrategular spine) on its sclerite II; females with 2, 3 or 4 réceptacles or recepta- 
cular clusters on the anterior margin of the bursa copulatrix (if 2, then thèse more wi- 
dely separated from each other than in Ryuthela), or with 2 anteriorly and 2 dorsally on 
the bursa, or with 2 anteroventrally and 2 anterodorsally. Distinguished from Liphistius 
Schiôdte, 1849 by the présence of 7-8 spinnerets (posterior médians more or less dis- 
tinctly fused) and by the lack of clavate trichobothria on tarsi of legs and palps; maies 
without retrolateral tibial apophysis on palp, palpai organ with large tegulum carrying 
two prominent apophyses, contrategulum and subtegulum without apophyses, para- 
cymbium without cumulus; females with leathery but unsclerotised walls of the bursa 
copulatrix devoid of thickened rim and vesicles, gland pores usually restricted to 
réceptacles or receptacular clusters situated anteriorly, dorsally or only slightly antero- 
ventrally on the bursa. 



602 



P. J. SCHWENDINGER & H. ONO 



The two species from southern Vietnam 

Heptathela australis (Ono, 2002) Figs 1-20, 55-58, 67 

Songthela australis Ono, 2002a: 120-122, figs 1-8 (description of female). 
Heptathela australis (Ono, 2002).- Platnick, 201 1 (transferred without explanation in an earlier 
version of that online catalogue; this generic placement is maintained here). 

MATERIAL EXAMINED: NSMT-Ar 9617 (1 maie), IEBR, without registration numbers 
(1 maie and 1 female), MHNG, without registration numbers (ail other spécimens), sample SV- 
03/18; 9 maies (matured early June 2004, 2.VII.2004, 30.VII.2004, 30.VI.2006, 10.V.2007, 
24.V.2007, May 2007, 17.IV.2008, 27.IV.2008), 10 females and 1 juvénile; near Dambri 
Waterfall, about 18 km N of Bao Loc, 11°38 , 42"N, 107°44 , 37"E, 850 m; 1 .-2.IX.2003; leg. P. 
J. Schwendinger. - MHNG, without registration number, sample SV-03/17; 1 maie; Lam Dong 
Province, off the road from Da Hoa (= Da Huoai) to Bao Loc, about 13 km SW of Bao Loc, 
11°27'20 ,, N, 107°43'04"E, 690 m; 3 1.VIII.2003; leg. P. J. Schwendinger. 

Emended diagnosis: Maies of H. australis differ from those of H. tonkinensis 
by a narrower cymbium with a longer and narrower elongate proventral-distal pro- 
jection; paracymbium much longer and basally much narrower, almost forming a right 
angle with the cymbium; an unpigmented and unsclerotised zone présent distoventrally 
on cymbium; marginal tegular apophysis much longer and pointed, terminal tegular 
apophysis with its dorsal edge much less projecting and less dentate; contrategulum 
with two distal edges running parallel to each other: an outer sharp edge and an inner 
dentate edge; the sharp edge with a pronounced, beak-like extension prolaterally, the 
dentate edge not connected with the isolated denticles in the ventro-proximal part of 
the contrategulum; conductor shorter, situated close to the embolus, its apex without 
denticles; embolus longer. Females of H. australis are similar to those of H. hang- 
zhouensis, differing by larger posterior "bursae" (= receptacular clusters) (see Ono, 
2002a: 120, referring to Song & Haupt, 1984: fig. 3c-d). Another distinction is found 
in the shape of the bursa copulatrix in dorsal and ventral view: longer and trapezoidal 
or triangular in H. australis, shorter and rectangular in H. hangzhouensis . 

Description of MALE (matured 10.V.2007): Colouration in alcohol (live spé- 
cimens distinctly darker): Ground colour of carapace light brown, with indistinct dark 
brown W-shaped pattern behind dark ocular mound and with dark brown band along 
anterior and latéral margins. Opisthosoma cream, mottled with light grey; tergites uni- 
formly light greyish brown; sternites light orange-brown; spinnerets cream. Chelicerae 
light brown distally, cream proximally. Legs and palps dorsally uniformly light reddish 
brown except for a dark red-brown distal zone of the cymbium (not including cymbial 
projection); no dark annulation présent. Ventral side of body and limbs generally 
lighter than dorsal side. 

Total length 15.2. Carapace 6.4 long, 5.2 wide, set with a few short, peg-like 
hairs (with blunt tips) on margin (mostly anteriorly and posteriorly), behind ocular 
mound and on coxal élévations, only few longer pointed hairs running over ocular 
mound in a longitudinal row. Eye group 0.92 long, 1 .05 wide. Eye sizes and interdis- 
tances: AME 0.04, ALE 0.59, PME 0.30, PLE 0.53; AME-AME 0.09, AME-ALE 1.14, 
PME-PME 0.04, PME-PLE 0.12, ALE-PLE 0.08. MOQ 0.43 long, front width 0.19, 
back width 0.46. Labium 0.4 long, 1.2 wide. Sternum 2.5 long, 2.2 wide (1.2 on ven- 
tral surface). Anterior margin of sternum distinctly elevated, thus a deep suture présent 
between sternum and labium. Maxillae 2.2 long, 1.4 wide. Promargin of cheliceral 



HEPTA THELA FROM VIETNAM 



603 



groove with 10 small tccth on cach chelicera. Paired tarsal claws with 4-5 tccth; un- 
paired claws without denticles. Measurcmcnts of limbs: palp 1 1.4 (3.4 + 1.9 + 3.9 + 
2.2); leg I 18.7 (5.1 + 2.4 + 3.8 + 4.9 + 2.5); leg II 19.1 (5.0 + 2.4 + 3.7 + 5.3 + 2.7); 
leg III 20.4 (4.9 + 2.5 + 3.8 + 6.2 + 3.0); leg IV 26.5 (6.5 + 2.7 + 5.0 + 8.6 + 3.7). 
"Tibial spurs" (sensu Platnick & Goloboff, 1985) absent on ail legs. Opisthosoma 5.7 
long, 3.6 wide; posterior margin of génital sternite (= sternite of second opisthosomal 
segment) widely rounded, with slightly protruding médian part (Fig. 1 ). 

Palp (Figs 2-15) with a quite long and narrow cymbial projection (Fig. 2); 
distoventral zone of cymbium (below subtegulum) unpigmented and unsclerotised, 
formed by extension of unpigmented and unsclerotised inner (retrolateral distal) side 
of cymbium onto its ventral side (Fig. 3); paracymbium relatively long and narrow, 
with only indistinct proximal tubercle; cymbium and paracymbium almost at right 
angles (Fig. 3). Tegulum with two large apophyses: 1) marginal apophysis long, 
pointed, distad-directed, with a sharp edge running across it from prolateral to retro- 
lateral side (Figs 2-4); 2) terminal apophysis retrolaterad- and slightly proximad- 
directed, with an abruptly narrowed and slightly distad-bent apex (Fig. 3), dorsal side 
of apophysis only moderately extended and carrying a low, weakly dentate edge, dorsal 
and ventral sides of apophysis almost parallel to each other (Figs 3, 5, 10-14); edges of 
both apophyses not in contact with each other. Contrategulum with a few isolated 
denticles ventro-proximally and with two parallel distal edges, the outer one sharp, the 
inner one finely dentate, both unifonnely pigmented (Fig. 3); sharp edge prolaterally 
abruptly truncate, with a beak-like extension pointing toward marginal apophysis of 
tegulum (Fig. 2). Conductor arising ventro-proximally on embolus (= bulbal sclerite 
III), its proximal half fairly wide and fused with base of embolus, its distal half free, 
blade-like, with continuously narrowing sides and pointed apex (Figs 3-9). Embolus 
largely sclerotised, prolateral half strengthened by numerous longitudinal ribs (Figs 
2-5); terminal fringe around opening of sperm duct (= spermophore) hyaline and finely 
serrate (Figs 3-4, 6, 8-9). 

Description of female: See Ono (2002a: 120-122, figs 1-8). Anterior margin 
of sternum elevated as in maie, thus also with a deep suture between sternum and 
labium. "Tibial spurs" présent on legs I-HL Vulva as in Figs 16-19, 55-58. 

Variation: Range of measurements in maies (n=10) and in newly collected 
females with clearly developed vulva (n=10; in parenthèses): body length 12.7-17.6 
(15.3-24.1), carapace length 5.4-7.1 (6.0-9.4), carapace width 4.2-6.2 (4.8-8.1). 
Variation in the overall shape of the vulvae of four new females is not strongly pro- 
nounced (Figs 16-18, 55). The largest female (only it) has a dark brown spot on the 
ventral wall of the bursa copulatrix, and several small sclerotised vesicles (as the ones 
on the receptacular clusters), penetrated by gland pores, sitting on the dorsal wall of the 
bursa copulatrix, in between the four receptacular clusters (Fig. 16). The latter charac- 
ter can be an atavism (see Discussion: évolution of female copulatory organs). In one 
female the posterior margin of the génital sternite is indented (probably from an injury 
sustained in an earlier instar; Fig. 20). For variation in the shape of the palpai organ, 
see Figs 6-9 (conductor), Figs 3, 10 (marginal tegular apophysis) and Figs 5, 10-14 
(terminal tegular apophysis). One maie has a proximally narrow paracymbium 
(Fig. 15). The number of isolated denticles ventro-proximally on the contrategulum 
varies from 2-7. 



604 



P. J. SCHWENDINGER & H. ONO 




FlGS 1-15 

Heptathela australis (Ono, 2002), maie (mature 10.V.2007) from the type locality (1-6), five 
other maies from the type locality (7, 11; 9; 10; 12; 13), maie from Dao Hoa - Bao Loc (8, 14- 
15). (1) Posterior margin of génital sternite, ventral view. (2) Distal part of left palp, prolateral 
view. (3) Same, ventral view. (4) Same, retrolateral view. (5) Same, distal view. (6, 8-9) Embolus 
and conductor, proventral view. (7) Conductor, proventral view. (10) Marginal and terminal apo- 



HEPTA THELA FROM VIETNAM 



605 




Figs 16-20 



Heptathela australis (Ono, 2002). (16) Vulva of largest female, dorsal view; dorsal and latéral 
walls of génital atrium not shown. (17-18) Bursae copulatrices of two other females, dorsal view. 
(19) Schéma of vulva, latéral view (the oviduct is actually anterodorsad- and not posterodorsad- 
directed). (20) Posterior margin of deformed génital sternite, ventral view. BC = bursa copula- 
trix; GA= génital atrium; Od = oviduct; RC = receptacular cluster. Scale lines 1 .0 mm. 

RELATIONSHIPS: Despite pronounced différences in female copulatory organs, 
H. australis appears most closely related to N. nui sp. n. Their palps share distinct 
synapomorphies, and both species occur in the same mountain range, about 80 km 
apart. Their closest known relatives are H. tonkinensis and H. tomokunii from northern 
Vietnam, but more closely related species most likely await discovery in the mountains 
between Da Lat and Hanoi. The maie of the geographically closer H. cucphuongensis 
(from south of Hanoi) is unknown and therefore the relationships of this species are not 
yet clear. 



physis of tegulum, ventral view. (11-14) Terminal apophysis of tegulum, ventral view. (15) 
Outline of paracymbium, ventral view. Co = conductor; CP = cymbial projection; CT = contra- 
tegulum; Cy = cymbium; DE = dentate distal edge of contrategulum; DT = dorsal extension of 
terminal apophysis of tegulum; E = embolus; MA = marginal apophysis of tegulum; PC = para- 
cymbium; PE = prolateral extension of sharp distal edge of contrategulum; SE = sharp distal 
edge of contrategulum; ST = subtegulum; T = tegulum; TA = terminal apophysis of tegulum; 
Z = unpigmented distoventral zone of cymbium. Scale lines 1 .0 mm. 



606 



P. J. SCHWENDINGER & H. ONO 



Distribution: Known only from two localities near Bao Loc village in 
southern Vietnam. 

BiOLOGY: The mature maie was discovered when scraping leaf litter from the 
floor of an evergreen broadleaf forest. He probably had left his burrow in search of 
females and was taking temporary shelter in the leaf litter during the day. Ail other 
spiders were extracted from their burrows in sun-exposed earth banks (with little 
végétation) on one side of a dirt road in an old secondary forest. None of the burrows 
inspected at the beginning of September contained mature maies, and by then the 
mating period was probably at its end. 

Ail burrows were closed with a single trapdoor that had its hinge on the upper 
side of the entrance; the lower rim of the burrow entrance was slightly protruding. 
Trapdoors of females were up to 3.3 cm wide and 2.5 cm long; those of penultimate 
maies 2.0-2.3 and 1.5-1.8 cm, respectively. No spiders were seen when their doors 
were carefully opened with forceps for inspection. They were resting in the depth of 
their burrows and could not be provoked to corne to the entrance by inserting a flexible 
grass stem and disturbing them (which often works for extracting spiders of the genus 
Liphistius). Ail thèse spiders had to be dug up. Large and medium-sized spiders 
(including adult maies) react to disturbance by repeatedly raising their body above the 
ground while the leg tarsi remained on the ground (a behaviour called "tiptoeing") and 
by spreading their chelicerae, as large Liphistius also do. Four egg cases (2.5-3.5 cm in 
diameter and 1.4-1.9 cm high) were built between the end of November and late 
December 2003. Three of them held clusters of 129-164 beige or light yellow eggs 
wrapped in a thin skin made of a coagulated vulval sécrétion and some fine silken 
threads (as présent in ail liphistiid egg cases examined, but less distinct in Liphistius), 
resting on a mesh of fine threads above the bottom of the inner chamber of the egg 
case. Ninery-nine third (or later?) instar juvéniles were taken from the fourth egg case 
in mid-April 2004. Five of them were maies that reached maturity in June 2006, May 
2007 and April 2008, which is presumably earlier in the year than in nature (since the 
adult maie was found at the end of August). Development from eggs to adult maies in 
captivity thus took between one and a half years and three years and four months. 
Females which laid eggs between November and December moulted in the following 
M arc h, May and June. 

Three of the large females, which were collected in the field and subsequently 
laid eggs in Geneva, each carried more than 50 ectoparasitic mites of différent instars 
on their prosoma. Thèse mites were usually sitting on the carapace, but when disturbed 
took shelter on the ventral side, between the coxae and the sternum, and between the 
prosoma and the opisthosoma. They pierced the carapace at numerous places and left 
clearly visible dark round scars that make the spiders look as if they had measles 
(Fig. 67). The carapace, a quite strongly sclerotised plate, is an unlikely place for 
sucking body fluids. Surprisingly, no scars or any other signs of damage are visible on 
the soft membranes between the sclerites of the prosoma or on the opisthosoma. 



HEPTATHELA FROM VIETNAM 



607 



Heptathela nui sp. n. Figs 21-33, 68 

HOLOTYPE: MHNG, without registration number, sample SV-03/21; maie (matured 
10. VI. 2004); Vietnam, Lam Dong Provinee, Mt Penhatt near Quang Trung Réservoir, about 
8 km SW ofDa Lat, 11°52 , 37 ,, N, 108°25'58"E, 1500 m; 6. IX. 2003; leg. P. J. Schwendinger. 

PARATYPES: MHNG, without registration number, sample SV-03/21; 2 maies (matured 
3.V.2004, 10.VII.2004) and 3 females ("allotype" constructed egg case in XII.2003 and moultcd 
5.IV.2004); collected together with the holotype. - NSMT-Ar 9618, 9619 (1 maie, 1 female), 
IEBR, without registration numbers (1 maie, 1 female), MHNG, without registration numbers 
(ail other spécimens), sample SV-03/22; 3 maies (matured early June 2004, 15. VI. 2004, 
21.IV.2005) and 4 females; Cam Ly Waterfall, western outskirts of Da Lat city, 11°56'37 ,, N, 
108 o 25'14"E, 1450 m; 8.-9.IX.2003; leg. P. J. Schwendinger. 

ETYMOLOGY: "Nui", a noun in apposition, is the Vietnamese word for 
"mountain". 

DiAGNOSlS: Maies of the new species can be distinguished from those of the 
closely related H. australis by: anterior margin of sternum low; paracymbium stronger, 
with distinct proximal tubercle; conductor narrower, with more strongly bent apex; 
tegulum with larger marginal apophysis, terminal apophysis more continuously 
narrowing towards apex, its dorsal and ventral sides at an acute angle to each other; 
prolateral side of contrategulum with light-coloured central portion; sharp (outer) distal 
edge of contrategulum convex, broadly rounded, without beak-like extension; denticles 
on inner edge of contrategulum running down to ventro-proximal area in a continuous 
row; embolus shorter, its terminal fringe more coarsely serrate. Females of H. nui 
sp. n. differ from those of H. australis by: bursa copulatrix shorter, with more strongly 
inclined latéral margins; receptacular clusters smaller, with more or less distinct stalks, 
ail four sitting on the anterior margin of the bursa copulatrix. 

DESCRIPTION: MALE (holotype). Colouration in alcohol (live spécimens dis- 
tinctly darker): Carapace and legs light reddish brown; ocular mound dark; bulbal 
sclerites and distal part of cymbium red-brown, cymbial projection light brown. No 
annulation on legs or dark pattern on carapace and opisthosomal tergites. Ventral side 
of body and limbs generally lighter than dorsal side. Opisthosoma cream, mottled with 
light grey; tergites uniformly light greyish brown. 

Total length 14.4. Carapace 6.3 long, 5.0 wide, set with a few short, peg-like 
hairs (with blunt tips) on margin (mostly anteriorly and posteriorly), behind ocular 
mound and on coxal élévations; only few long pointed hairs running over ocular 
mound in a longitudinal row. Eye group 0.87 long, 0.99 wide anteriorly, 0.91 pos- 
teriorly. Eye sizes and interdistances: AME 0.04, ALE 0.60, PME 0.32, PLE 0.50; 
AME-AME 0.14, AME-ALE 0.07, PME-PME 0.05, PME-PLE 0.08, ALE-PLE 0.06. 
MOQ 0.43 long, front width 0.21, back width 0.52. Labium 0.5 long, 1.2 wide. 
Sternum 2.7 long, 2.2 wide (1.3 on ventral surface). Anterior margin of sternum low, 
thus only a shallow furrow présent between sternum and labium. Maxillae 2.2 long, 1 .4 
wide. Promargin of cheliceral groove with 10 teeth on right chelicera, 12 on left. Paired 
tarsal claws with 3-5 teeth; unpaired claws without denticles. Measurements of limbs: 
palp 12.1 (3.6 + 2.1 + 4.1 + 2.3); leg I 19.7 (5.6 + 2.5 + 4.0 + 5.0 + 2.6); leg II 19.3 
(5.2 + 2.4 + 3.8 + 5.2 + 2.7); leg III 20.6 (5.0 + 2.5 + 3.8 + 6.2 + 3.1); leg IV 25.9 (6.6 
+ 2.7 + 4.9 + 8.2 + 3.5). "Tibial spurs" absent on ail legs. Opisthosoma 5.6 long, 3.7 
wide; posterior margin of génital sternite widely rounded, with an indistinct invagi- 
nation in the middle (Fig. 21). 



608 



P. J. SCHWENDINGER & H. ONO 



Palp (Figs 22-26) with paracymbium carrying a proximal tubercle (Fig. 23); 
cymbium and paracymbium almost at right angles to each other; distoventral zone of 
cymbium (below subtegulum) unpigmented and unsclerotised (Fig. 23). Tegulum with 
a very long, pointed, distad-directed marginal apophysis with a sharp edge (Figs 
22-25), and with a retrolaterad- and slightly proximad-directed terminal apophysis 
continuously narrowing to a rounded, distad-bent apex (Figs 23, 25); dorsal side of 
terminal apophysis less extended than in H. australis, carrying a weakly dentate edge 
(Fig. 25) not being in contact with edge on marginal apophysis. Contrategulum with a 
cream central portion on prolateral side (Fig. 22); two parallel distal edges présent: the 
outer one sharp, the inner one finely dentate (Fig. 23); sharp edge convex, widely 
rounded, without beak-like extension (Fig. 22); row of denticles on inner edge running 
down to near ventro-proximal margin of contrategulum and there forming two rows of 
denticles together with a few irregularly arranged additional denticles situated more 
proximally (Fig. 23). Conductor situated ventro-proximally on embolus; proximal 
portion of conductor fairly wide and fused with embolie base; distal portion of 
conductor free, narrow, blade-like, with slightly hook-like apex (Figs 23-26). Embolus 
largely sclerotised, prolateral half strengthened by numerous longitudinal ribs (Figs 22, 
25); terminal fringe around opening of sperm duct (= spermophore) hyaline, coarsely 
serrate, with fairly deep indentations (Figs 22-24, 26). 

FEMALE ("allotype"). Colouration similar to that of maie but slightly lighter; 
palpai tarsi brown. 

Total length 21.6. Carapace 8.3 long, 6.3 wide. Eye group 0.97 long, 1.03 wide 
anteriorly, 1.00 posteriorly. Eye sizes and interdistances: AME 0.06, ALE 0.60, PME 
0.35, PLE 0.50; AME-AME 0.19, AME- ALE 0.14, PME-PME 0.04, PME-PLE 0.06, 
ALE-PLE 0.06. MOQ 0.49 long, front width 0.30, back width 0.52. Labium 1.1 long, 
2.0 wide. Sternum 3.6 long, 2.5 wide (1.9 on ventral surface), its anterior margin 
distinctly elevated, thus separated from labium by a deep suture (not so in maies). 
Maxillae 2.8 long, 1.9 wide. Promargin of cheliceral groove with 11 teeth on each 
chelicera. Paired tarsal claws of legs with 2-4 teeth, unpaired claws without denticles; 
each palpai claw with 2 denticles. Measurements of limbs: palp 13.3 (4.5 + 2.5 + 3.0 
+ 3.3); leg I 15.2 (4.9 + 2.6 + 3.0 + 3.0 + 1.7); leg II 14.6 (4.4 + 2.6 + 2.7 + 3.1 + 1.8); 
leg III 15.5 (4.3 + 2.7 + 2.7 + 3.8 + 2.0); leg IV 21.9 (6.1 + 3.0 + 3.9 + 6.2 + 2.7). 
'Tibial spurs" présent on legs I-III. Opisthosoma 7.7 long, 6.4 wide. Posterior margin 
of génital sternite broadly rounded, with a slight médian invagination. 

Vulva (Fig. 31). Bursa copulatrix short, plano-convex in dorsal and ventral 
view, its anterior margin widely rounded and carrying four anteriad-directed recepta- 
cular clusters on indistinct short stalks, the médian pair slightly larger than the latéral 
one. Génital atrium densely set with fine hairs on dorsal and ventral walls. 

Variation: Range of measurements in maies (n=6) and in females with clearly 
developed vulvae (n=7; in parenthèses) examined: Body length 13.2-16.5 (16.4-22.5), 
carapace length 5.2-6.5 (6.1-8.2), carapace width 4.2-5.3 (4.9-6.5). For variation in the 
shape of the vulva in three females, see Figs 31-33. In one female the receptacular 
clusters are raised on relatively long stalks (Fig. 33), in two other females the stalks are 
stout and rather indistinct (Figs 3 1-32). Variation in the shape of the palpai organ is less 
pronounced than in H. australis: the marginal tegular apophysis carries a subterminal 



HEPTATHELA F ROM VIETNAM 



609 




Figs 21-30 

Heptathela nui sp. n., maie holotype (21-26) and three maie paratypes from Cam Ly Waterfall 
(27-28; 29; 30). (21) Posterior margin of génital sternite, ventral view. (22) Distal part of left 
palp, prolateral view. (23) Same, ventral view. (24) Same, retrolateral view. (25) Same, distal 
view. (26) Embolus and conductor, proventral view. (27) Conductor, proventral view. (28) 
Marginal and terminal apophysis of tegulum, ventral view. (29) Marginal apophysis of tegulum, 
ventral view. (30) Outline of paracymbium, ventral view. PT = proximal tubercle of para- 
cymbium; other abbreviations as in Figs 1-15. Scale lines 1.0 mm. 



610 



P. J. SCHWENDINGER & H. ONO 




Figs 31-33 

Heptathela nui sp. n., three female paratypes. (31) Vulva of "allotype", dorsal view; dorsal and 
latéral walls of génital atrium not shown. (32-33) Bursae copulatrices of two other females, dor- 
sal view. Abbreviations as in Figs 16-20. Scale lines 1.0 mm. 

denticle in two maies (Figs 25, 28), in others it is bare (Fig. 29); the proximal tubercle 
of the paracymbium is very distinct in two maies (Figs 23, 30), less so in others; in 
some maies the scattered denticles below the continuous row of denticles on the 
ventro-proximal side of the contrategulum are indistinct. In one female the posterior 
médian spinnerets are fused only along their entai sides but the apices are still free, 
whereas in ail other conspecific spiders examined thèse spinnerets are completely 
fused. 

Remarks: The paired leg claws of the "allotype" are inflated, especially on the 
posterior legs. This is not so in any of the other spécimens of both species examined 
and thus it was probably caused by préservation. Similar rare cases of inflated claws 
were also observed in alcohol-preserved spécimens of mygalomorph spiders. 

Relationships: Heptathela nui sp. n. possesses a vulva similar to that of 
H. abca from northern Vietnam, and to H. bristowei Gertsch, 1967, H. sinensis and 
H. schensiensis (Schenkel, 1953) from central China. However, strong similarity and 
several likely synapomorphies in the maie palps of H. australis and H. nui sp. n. show 
that thèse species are the most closely related. See also Discussion - Systematics. 

Distribution: Known only from two localities near Da Lat city in southern 
Vietnam. 

BiOLOGY: The spiders examined were collected from a steam bank in an ever- 
green hill forest, from the recess of an earth bank at the side of a path in a pine forest 
(both at Mt Penhatt), and from an earth bank in a park with pine trees (at Cam Ly 
Waterfall). Burrows were as those of H. australis, but at the Cam Ly site (not so on Mt 
Penhatt) they ail had several grass stems attached to their entrances to enlarge the 
sensory area. "Twig-lining" is known from individual populations of several species of 



HEPTATHELA FROM VIETNAM 



mygalomorph trapdoor spiders in Australia, North America and Japan (see Haupt, 1995 
for a summary), and it was also reported for Ryuthela populations (Haupt, 1983: 286) 
and for some H. abca at the type locality (Ono, 1999: 41, figs 13-14). Trapdoors of 
females were up to 2.6 cm wide and 2.0 cm long; those of penultimate maies 1.6-2.1 
and 1.1-1.6 cm, respectively. Large and medium-sized spiders react to disturbance by 
"tiptoeing", spreading their chelicerae and raising their palps. One egg case (2.5 cm in 
diameter and 1.7 cm high) was built on 19. XII. 2004. It held 83 light yellow eggs 
suspended above the bottom of the inner chamber. The female with eggs moulted in 
early April of the following year; other females moulted in January, March, April, 
September. October and November, usually twice per year. Maies (collected in early 
September 2003) became adult between May and July 2004 and in April 2005. 

The largest female (gravid; died before laying its eggs) had about 50 ecto- 
parasitic mites of the genus Ljunghia on its prosoma. Thèse caused the same kind of 
scars on the carapace of the spider (Fig. 68) as described earlier in this paper for 
H. australis. 

Species examined for comparison 

Heptathela tonkinensis (Bristowe, 1933) Figs 34-38 

Liphistius birmanicus Thorell, 1897.- Simon, 1909: 70-71 (misidentification; description of 
maie). 

Liphistius tonkinensis Bristowe in Bristowe & Millot, 1933: 1022, text-fig. 4 (naming of species; 

illustration of palp of maie holotype).- Bristowe, 1976: 4 (listing). 
Heptathela tonkinensis (Bristowe, 1933).- Haupt, 1983: 284-285, figs 8a-b, d. 13e (transfer; 

description and illustration of palp of maie holotype); Platnick & Sedgwick, 1984: 

3 (transfer). 

Vinatela tonkinensis (Bristowe, 1933).- Ono, 2000: 150 (transfer). 

Nanthela tonkinensis (Bristowe, 1933).- Haupt, 2003: 69, 71, figs 51a-b (transfer; illustration of 

palp of maie holotype). 
Here transferred back to Heptathela. 

MATERJAL EXAMINED: MNHN 29170, AR-4104; maie holotype; Vietnam, Tonkin, forêt 
de Khà là; no collecting date; leg. Biaise. 

Emended DIAGNOSIS: The previously published descriptions of the holotype do 
not mention some of the following characters: cymbial projection quite broad and dis- 
tinctly inclined from axis of cymbium (Fig. 34); distoventral zone of cymbium (below 
subtegulum) pigmented and sclerotised as normal; paracymbium short and exceptio- 
nally wide at base, at an acute angle to axis of cymbium (Fig. 35); contrategulum 
distally with a broad band of denticles (arranged in several irregular rows) running 
down to near ventro-proximal margin of contrategulum, no sharp distal edge présent 
(Fig. 36); marginal apophysis of tegulum fairly short, compressed and arched, with a 
sharp distal edge running from prolateral to retrolateral side (Figs 34, 37-38); terminal 
apophysis of tegulum beak-like, dorsally with a large and broadly arched extension 
with a coarsely dentate edge (Figs 34, 36-38) not in contact with edge on marginal 
apophysis; conductor long (about as long as embolus), its distal part fairly distant from 
embolus (Figs 34, 36). "Tibial spurs" absent on ail legs. Posterior margin of génital 
sternite widely and unifonnly rounded, without protruding médian part. Anterior 
margin of sternum not markedly elevated, suture to labium therefore shallow (as in 
maie of H. nui sp. n.). 



612 



P. J. SCHWENDINGER & H. ONO 




/ 



Figs 34-38 

Heptathela tonkinensis (Bristowe, 1933), maie holotype. (34) Distal part of left palp, prolateral 
view. (35) Paracymbium of left palp, ventral view. (36) Left palpai organ, distal view. (37) 
Marginal and terminal apophysis of tegulum of left palp, ventral view. (38) Same, right palp. 
Abbreviations as in Figs 1-15. Scale lines 1.0 mm. 

Remarks: The retrolateral view of the right palp of the holotype given by 
Bristowe (Bristowe & Millot. 1933: text-fig. 4) is detailed and correct, but it gives the 
false impression of a quite long and pointed marginal tegular apophysis, as présent in 
H. australis and H. nui sp. n. A comparison of thèse three species showed that ail of 
them have a fairly sharp distal edge on their marginal tegular apophysis, and in pro- 
and retrolateral view this apophysis always looks pointed (see also Fig. 34). Ventral 
and dorsal views show that in H. tonkinensis this apophysis is actually scale-like, fairly 
short and moderately arched (Figs 37-38; Haupt, 1983: fig 8b; Haupt, 2003: fig. 51B), 
whereas in H. australis and H. nui sp. n. it is long, triangular and truly pointed (Figs 3, 
5. 10,23,25,28-29). 

RelationshipS: Among the hepthatheline species of which maies are known, 
H. tonkinensis currently appears most closely related to H. tomokunii; both are more 
distantly related to H. australis and H. nui sp. n. Heptathela cucphuongensis (known 
only from females), which also occurs in northern Vietnam (south of Hanoi), is 
geographically closer and possibly also more closely related to H. tonkinensis and 
H. tomokunii than to H. australis and H. nui sp. n. from the south of the country. 

DISTRIBUTION: Heptathela tonkinensis is known only from the holotype 
collected in a forest near Khà là (this spelling is given on the laeser-printed label in the 
tube with the spécimen; in the original description the name of the locality is spelled 
"Kha-lé"), in the valley of the Song (= river) Luc Nam, northeast of Hanoi (Simon, 
1909: 69, 71). This locality probably corresponds to the village of Kha Le, about 15 
km east of the township of Luc Nam in Bac Giang Province, northern Vietnam. 



HEPTATHELA FROM VIETNAM 



613 



Heptathela tomokunii Ono, 1997 Figs 39-43 

Heptathela tomokunii Ono, 1997a: 24-26, figs 1-8 (description of maie and female). 

MATERIAL EXAMINED: NSMT-Ar 3396-3398; maie holotype (matured 8.X.1995), 
female "allotype", 1 female paratvpe: Vietnam, Vinh Phu Province, Tarn Dao, 900 m; 
15./21.IX.1995; leg. H. Ono. 

Remarks: Diagnosis and description were given by Ono (1997a). The 
following can be added: cymbial projection only moderately developed, quite wide 
(Fig. 39); paracymbium short and wide at base, at an acute angle to axis of cymbium 
(Ono, 1997a: fig. 2), similar to that of H. tonkinensis; subtegulum exceptionally long 
(Fig. 40); contrategulum with denticles on distal edge modified into a séries of parallel 
ribs running down to 1-2 isolated ventral denticles near proximal margin (Figs 39-40); 
tegulum with quite low, narrowly arched marginal apophysis and with quite short 
terminal apophysis (Figs 39, 41) carrying a pronounced. strongly bulging dorsal 
extension with a coasely dentate edge (Fig. 39; Ono, 1997a: fig. 4) not in contact with 
edge on marginal apophysis; génital atrium with some hairs on ventral side near 
posterior and latéral margins (Fig. 42); bursa copulatrix short, anterior margin widely 
arched; ail réceptacles simple and smooth [Fig. 43; also confirmed for the "allotype", 
though the corresponding illustration (Ono, 1997a: fig. 6) indicates indistinctly cauli- 
flower-shaped réceptacles], not developed as caulifower-shaped receptacular clusters 
as in other Heptathelinae examined; larger latéral pair of réceptacles sessile, slightly 
displaced onto dorsal wall of bursa copulatrix; smaller médian pair of réceptacles 
stalked [on common socket in paratype (Fig. 43), on individual stalks in "allotype" 
(Ono, 1997a: fig. 6)] and slightly displaced onto ventral wall (Fig. 43). 'Tibial spurs" 
présent on legs I-III of female, on last exuvia of maie and also on mature maie (with a 
few "spurs" reduced). 

RELATIONSHIPS: Due to similarities in the palpai organ of maies (moderately 
developed cymbial projection; short, wide paracymbium at an acute angle to the axis 
of the cymbium; long subtegulum; low, arched marginal tegular apophysis; pronoun- 
ced dorsal extension of terminal tegular apophysis), H. tomokunii appears to be more 
closely related to H. tonkinensis (female unknown) than to H. australis and H. nui 
sp. n. More or less pronounced displacements of the latéral pair of réceptacles onto the 
dorsal wall of the bursa copulatrix in H. tomokunii, H. australis and some Ryuthela 
species are considered homoplasic (see Discussion). 

DISTRIBUTION: This species is only known from the type locality. the Tarn Dao 
hill station, about 85 km northwest of Hanoi, in northern Vietnam. 

Heptathela kikuyai Ono, 1998 Figs 44-45 

Heptathela kikuyai Ono, 1998: 16-19, figs 11-16 (description of maie and female). 

MATERIAL EXAMINED: NSMT-Ar 8718; 1 maie; Japan. Kyushu Island, Oita Préfecture, 
Hita-shi; 15.X.2006; leg. T. Irie. - NSMT-Ar 3545; 2 females; Kyushu Island, Miyazaki 
Préfecture, Nishi-ushuki-gun, Takachiho-cho; 8.-14.IX. 1 996; leg. H. Ono. 

Distribution: Known from several localities in the northwestern part of 
Kyushu Island, Japan. 



614 



P. J. SCHWENDINGER & H. ONO 




Figs 39-43 

Hepiathela tomokunii Ono, 1997, maie holotype (39-41) and female paratype (42-43). (39) 
Distal part of left palp, prolateral view. (40) Left palpai organ, ventral view. (41) Marginal and 
terminal apophysis of tegulum of left palp, ventral view. (42) Vulva, dorsal view; dorsal and 
latéral walls of génital atrium not shown. (43) Bursa copulatrix, dorsal view. R= réceptacle; other 
abbreviations as in Figs 1-20. Scale lines 1.0 mm. 



Heptathela kimurai (Kishida, 1920) Figs 46-47 

Liphistius kimurai Kishida, 1920: 362, fig. 1 (description of maie and female); type species of 
Heptathela. For a complète synonymy, see Platnick (2011). 

MATERIAL EXAMINED: NSMT-Ar 3549; 1 maie; Japan, Kyushu Island, Kagoshima-shi; 
7.XI.1970; leg. K. Tanaka. - NSMT-Ar 8717; 1 female; Kyushu Island, Kagoshima-shi, 
Shiroyama; 25.XI.1997; leg. H. Ono. 

Distribution: Known from several localities in the southern part of Kyushu 
Island, Japan. 



HEPTATHELA FROM VIETNAM 



615 




Figs 44-50 

Heptathela kikuyai Ono, 1998, maie (44-45); Heptathela kimurai (Kishida, 1920), maie (46-47); 
Ryuthela owadai Ono, 1997, maie holotype (48-50). (44, 46, 49) Left cymbium, prolateral view. 
(45, 47, 50) Left palpai organ (45 and 47 slightly expanded), ventral view. (48) Same, retrolateral 
view. CS = contrategular spine; other abbreviations as in Figs 1-15. Scale lines 1.0 mm. 



616 



P. J. SCHWENDINGER & H. ONO 



Ryuthela owadai Ono, 1 997 Figs 48-50 

Ryuthela owadai Ono, 1997b: 155-157, figs 15-18 (description of maie).- Ono, 2001: 151-153, 
figs 1-3 (description of female). 

MATERIAL EXAMINED: NSMT-Ar 3459; maie holotype; Japan, Okinawa Préfecture, 
Tokashiki Island, Aharen, about 1 00 m; 1 1 .X. 1 990; leg. M. Owada. - NSMT-Ar 87 1 7; 1 female; 
Okinawa, Tokashiki Island, Tokashiku, 30 m; 17.1.2001; leg. H. Ono. 

Distribution: Known only from the Tokashikijima Island, Ryukyu Islands, 

Japan. 

Ryuthela nishihirai (Haupt, 1979) 

Heptathela nishihirai Haupt, 1979: 356-362, 365, figs 6-13 (description of maie and female); 
type species of Ryuthela. For a complète synonymy, see Platnick (201 1). 

MATERIAL EXAMINED: NSMT-Ar 422-423; 1 maie (matured X.1977) and 1 female 
syntypes; Japan, Shuri, Okinawa, Ryukyu Islands; 15.111.1976; leg. M. Nishihira & J. Haupt. - 
MHNG (coll. S. Heimer), without registration number; 1 female; Japan, Ryukyu Islands, without 
further data. 

Distribution: Known from several localities on Okinawa Island, Japan. 
Liphistiinae Thorell, 1869 

Liphistius ornatus Ono & Schwendinger, 1990 Figs 51-54 

Liphistius ornatus Ono & Schwendinger, 1990: 166-170, figs 1-8 (description of maie and 
female). 

MATERIAL EXAMINED: MHNG, without registration number; maie holotype, 1 maie 
paratype, 4 female paratypes; Thailand, Chanthaburi Province, Khao Soi Dao Wildlife 
Sanctuary, 300-400 m; 9.V.1987; leg. P. Schwendinger. 

Distribution: Known only from the type locality in the southern part of 
eastern Thailand. 

Liphistius thaleri Schwendinger, 2009 Figs 59-64 

Liphistius thaleri Schwendinger, 2009: 1255-1260, figs 1-12 (description of maie and female). 

MATERIAL EXAMINED: MHNG, without registration number; maie holotype, 5 maie 
paratypes, 8 female paratypes; Thailand, Trang Province, Libong Island, 30 m; 20. VII. 2005; leg. 
P. Schwendinger. 

Distribution: Known only from Libong Island in southern Thailand. 
DISCUSSION 

Taxonomic characters and terminology: Différent and partly contra - 
dictory terminology (for a summary see Table 1) for génital structures of liphistiid 
spiders has been used in taxonomic publications by various authors. 

Conductor. Haupt (1979: 358, 360-361, figs 8-10 and subséquent papers of the 
same author) used the term "conductor" for a distad-directed spine-like structure on the 
palpai organ of Ryuthela maies. This spine arises on the ventral side of the contra - 
tegulum and is actually a strongly elongate denticle, the last (proximal-most) one in a 
séries (continuous or interrupted) of denticles that run down from the distal edge of the 
contrategulum in ail heptatheline palps examined (denticles indistinctly developed in 



HEPTATHELA FROM VIETNAM 



617 




Figs 51-54 

Liphistius ornatus Ono & Schwendinger, 1990, maie holotype (51-52) and female paratype 
(53-54). (51) Distal part of left palp, ventral view. (52) Same, retrolateral view. (53) Vulva, 
dorsal view; dorsal and latéral walls of bursa capulatrix and génital atrium not shown. (54) Same, 
ventral view. CDO = central dorsal opening; Co = conductor; CT = contrategulum; Cy = 
cymbium; DEC = distal edge of "contrategulum"; DET = dorsodistal edge of "tegulum"; DPE = 
dentate proximal edge of "contrategulum"; DT = dorsal extension of terminal apohysis of 
tegulum; E = embolus; GA = génital atrium; MA = marginal apophysis of tegulum; P = medium- 
sized pore (leading to ampulliform vesicle); PC = paracymbium; PeP = paraembolic plate; PPI 
= poreplate; RC = receptacular cluster; SE = sharp distal edge of contrategulum; ST = sub- 
tegulum; T = tegulum; TiA = tibial apophsis; V = ampulliform vesicle. Scale lines 1.0 mm. 
Modified from Ono & Schwendinger, 1990: figs 1-4, with permission from the Bulletin of the 
National Science Muséum (Tokyo). 

H. kimurai, Fig. 47; modified to ribs in H. tomokunii, Figs 39-40). In R. owadai the 
three proximal denticles are elongate to various degrees (Figs 48, 50; Ono, 1997b: figs 
16-18). Thus the "conductor" of Ryuthela is not fused with the contrategulum as 
suggested by Haupt (1983: 286; 2003: 71), it is an original part of the contrategulum 
and as such also of the bulbal sclerite II sensu Kraus (1978: 237, figs 2-4). In other 
heptatheline gênera the conductor is a structure of various shapes that arises from the 
ventral base of the embolus (or embolus complex) and thus belongs to bulbal sclerite 
III. Consequently the "conductors" of both groups are not homologous and therefore 
should not both be called the same. In his descriptions of various Ryuthela species Ono 
correctly notes: "contrategulum with a basai spine; conductor not developed" (e.g., 
Ono, 1997b). The elongate basai denticle of the contrategulum, in the following called 
contrategular spine, is unique within the Liphistiidae and a clear synapomorphy for 
species in Ryuthela. Raven (1985: 15, 16) also did not recognize the différent origins 
of the Ryuthela and Heptathela "conductors" but just their différent shapes, and gave 
this as one of the reasons to synonymise Ryuthela with Heptathela {Ryuthela was later 
removed from synonymy by Haupt, 2003: 71). 



618 



P. J. SCHWENDINGER & H. ONO 



The conductor of Heptathela is on the same bulbal sclerite (III) and essentially 
in the same position as the "paraembolic plate" (also called "scale-like plate of the 
ventral embolus edge", "proximal edge of embolus" and "Basalkante des Embolus") 
présent in many species of Liphistius (especially in those of the trang-group; Figs 
51-52). Thèse sclerites differ only in shape and in distance to the embolus proper: in 
Liphistius the paraembolic plate is wide, short or long, fairly remote from the embolus 
and slightly inclined from it, whereas in non-Ryuthela heptathelines the conductor is 
wide or narrow, long, usually lies much closer to the embolus and runs more or less 
parallel to its surface. The relative position of the conductor in H. tonkinensis (type 
species of "NanthelcT; Figs 34, 36) is intermediate between that of Liphistius and 
Heptathela. Différences in shape [paraembolic plate scale-like (Fig. 51) or only deve- 
loped as a low wide edge in Liphistius (Schwendinger, 1998: fig. 1A-D, I); conductor 
leaf-shaped in Heptathela from Japan (Haupt, 2003: fig. 52F-K, M-P, fig. 62G-L), 
proximally broad and usually carrying three more or less pronounced, acutely pointed 
tips in "Sinothela" (Haupt, 2003: fig. 52L, fig. 62D-F), blade-like, spiniform and 
smooth, or spiniform and apically dentate in "Nanthela" (Haupt, 2003: fig. 62C)] do 
not exclude that thèse structures are homologous. This was stated by Raven (1985: 15) 
and vigorously refuted by Haupt (2003: 92); we here support Raven's view. 

The terms "conductor" and "contrategulum" were confused in Ono's (1998) 
taxonomic treatment of the maies of five Heptathela species from Kyushu Island 
(Japan). What is correctly referred to as "conductor" in the descriptive text (except on 
page 21), is incorrectly called "contrategulum" in the legends to the corresponding 
figures. 

Tegulum: Tegulum and contrategulum are the two more or less distinctly 
outlined parts of the ring-shaped bulbal sclerite II of the palpai organ in Liphistiidae. 
In the Heptathelinae the tegulum extends over the dorsal and retrodorsal side of the 
bulb and carries two apophyses: 1) Dorsally lies the distad-directed marginal apophysis 
over which runs a sharp distal edge from the prolateral to the retrolateral side. This apo- 
physis is long and pointed in H. australis and H. nui sp. n. (Figs 2-5, 10, 22-25, 28-29), 
short and arched in the other heptatheline species examined (Figs 34, 37-38, 39, 45, 
47-48). 2) Retrolaterally lies the retrolaterad-directed terminal apophysis over which 
runs a more or less coarsely dentate (least developed in R. owadai) dorsal edge 
perpendicular to the distal edge on the marginal apophysis. The terminal tegular apo- 
physis is distinctly developed in "Nanthela" and Heptathela (Figs 3, 5, 10-14, 23, 25, 
28, 34, 36-39, 41, 45, 47), indistinctly so in Ryuthela (Fig. 48). In H. australis and H. 
nui sp. n. the dorsal side of the terminal tegular apophysis is only moderately extended 
and slightly arched, with the dorsal dentate edge only little elevated (Figs 5, 25), in the 
other Heptathelinae examined it is strongly protruding into a widely arched extension 
(Figs 34, 36, 39, 47-48; not visible in Fig. 45). In H. kikuyai and H. kimurai there is a 
coarsely dentate third edge on the tegulum (= the "first edge" of Song & Haupt, 1984: 
444) which starts at the point where the edges of the marginal and terminal tegular apo- 
physes meet, and from there it runs down the prodorsal side to the base of the tegulum. 
This edge is not produced into an apophysis as on the other two tegular edges. This 
condition appears to be characteristic for H. kimurai (the type species of Heptathela) 
and its closest relatives. In the other Heptathelinae the edges of the marginal and 
terminal apophyses do not meet prodorsally and no third edge is présent. 



HEPTATHELA FROM VIF.TNAM 



619 



In Liphistius the tegulum is less obvious, because it seemingly does not possess 
the same apophyses as the tegulum of the Heptathelinae. The so-called "tegular 
process" of some Liphistius species arises from the opposite (= ventral to proventral) 
side of bulbal sclerite II and thus belongs to the contrategulum. The contrategular 
process is probably apomorphic for Liphistius, as is also the subtegular apophysis (both 
présent only in some species); both apophyses are absent in ail Heptathelinae. What 
was previously called "tegulum" in Liphistius therefore does not correspond to the te- 
gulum in the Heptathelinae. There is, however, a sclerotised area on the retrodorsal side 
of bulbal sclerite II in Liphistius that is connected to the larger part (= the contra- 
tegulum) of bulbal sclerite II by a narrow sclerotised area and that carries a sharp distal 
edge in most species and a dentate marginal or submarginal proximal edge in ail 
species (Fig. 52, Schwendinger, 1996: figs 1, 12, 21, 31, 39, 46, 55, 65, 74, cf. Figs 4, 
24). Thèse two edges, though being more two-dimensional, resemble the marginal and 
terminal tegular apophyses of Heptathelinae and they lie at the same position on the 
palpai organ. Haupt (1983: figs le, 3-4) called the sclerite of the Liphistius palp which 
carries thèse two edges the "contrategulum", and that appellation was followed in pa- 
pers on Liphistius by Ono and by Schwendinger. We assume that Haupt regarded this 
sclerite as the "contrategulum" because of a misinterpretation of his own illustration 
(see Haupt, 1983: fig. le) that shows a distinctly expanded maie palp of L. malayanus 
Abraham, 1923 in which the so-called "contrategulum (K)" was rotated to the ventral 
side where the contrategulum is situated in the Heptathelinae (see Haupt, 1983: 
fig. lb). In his illustrations of non-expanded Liphistius palps (see Haupt, 1983: figs 
3b-c, 4a-b), however, the so-called "contrategulum (K)" lies on the other side, retro- 
dorsally, where the tegulum of the Heptathelinae is situated. Correspondingly the latter 
figures show the so-called "tegulum (T)" proventrally, where the Heptathelinae have 
the contrategulum. Haupt has obviously confused thèse terms in both taxa. The 
tegulum of Liphistius (hitherto incorrectly called "contrategulum") is much shorter and 
narrower than that of the Heptathelinae, and it is confined to the retrodorsal side of 
bulbal sclerite II. The dentate proximal or subproximal edge of the tegulum of 
Liphistius is presumably homologous with the dentale edge on the dorsal side (pro- 
truded into a dorsal extension in some species) of the terminal tegular apophysis of the 
Heptathelinae, and the more or less distinctly developed distal edge of the Liphistius 
tegulum is presumably homologous with the marginal apophysis of the Heptathelinae 
(Fig. 52 cf. Figs 4, 24). A fairly simple tegulum (more reduced than in Heptathela but 
less than in Liphistius) is also présent in Ryuthela owadai (Fig. 48). We consider the 
small tegula of Liphistius and Ryuthela as homoplasic réductions, which implies that 
the palpai organs of Liphistius and (to a lesser degree) Ryuthela are more derived than 
those of Heptathela. 

Contrategulum: Haupt (1983: 277) introduced the term "contrategulum" for the 
prolateral to ventral part of bulbal sclerite II that opposes the tegulum. In the 
Heptathelinae tegulum and contrategulum are more or less distinctly separated by a 
wide membranous zone (haematodocha) retroventrally, thus forming a broken ring. 
Prodorsally, however, both parts are widely fused and it is difficult to tell where the 
contrategulum ends and the tegulum begins. An indicator is the sharp distal edge of the 
contrategulum portion which more or less abruptly ends prolaterally in H. australis 



620 



P. J. SCHWENDINGER & H. ONO 



(Fig. 2), H. nui sp. n. (Fig. 22), H. kikuyai and H. kimurai, wereas in H. tonkinensis 
(Figs 34, 36), H. tomokunii (Fig. 39) and R. owadai (Fig. 48) it fades out between the 
marginal tegular apophysis and the embolus. More proximally at that place contrate- 
gulum and tegulum are usually fused with each other without a transition, though in 
Hepthatela maies from Japan examined a small invagination in the proximal margin of 
bulbal sclerite II presumably marks the border between both parts. In Ryuthela this pat- 
tern is modified and the distinction between contrategulum and tegulum is more visible 
on the dorsal side of the palpai organ: the contrategulum is much wider than in other 
heptathelines (as wide as in Liphistius) and extends distally past the tegulum to the 
retrolateral side in an almost complète circle; a deep, almost horizontal furrow divides 
both sclerites dorsally (Fig. 48), and only on the ventral side are tegulum and contra- 
tegulum completely fused (Fig. 50). In R. nishihirai tegulum and contrategulum appear 
to be completely separated (see Haupt, 1979: 360, fig. 9). In Liphistius the contra- 
tegulum is much more extensive than the tegulum, almost completely embracing the 
middle portion of the palpai organ (as also in Ryuthela, but there the tegulum is much 
wider than in Liphistius and overlaps the contrategulum). Retrolaterally the large 
contrategulum and the much smaller tegulum are connected by sclerotised bridges (the 
more dorsal one always narrower than the more ventral one) and form a closed ring 
(Fig. 52). What was previously called "tegular process", "distal edge of tegulum" and 
"dorsal edge of tegulum" in various publications on Liphistius by Haupt and by 
Schwendinger, are in fact parts of the contrategulum. 

In Heptathelinae the contrategulum usually carries on its distal edge (which is 
divided into a sharp outer edge and a dentate inner edge in H. australis and H. nui 
sp. n.) a row of denticles which are more or less distinctly connected to a few (in some 
species isolated) denticles ventro-proximally. Thèse denticles were subject to evolu - 
tionary modifications of taxonomic significance. In several species they have 
multiplied and become more than one row deep: in H. nui sp. n. two rows proximally 
(Fig. 23), in R. owadai three rows proximally (Fig. 50), in H. kikuyai several rows 
proximally (Fig. 45) and in H. tonkinensis several irregular rows on most of its distal 
edge (Fig. 36); in H. tomokunii (Figs 39-40) and seemingly also in H. hongkong Song 
& Wu, 1997 (Haupt, 2003: fig. 51D) they have transformed into a séries of parallel 
edges or ribs. Such ribs are also présent in some Liphistius species where they are 
probably homoplasic (see Schwendinger, 1995: figs 22-23, 26-27; Schwendinger, 
1996: figs 12, 15-17; Schwendinger, 1998; fig. 5b, e-f). In Ryuthela spp. the basalmost 
of thèse denticles (in R. owadai to a lesser degree also two adjacent ones; Figs 48, 50) 
has become strongly elongate and spine-like. This. modified contrategular denticle of 
Ryuthela is clearly not homologous to the conductor (or paraembolic plate) of other 
Liphistiidae (see discussion of conductor), but it présents a clear synapomorphy for 
species in Ryuthela. 

Cymbial projection: Haupt (2003: 69, 94, character 23 in table 11 and fig. 61) 
considers the elongate pro ventral cymbial lobe as "autapomorphic" (Haupt, 2003: 94; 
it should read "synapomorphic") and distinctive for maies of species in the genus 
"Nanthela ". However, this modification is also présent in two Heptathela species from 
Japan examined: H. kikuyai (Fig. 44) and H. kimurai (Fig. 46; the type species), and it 
is also discernible in Song & Haupt's (1984: fig. le) illustration of the maie palp of 



HEPTATUELA KROM VIHTNAM 



621 



H. schensiensis (previously in "Songthela") and in Haupfs (1983: fig. lOa-d) illus- 
trations of the maie palp of Ryuthela nishihirai and R. ishigakiensis Haupt, 1983. In 
R. owadai the cymbial projection is only weakly developed (Fig. 49). In some 
Liphistius species (e.g., L. bristowei Platnick & Sedgwick, 1984 and L. batuensis 
Abraham, 1923; in MHNG examined), on the other hand, it is the prodorsal cymbial 
lobe (instead of the proventral one) that is moderately elongate. Thus it appears that 
one of the cymbial lobes (the proventral one in Heptathelinae, the prodorsal one in 
Liphistius) has become elongated to various degrees independently in différent phylo- 
genetic lineages. The taxonomic value of this character at the generic level has thus 
been overestimated. 

Vulva: HEPTATHELINAE. The vulva of heptatheline spiders is composed of 
two main parts (see Fig. 19): 1) The posterior part is a long and wide entrance area 
(= the génital atrium) furnished with numerous hairs on the dorsal and ventral walls in 
H. australis (Fig. 16), H. nui sp. n. (Fig. 31) and H. tomokunii (Fig. 42), but only on 
the dorsal wall in H. kikuyai, H. kimurai, R. owadai and R. nishihirai. The génital 
atrium is overlapped by a more or less tongue-shaped projection of the posteromedian 
margin of the génital sternite. 2) The anterior part of the vulva is a blind-ending pouch 
(= the bursa copulatrix) in the form of a flattened bell (see Ono, 2002a: fig. 3), plano- 
convex or trapezoidal in dorsal and ventral view, with walls consisting of a light 
yellow, tough, leathery but not sclerotised cuticle (Figs 16-19, 31-33, 42-43, 55). As 
already stated by Forster (1980: 280), there is no sclerotised structure in the vulva of 
heptatheline spiders that can be called a génital plate (an incorrect term used by several 
authors; see Table 1]. Usually no pores are présent in the walls of the bursa copulatrix 
(Figs 55-56), but the largest H. australis female has tiny pores on the small vesicles 
that lie scattered between the receptacular clusters (Fig. 16). This could be an atavism 
(see Evolution of female copulatory organs). From the dorsal side of the vulva, where 
both main parts meet at an obtuse angle, a short and wide, dorsoventrally depressed 
membranous collar (= the utérus externus) leads dorsad to the mesodermal part of the 
oviduct (= the utérus internus) and further to the ovary (Fig. 19). In the illustrations by 
Haupt (1983: fig. 2b) the entrance to the utérus externus is erroneously shown at the 
anterior end of the bursa copulatrix, which is in fact a cul-de-sac. The anterior margin 
of the bursa copulatrix carries between one and four sessile or stalked, mostly cauli- 
flower-shaped protubérances (= the receptacular clusters) composed of numerous more 
or less strongly sclerotised vesicles (= the réceptacles). The composite structure of the 
receptacular clusters in heptathelines (for Ryuthela nishihirai) was first recognized by 
Haupt (1979: 365). The walls of thèse vesicles are perforated by tiny micropores 
through which surrounding glands empty. The proximal, cuticular portions of the gland 
ducts that are sticking out of the micropores like fine hairs (Figs 56-58; SEM-micro- 
graphs taken from exuviae) clearly show that thèse are indeed gland pores. In the two 
females of H. tomokunii examined thèse anterior protubérances on the bursa copulatrix 
are simple in shape and have a fairly smooth surface (though also perforated by micro- 
pores; Figs 42-43) and may be called réceptacles. This indicates a trend towards 
simplification, which is also seen in the receptacular clusters of some Liphistius species 
that have become smooth and digitiform (Sedgwick & Schwendinger, 1990; fig. 6; 
Schwendinger, 1996: figs 18-20, 52-54). In H. australis and other species previously 



622 



P. J. SCHWENDINGER & H. ONO 




Figs 55-58 

Heptathela australis (Ono, 2002), SEM-micrographs of genitalia from the exuvia of one female, 
dorsal views. (55) Entire bursa copulatrix with receptacular clusters. (56) Border between dorsal 
wall of bursa copulatrix (without pores) and lower left receptacular cluster (with gland pores). 
(57) Lower left receptacular cluster. (58) Détail of the same showing gland pores and cuticular 
parts of gland ducts. BC = bursa copulatrix; GD = cuticular (proximal) part of gland duct; GP = 
gland pore (micropore); RC = receptacular cluster. Scale Unes: 14 um (58), 31 um (56), 56 um 
(57), 236 um (55). 

placed in "Songthela " by Ono (2000), two of thèse receptacular clusters (the latéral 
ones) have become displaced onto the dorsal wall of the bursa copulatrix and are 
directed dorsad instead of anteriad (Figs 16-19, 55). 

Illustrations in the original description of H. goulouensis Yin, 2001 show a very 
différent type of vulva (without a bursa copulatrix but instead with a pair of bifid 
"curved, tube-like stalks", each carrying a médian and a latéral réceptacle or recep- 
tacular cluster; Yin, 2001: figs 5-6, 8-9) which is similar to the vulvae of certain 
mygalomorph spiders. If correct, this would be most unusual for liphistiid spiders. A 



HEPTATHELA FROM VIETNAM 



623 



re-examination of thèse spécimens may show that the true boundaries of the bursa 
copulatrix were not recognized and that the "stalks" are actually zones of différent pig- 
mentation, or a mass of sperm inside the bursa copulatrix. Similar "stalks" and a short 
bursa copulatrix were illustrated for the holotype of H. sapana (Ono, 2010: fig. 16). 
The so called "didymous phenomenon" displayed by the female paratype of H. gou- 
louensis (Yin, 2001: figs 8-9) is most likely based on a spider that was preserved prior 
to moulting, which has the new cuticle of the vulva already formed underneath the old 
cuticle. 

The number of réceptacles or receptacular clusters and their position on the 
bursa copulatrix is variable and this has been used to recognize groups of species which 
were given generic rank. Ryuthela is characterized by only two anterior receptacular 
clusters situated close to each other (Haupt, 2003: 71, fig. 53), but a rudimentary pair 
of additional (latéral) receptacular clusters is still présent in the female "allotype" of 
R. owadai (see Ono, 2001: figs 2-3). This is not so in the corresponding female para- 
type examined. Non-Ryuthela Heptathelinae usually have four more or less distinctly 
developed réceptacles or receptacular clusters, but distinction from Ryuthela is not 
clear-cut. Latéral receptacular clusters have become more or less fused with médian 
ones in several Heptathela species (see Ono, 1998: figs 9-10, 15-16, 17-20, 31-32; 
Haupt, 2003: fig. 54), resulting in only two anteromedian receptacular clusters which 
are generally more widely separated from each other than in Ryuthela. The holotypes 
of H. hunanensis Song & Haupt, 1984 and of H. cucphuongensis have the médian pair 
fused and the latéral pair still isolated, resulting in a vulva with three anteromedian 
receptacular clusters (Song & Haupt, 1984: fig. 3e; Ono, 1999: fig. 8). Ono (2000) used 
this intermediate character state to establish the genus "Vinathela" (misspelled 
"Viathela " and placed in the synonymy of Heptathela by Haupt, 2003: 71, 79). Fusion 
of receptacular clusters has gone even further: the illustrated vulva of R. iheyana Ono, 

2002 has a single, wide and short anteromedian receptacular cluster (Ono, 2002b: figs 
2-3); the holotype and one female paratype of R. sasakii Ono, 1997 have a single, long 
and narrow anteromedian receptacular custer (Ono, 1997b: figs 5-8). We consider ail 
thèse forms as intermediate stages of réduction and fusion of the four anterior recepta- 
cular clusters that are plesiomorphically présent in heptatheline spiders. Such modi - 
fications probably have evolved independently in différent phylogenetic lineages and 
should be treated with caution when establishing relationships between species. 

Two receptacular clusters displaced to the dorsal side of the bursa copulatrix 
(Figs 16-19, 55) is another kind of modification in heptatheline vulvae. This character 
state was used by Ono (2000) to establish the genus "Songthela", but as H. australis 
(described in "Songthela ") has it and the obviously closely related H. nui sp. n. not, it 
can no longer be used as distinctive on the generic level. Ail the more so because se- 
veral transitional steps exist: the latéral receptacular clusters of H. tomokunii (see Figs 
42-43; Ono, 1997a: fig. 6), H. sinensis, H. schensiensis (see Song & Haupt, 1984: fig. 
3a-b, f) [the latter two placed in "Abcathela " by Ono, 2000, in "Sinothela " by Haupt, 

2003 and then in "Songthela" by Platnick, in an earlier version of his 2011 online 
catalogue] and of H. yunnanensis Song & Haupt, 1984 (see Song & Haupt, 1984: 
fig. 3f) are also slightly displaced onto the dorsal wall of the bursa copulatrix, but not 
as far back as in H. australis (Figs 16-18, 55) and as in H. hangzhouensis (see Song & 



624 



P. J. SCHWENDINGER & H. ONO 



Haupt, 1984: fig. 3c-d) (both species previously in "Songthela"). The médian pair of 
réceptacles (reduced receptacular clusters) of H. tomokunii (Figs 42-43; in addition to 
the latéral pair which was slightly displaced onto the dorsal wall) and of some Ryuthela 
species (see e.g., Ono, 1997b: figs 5-14, 19-22) is even sitting slightly below the 
anterior margin of the bursa copulatrix, on its ventral wall (thus on the same side as the 
receptacular cluster of Liphistius). 

LIPHISTIIDAE. The heptatheline vulva is largely différent from the vulva of 
Liphistius which carries no anterior receptacular clusters. In that genus the dorsal wall 
of the bursa copulatrix is a thin transparent membrane that never carries any modifi- 
cations apart from a pigmented spot opposing the macropore in the ventral wall in a 
few large females. (In Liphistius the entrance to the utérus externus is also situated at 
the border between atrium and bursa copulatrix, not at the anterior end of the bursa 
copulatrix as shown in Haupt, 1983: fig. 2a). The ventral wall of the bursa copulatrix 
in Liphistius is strongly sclerotised and includes a true génital plate (= the poreplate) 
perforated dorsally by medium-sized pores that lead inside small ampulliform vesicles 
on the ventral side (Figs 53-54, 59-60). The appellation "gland pores" used in a paper 
by Schwendinger (1996) for thèse vesicles is incorrect; the gland pores (= micropores) 
are actually on the ouside of the vesicles (ventrally on the poreplate), as clearly visible 
in SEM-micrographs by Kraus (1978: fig. 8; see also Fig. 62). Kraus (1978: 242-243, 
figs 6-8) called thèse vesicles "circular structures [or circular areas] . . . interpreted . . . 
as a pore system for glands . . . [which] must not be confused with the receptacula". 
Kraus (1978: fig. 8) first showed SEM-micrographs of micropores in Liphistius and 
recognized the function of the ampulliform vesicles, but the corresponding lettering is 
misleading [in fig. 8 the ventral surface of a vesicle is labelled as "gland pore", in 
fig. 6 the opening (= medium-sized pore) of a vesicle on the dorsal side of the pore- 
plate]. Sperm that enters into the ampulliform vesicles from the dorsal side of the 
poreplate is thus supplied with (or flushed out by) a sécrétion of the receptacular glands 
from the other side. Thus the vesicles on the ventral side of the Liphistius poreplate are 
sperm réceptacles. 

On the dorsal side of the Liphistius poreplate a fairly large central opening (or 
macropore; CDO in Figs 53, 59) leads to an unpaired, strongly sclerotised réceptacle 
or (in most species) receptacular cluster (Figs 54, 60-61; Haupt, 2003: 68 called this an 
"unpaired ventral sac") on the ventral side of the poreplate. The central receptacular 
cluster is perforated by gland pores, and it varies in shape from simple and digitiform 
(in that case called a réceptacle, see Sedgwick & Schwendinger, 1990; fig. 6; 
Schwendinger, 1996: figs 18-20, 52-54) to complex and cauliflower- or popcorn- 
shaped (Kraus, 1978: figs 5-8; Platnick & Sedgwick, 1984: figs 69, 80, 87). In the 
heptatheline vulva no réceptacle or receptacular cluster is found at this position 
(although shown there in Haupt, 1983: fig. 2b), but a slight displacement of the médian 
pair onto the ventral side of the bursal copulatrix can be seen in H. tomokunii (Figs 



Figs 59-64 

Liphistius thaleri Schwendinger, 2009, SEM-micrographs of vulvae from the exuviae of three 
female paratypes. (59) Ventral wall of opened bursa copulatrix and génital atrium, dorsal view. 
(60) Same. ventral view. (61) Receptacular cluster. (62) Ampulliform vesicle. (63) Anterolateral 



iii.r I 1 TU I I I 1 ROM \ II Ï YW1 



625 




protubérance on rim of poreplate. (64) Posterolateral protubérance on rim of poreplate. ALP = 
anterolateral protubérance on rim of poreplate; AMP = anteromedian protubérance on rim of 
poreplate; CDO = central dorsal opening (macropore) to receptacular cluster on other side; GA 
= génital atrium; GD = cuticular (proximal) part of gland duct; GP = gland pore (micropore); P 
= medium-sized pore (leading to ampulliform vesicle); PLP = posterolateral protubérance on rim 
of poreplate; PP1 = poreplate; RC = receptacular cluster; V = ampulliform vesicle. Scale lines: 
14 um (64), 16 um (62), 32 um (63), 35 um (61), 195 um (59), 205 um (60). 



626 



P. J. SCHWENDINGER & H. ONO 



42-43) and in several Ryuthela species (Ono, 1997b: figs 9-14, 19-22). In R. sasakii 
this pair has become fused. it possesses a common unpaired, slightly ventrally situa- 
ted, fairly large opening (macropore; see Ono, 1997b: figs 5, 7), and thus resembles the 
receptacular cluster in Liphistius. 

Several Liphistius species possess 1-3 pairs of ventrad-directed protubérances 
on the thickened anterior and latéral margins of the poreplate (Fig. 60). Thèse appear 
to be synapomorphic for members of superspecies D of the /ra«g-group (co-occurring 
with a synapomorphic subtegular apophysis in maies of thèse species, see 
Schwendinger. 2009; subtegular apophyses in the biistowei-group are homoplasic), but 
also occur in distantly related species in other species groups (e.g., Platnick & 
Sedgwick. 1984: figs 13, 21, 78; Schwendinger, 1990: figs 2-4, 37-39, 43-45, 47-49, 
53-56). Thèse protubérances are hollow, in contact with the bursa copulatrix through 
medium-sized pores on the dorsal side of the poreplate, and their walls are perforated 
with micropores through which gland ducts enter (Figs 60, 63-64). Therefore the ven- 
tral marginal protubérances are functional sperm réceptacles. Function and position of 
thèse protubérances suggest that they are homologous with the receptacular clusters of 
the heptatheline vulva, but considering that the poreplate in Liphistius itself is an apo- 
mophic structure, that is quite unlikely. We regard thèse protubérances as outgrowths 
on the thickened margin of the poreplate, which have developed several times inde- 
pendently in Liphistius. They are useful for species distinction, and their number and 
position on the poreplate allow recognizing relationships between species. 

Micropores are not only found in the elevated parts of the ventral side of the 
poreplate, but also in the flat parts between them (Figs 60, 62). Most of the pores there 
are grouped and sitting in shallow dépressions, with membranous gland duct bases 
sticking out of them, and thèse groups of sessile micropores are in contact with 
medium-sized pores on the dorsal side of the poreplate. Thus essentially the whole of 
the poreplate serves as a sperm réceptacle. 

Evolution of female copulatory organs: The fact that two very différent types of 
vulvae exist within the Liphistiidae raises the questions: how did they evolve and 
which one is more derived than the other? Five hypothèses were put forward in an 
attempt to explain the évolution of female copulatory organs in Liphistiidae and in 
Araneae as a whole: 

1) Platnick & Gertsch (1976: 6), and subsequently Platnick (1977: 13), pos- 
tulated that in ancestral spiders the palps were pressed simultaneously into the female 
génital tract during copulation, and that the tips of embolus and conductor of each 
palpai organ left imprints in the wall of the vulva which developed into réceptacles. A 
vulva with four réceptacles, as found in most Heptathelinae and in some basai 
Mygalomorphae and basai Araneomorphae, is thus plesiomorphic for the Araneae. 

2) An alternative hypothesis, put forward by Forster (1980: 277, fig. 12) and 
further elaborated by Forster, Platnick & Gray (1987: 94-98), présents an archetypical 
spider vulva with only a large and simple bursa copulatrix supported by a secretory 
gland System that empties through pores in the anterior wall of the bursa. According to 
this hypothesis. numerous réceptacles (vesicles) developed later by invagination 
(better called exsacculation) of the bursal wall where the gland pores are situated. In 
this view the Liphistius vulva is relatively primitive because it still has numerous indi- 



HEPTATHELA F ROM VIETNAM 



627 



Table 1. Différent terminology used for homologous génital characters in three groups of 
liphistiid spiders by various authors in 14 selected publications. Forster (1980 1 ); Haupt (1979 2 , 
1983 3 , 2003 4 ); Kraus (1978 5 ); Ono (1997b 6 ); Ono (1998 7 ); Ono & Nishikawa (1989 8 ); 
Schvvendinger (1990 9 , 1996 10 , 2009 11 ), Song & Haupt ( 1984 12 ), Yin (2001 ,3 ), présent paper 14 . 
Abbreviations (in parenthèses) correspond to those given in Figs 1-64. Terms between inverted 
commas are considered as inappropriate or incorrect. 



Heptathela (see Figs 1-47, 
55-58, 67-68) 



Ryuthela (see Figs 48-50) 



Liphistius (see Figs 51-54, 
59-64) 



conductor (Co) 2 ' 3 > 4 < 7 > 12 < 14 , 
"contrategulum" (CT)7 



ventro-proximal denticles on 
contrategulum 14 



contrategulum (CT) 3 - 4 - 7 - 8 - 
12, 14 



tegulum (T) 



3,4, 7, 8, 12, 14 



terminal apophysis of 
tegulum 12 ; third tegular 
edge 12 ; dentate edge of 
dorsal extension of terminal 
apophysis of tegulum (DT) 14 

marginal apophysis of 
tegulum (MA) 12 ' 14 ; second 
tegular edge 12 



génital plate 4 - 12 ' 13 ; Génital- 
platte 3 ; dorsal and ventral 
walls of bursa copulatrix 14 

receptacula 1 - 3 - 4 - 12 - 13 ; 
réceptacles (R) 14 ; 
receptacula organ 13 ; 
receptacular clusters (RC) 14 ; 
spermathecae 7 ' 8 



"conductor" 2 ' 3 ' 4 ; spine on 
contrategulum 6 ; 
contrategular spine (CS) 14 

Medianapophyse 2 ; 
contrategulum (CT) 6 - 14 

tegulum (T) 2 - 3 - 4 - 6 - 14 

dentate edge of dorsal 
extension of terminal 
apophysis of tegulum (DT) 14 



marginal apophysis of 
tegulum (MA) 14 



Genitalplatte 3 ; Atrialplatte 3 ; 
dorsal and ventral walls of 
bursa copulatrix 14 

receptacula 2 - 3 - 4 ; 
receptacular clusters (RC) 14 ; 
spermathecae 6 



para-embolic plate (PP) 10 - 11 - 14 ; 
"posterior" edge of embolus 9 ; 
scale-like proximal embolus 
edge 9 ; broad lamella at base of 
embolus 4 - 5 ; Chitinplatte auf 
Basalkante des Embolus 3 ; 
conductor 14 



"tegulum" (T) 3 - 4 - 9,10,11. 
contrategulum (CT) 14 

"contrategulum" (CT) 3 - 4 - 10 - 
tegulum (T) 14 

dentate proximal edge of 
"contrategulum" (DPE) 10 ; 
"ventral" edge of "contra- 
tegulum" 11 ; dentate edge of 
dorsal extension of terminal 
apophysis of tegulum (DT) 14 

distal edge of "contra- 
tegulum" (DEC) 10 ; "dorsal" 
edge of "contrategulum" 1 1 ; 
distal edge of tegulum 14 ; 
marginal apophysis of 
tegulum (MA) 14 

Genitalplatte 3 ; Atrialplatte 3 ; 
receptaculum 4 ; chitinous 
plate 5 ; inner plate 5 ; internai 
plate 5 ; poreplate (PP1) 1 - 14 

receptaculum 9 - 10 ' 14 ; 
receptacular system 5 ; central 
receptacular cluster (RC) 9 - 10 - 
14 ; pseudo-receptaculum 1 ; 
unpaired ventral sac 4 



vidual réceptacles [plus an autapomorphic unpaired "pseudo-receptaculum" (= the 
central receptacular cluster)] embedded in the poreplate, whereas the heptatheline 
vulva has developed further away from the ancestral form through a bilatéral aggrega - 
tion of réceptacles into receptacular clusters and through a réduction of the poreplate 
(Forster, Platnick & Gray, 1987: 98). 



628 



P. J. SCHWENDINGER & H. ONO 



3) A similar interprétation was given by Kraus (1978: 235, 249), who considers 
the Liphistius-type vulva (with an unpaired central receptaclular cluster) as the most 
primitive within the Araneae and postulated that vulvae with paired réceptacles (and 
also those with unpaired ones) derived from it. 

4) Haupt took éléments of the previous hypothèses to explain the évolution of 
female copulatory organs in spiders. He postulated an ancestral vulva with a large 
bursa copulatrix and irregularly arranged gland pores (Haupt, 1983: 288), as did 
Forster (1980), but referred to Kraus (1978) who proposed a différent hypothesis (see 
above). Localised réceptacles were assume to have formed later in co-evolution with 
the split tips of the embolus (Haupt, 1983: 288). In his cladogram of mesothelid syste- 
matics and in the corresponding character évaluation Haupt (1990: 136, fig. 1) follows 
Kraus (1978) and regards the vulva of Liphistius as plesiomorphic and that of 
Heptathelinae as apomorphic. Later, however, he noted that "it cannot be decided with 
certainty whether unpaired or paired receptacula in mesothelid spiders have to be 
considered as apomorphic for the group" (Haupt, 2003: 93). 

5) Raven (1985: 15-16) regarded the vulva of Ryuthela (with a single pair of 
anteromedian réceptacles) as plesiomorphic and assumed that the vulvae of Heptathela 
and Liphistius derived from it. 

In view of the evolutionary tendencies for fusion and posteriad-displacement (to 
the dorsal and ventral walls) of four anterior réceptacles or receptacular clusters in 
différent lineages of heptatheline spiders, we believe that the same has also happened 
during the évolution of Liphistius species. We thus favour a combination of hypothèses 
1 and 2 and consider the Liphistius vulva as derived from a proto-Heptathela vulva. 
That presumably resembled the vulvae of H. nui sp. n. (Figs 3 1-33) and of species from 
China and Vietnam previously placed in the genus "Abcathela" by Ono (2000), but 
had a much larger bursa copulatrix carrying four receptacular clusters on its anterior 
margin and possibly also scattered micro-receptacles on the walls of the bursa copu- 
latrix more posteriorly. Such isolated micro-receptacles (vesicles penetrated by gland 
pores) are still présent in the anterior part of the dorsal surface of the bursa copulatrix 
of the largest H. australis female examined (Fig. 16). This possibly is an atavism. 

The pronounced différences in the vulvae of Liphistius and Heptathela indicate 
that the séparation of both gênera goes back much further in time than the séparation 
of Heptathela and Ryuthela. The vulva of the latter is essentially a Heptathela vulva 
with a strongly reduced bursa copulatrix, with the receptacular clusters of each side 
fused to one another resulting in a single pair. 

This interprétation is in line with the hypothesis of Schwendinger (2009: 1265- 
1266) that the Mesothelae originated in Euramerica before its intégration into Pangaea, 
and from Euramerica spread eastward onto terranes that successively accreted to 
eastern Laurasia. In such a scénario, and knowing the very limited powers of dispersai 
of extant mesothelid spiders, one would rather expect to find the more basai extant 
species in southern and eastern China than further south, in Myanmar, Thailand, 
peninsular Malaysia and Sumatra. The latter lands mostly lie on the Sibumasu terrane 
which accreted to Laurasia later than the Chinese terranes and thus was colonized by 
Laurasian species later. This zoogeographic hypothesis and its implications for liphis- 
tiid phylogeny will be falsified as soon as an early (Palaeozoic or Mesozoic) meso- 
thelid fossil is discovered on land of Gondwanan origin. 



HEPTATIII I ! I ROM VIETNAM 



629 



"Tibial spurs": The function of thèse paired structures distally on leg tibiae 
I-III (Platnick & Goloboff, 1985: figs 1-2; Haupt, 2003: fig. 16B-D), which must not 
be confused with coupling spurs ventrally or prolaterally on tibia I or tibia II in maies 
of many mygalomorph spiders, is not yet understood. Platnick & Goloboff (1985) 
consider them as part of proprioreceptive sensé organs in which the "tibial spurs" press 
against smooth oval patches at the bases of metatarsi I — 1 1 1 (Platnick & Goloboff, 1985: 
figs 3-4; Haupt, 2003: fig. 16B, D). Haupt (2003: 22, 95, fig. 16), however, assumes 
that the "tibial spurs", situated close to lyriform organs, are more likely used to monitor 
hemolymph pressure and strains in the cuticle, because they are allegedly not in contact 
with the metatarsal patches ("the bristle has no contact to the circular area of thinner 
cuticle"). This statement is incorrect. On alcohol-preserved spécimens with stretched 
legs, or on a dry, stub-mounted and sputter-coated leg as shown by Haupt (2003: 16D), 
thèse structures are usually not in contact with each other, but they are so on live 
spiders sitting in ambush with slightly bent legs at the burrow entrance. Whatever the 
function of this sensory System is, its phylogenetic significance as a clear apomorphy 
of the Liphistiidae is undisputed. We found it to be présent in ail juvénile (also exuvia 
of penultimate maies) and female liphistiids ever examined by both of us. "Tibial 
spurs" (but not always the light smooth paired patches on metatarsi I-III) are usually 
absent in mature maies. In the maie holotype of H. tomokunii and in the maie paratype 
of L. ornatus (Ono & Schwendinger, 1990: 169-170) some "tibial spurs" are présent 
but probably no longer functional. 

Systematics: Discoveries of new species or of the missing sex of described 
species occasionally challenge the concepts of established supraspecific taxa. This is 
also the case in the two Heptathela species from southern Vietnam. The female of H. 
australis corresponds perfectly to Ono's (2000: 150, fig. 4D) concept of "Songthela", 
where this species was originally placed. Its newly discovered maie, however, 
possesses the two characters considered by Haupt (2003: 69) as diagnostic for 
"Nanthela " (a cymbial projection présent, conductor slender and much of its base 
fused to the embolus), and it also has a character considered in the same paper, on the 
same page, as diagnostic for Heptathela [paracymbium almost as long as cymbium 
(though relatively shorter in relation to the cymbium of Heptathela from Japan because 
that usually has no pronounced cymbial projection)]. The same type of maie palp is 
présent in the closely related H. nui sp. n., the female of which, however, possesses a 
vulva as in species previously placed in "Abcathela" (see Ono, 2000: 149-150, 
fig. 4C). This suggests that three of thèse four nominal gênera ("Abcathela", 
Heptathela, "Nanthela ", "Songthela ") do not reflect phylogenetic relationships. We 
rule out that H australis and H. nui sp. n. are only distantly related. Geographical 
proximity and strong overall resemblance of the maie palps show that thèse species are 
the most closely related. They share four likely synapomorphies in their maie palps: 1) 
a very pronounced cymbial projection (more than in other heptathelines); 2) a long and 
pointed marginal tegular apophysis; 3) two parallel distal edges (one sharp, the other 
dentate) on the contrategulum; 4) an unpigmented and unsclerotised distoventral zone 
on the cymbium. Consequently thèse two species cannot be kept in différent gênera, 
and at the moment it appears most suitable to place them in the genus Heptathela. 

The generic concept of "Songthela" is firmly rejected because the species 
included clearly do not represent a monophyletic group. Its single diagnostic character 



630 



P. J. SCHWENDINGER & H. ONO 



(the dorsally displaced latéral receptacular clusters on the bursa copulatrix) is also 
présent in species outside the genus, but it is not shared by the closely related species 
pair "Songthela " australis and Heptathela nui sp. n. The original concept of this genus 
does anyway no longer correspond to the currently included species apart from the type 
species, S. hangzhouensis (see Platnick, 2011). "Songthela" australis and "Song- 
thela "cipingensis Wang, 1989 were transferred to Heptathela in an earlier version of 
Platnick's (201 1) online catalogue, without giving an explanation. Three other species 
[H. heyangensis (Zhu & Wang, 1984), H. schensiensis and H. sinensis, ail with a 
différent type of vulva than in the type species] were included in "Songthela " by the 
synonymisation of "Abcathela" with "Sinothela" and then by the synonymisation of 
"Sinothela " with "Songthela ". 

The generic concept of "Nanthela" is also rejected because it is not based on 
any clear apomorphies. One of the two diagnostic characters, the cymbial projection is 
also présent in species outside "Nanthela ", e.g., in H. kikuyai (Fig. 44), H. kimurai 
(Fig. 46) and "Sinothela" schensiensis (see Song & Haupt, 1984: fig. lc-d). The other 
diagnostic character of "Nanthela ", the slender conductor, is possibly plesiomorphic 
(Haupt, 1990: 137, fig. 1; Haupt, 2003: fig. 61, table 11). Instead of retaining this 
poorly defined genus and transferring species from the well-established genus 
Heptathela to it {"Songthela" australis, H. nui sp. n., "Nanthela" tonkinensis, H. to - 
mokunii and presumably also other species form Vietnam form a distinct clade), we 
here place "Nanthela " in the synonymy of Heptathela. 

We want to point out that the systematics of the Heptathelinae (except for the 
fairly well-defined and monophyletic group of species currently placed in the genus 
Ryuthela), as proposed by Ono (2000), Haupt (2003), Platnick (201 1) and summarized 
in Table 2, are in dire need of a comprehensive revision based on genitalic characters 
of both sexes, and ideally also on molecular data. Until this is done (and the missing 
sexes are found), transferring or (in most cases) returning ail non-Ryuthela 
Heptathelinae to Heptathela solves at least the problem of having closely related 
species in différent gênera. This solution is not idéal. Ryuthela appears to be an 
offshoot from within Heptathela, leaving the latter paraphyletic. At the moment no 
other monophyletic groups of species, which are sufficiently distinct to warrant generic 
rank, can be recognized within Heptathela. However, a more comprehensive study 
including the missing sexes (and new species yet to be discovered) may show that 
distinct clades do exist in Heptathela as defined here. If so, the 34 nominal taxa here 
included in Heptathela could be split into several gênera again. In that case plenty of 
generic names will be available. 

Supplementary OBSERVATIONS: Venom glands and moulting position: While 
Haupt's (2003) monograph is a very useful and comprehensive compilation of the 
knowledge of the Mesothelae of that time, his characterisation of the group includes 
two misinterpretations: 

1) Haupt stated that mesothelid spiders do not possess venom glands 
("Mesothelae, however, lack such venom glands. . . . This can only mean that venom 
glands are an apomorphic character of Opisthothelae"; Haupt, 2003: 6), despite the 
detailed description and illustration of such a gland in Liphistius desultor Schiôdte, 
1849 by Millot (Bristowe & Millot, 1933: 1046, pl. 3, figs 1-2). This question has now 



HEPTATHELA KROM VIETNAM 



631 



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been settled by Foelix & Erb (2010a: 7-8, fig. 10; 2010b), who show that extant 
mesothelid spiders (at least females and juvéniles) have venom glands. In liphistiid 
spiders the venom probably does not play an important rôle during prey capture. Upon 
seizing prey, the spiders almost immediately start crushing it with their strong and un- 
usually mobile chelicera, which can be spread at a right angle to each other. Liphistius 
bites to one of us (PJS) had no recognizable toxic effects. 

2) He also stated that "the moulting spécimen turns round and lies on its back 
for the moulting process ..." (Haupt, 2003: 49-50, 95), in the same way as theraphosid 
spiders do. and he concludes "As this turning for moulting does not occur in other 
Megoperculata . . . it seems to be an autapomorphic habit characteristic of Araneae" 
(Haupt, 2003: 50). This is not the case. Haupt was presumably generalizing from a 
single observation captured in a photo (Haupt, 2003: fig. 35D) which shows the final 
moult of a L. trang Platnick & Sedgwick, 1984 maie lying on its back. Looking at the 
exuvia in this photo, however, it becomes clear that the spider actually moulted in a 
venter-down position. When pulling the anterior limbs out of the old cuticle, the 
posterior legs obviously remained stuck in it. As the spider struggled to get free, both 
legs IV and one leg III (still soft and flexible) bent backwards and the spider fell on its 
back. The damage to the discarded exuvia seen in another photo (Haupt, 2003: fig. 
35E), with the fourth pair of legs missing, clearly show that this was not a normal 
moult and that the spider 's fourth pair of legs probably got permanently stuck in the 
exuvia. An examination of this spécimen in Haupt's collection should confirm our 
interprétation. We have had the chance to observe dozens of moults of liphistiid spiders 
in captivity, and we have collected hundreds of exuviae in the field and in captivity. In 
ail cases the exuviae were in a venter-down position, with the leg claws and palpai 
claws gripping the substrate (Figs 65-66). In the interior of their burrows (in nature 
liphistiids always moult inside them) there is not enough space for the spiders to arch 
up and bend backward during moulting. They thus normally moult in the same position 
as uropygids and amblypygids. 

Ectoparasitic mites: The same kind of mites, as were collected from the two 
Heptathela species in southern Vietnam (Figs 67-68 show bite marks), were also found 
on Liphistius at several localities in Laos, Thailand and peninsular Malaysia. At one 
locality in eastern Thailand the majority of liphistiid spiders collected were carrying 
mites. They stay on their hosts permanently and pierce the hard carapaces and cheli- 
cerae but not the soft membranes. Thèse mites were identified as belonging to the 
laelapid genus Ljunghia and are currently being studied by I. Juvara-Bals & B. 
Halliday. Ljunghia includes obligate ectoparasites which naturally occur on primitive 
spiders in SE-Asia and Australia (Domrow, 1975), and which were also found on a 
Central American theraphosid spider in captivity (Moraza et al., 2009). So far only one 
Ljunghia species associated with liphistiid spiders is known, Ljunghia bhstowi 
(Finnegan. 1933) living on Liphistius malayanus in peninsular Malaysia (Finnegan. 
1933). Further species, including the one living on H. australis and H. nui sp. n., will 
be described soon. 

It appears worth pointing out that ail Heptathela spécimens from southern 
Vietnam that carried such mites were reproducing females that either laid eggs (3 H. 
australis) or were gravid (1 H. nui sp. n.). No mites were seen on juvéniles or mature 



HEPTATHELA FR()\1 VILTNAM 



633 




Figs 65-66 

Liphistius sumatranus Thorell, 1890, moulting of an immature maie in captivity. (65) Spider 
partly emerged from the old cuticle. (66) Spider fully emerged. Note that the animal does not lie 
on its back. 



634 



P. J. SCHWENDINGER & H. ONO 




Figs 67-68 

Dorsal view of prosomata of two females showing bite marks caused by ectoparasitic Ljunghia 
mites. (67) Heptathela australis (Ono, 2002); chelicerae spread due to préservation. (68) H. nui 
sp. n., paratype; chelicerae in normal position. 



HEPTATHELA FROM VIETNAM 



635 



maies. This may just be a coincidence, because in Liphistius such mites were also 
found on females that did not reproduce in captivity, and on immature and mature 
maies (unpublished observation). 

ACKNOWLEDGEMENTS 

The MHNG supported the first author's collecting trip to southern Vietnam. The 
Japan Society for the Promotion of Science (JSPS No. 2 1 540487) supported the second 
autor's study. Christine Rollard (MNHN) kindly sent us the holotype of Heptathela 
tonkinensis for examination. Ilinca Juvara-Bals (Cologny, Switzerland) identified the 
mites; Bruce Halliday (CSIRO, Canberra, Australia) confirmed her identification and 
provided additional information. Rainer Foelix (Aarau, Switzerland) pointed out the 
cuticular bases of the receptacular glands and commented on the ultrastructure of spi- 
der genitalia. Norman Platnick (American Muséum of Natural History, New York, 
USA) kindly reviewed the manuscript and pointed out hypothèses and publications that 
we have overlooked. Yoshimi Watanabe (NSMT) produced most of the ink drawings, 
Christina Lehmann-Graber (MHNG) completed them and added other ones, André 
Piuz (MHNG) took the SEM micrographs, Florence Marteau and Lionel Monod (both 
MHNG) arranged ail illustrations into plates. John Hollier (MHNG) checked the 
English text and suggested improvements. The Bulletin of the National Science 
Muséum (Tokyo), renamed the Bulletin of the National Muséum of Nature and 
Science, allowed us to reproduce figures 51-54 from Ono & Schwendinger (1990). 

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BRISTOWE, W. S. & MlLLOT, J. 1933. The liphistiid spiders [by W. S. Bristowe], with an 

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Revue suisse de Zoologie 118 (4): 639-649; décembre 201 1 



A new genus and species of Spilomelinae (Lepidoptera, Pyralidae) 
from the Galapagos Islands, Ecuador 

Bernard LANDRY 1 , Lazaro ROQUE- ALBELO 2 & James E. HAYDEN 3 

1 Muséum d'histoire naturelle, C.R 6434, 1211 Geneva 6, Switzerland. 
E-mail: bernard.landry@ville-ge.ch 

2 Ecologia Environment, 1025 Wellington St., West Perth, W.A. Australia and Curtin 
Institute for Biodiversity and Climate, Curtin University, PO Box U1987, Perth, 
WA 6845, Australia. E-mail: lazaro.roque-albelo@ecologia.com.au 

3 Florida State Collection of Arthropods, FDACS, Division of Plant Industry, 
P.O. Box 147100, Gainesville, Florida, U.S.A., 32614. 

E-mail: james.hayden@freshfromflorida.com 



A new genus and species of Spilomelinae (Lepidoptera, Pyralidae) from 
the Galapagos Islands, Ecuador. - Cheverella galapagensis Landry, 
gen. n. and sp. n. is described as an endémie of the Galapagos Islands, 
Ecuador. Based on a combination of two apomorphies (reduced uncus and 
présence of setose pads on anterodorsal extensions of the maie valva 
medially) this taxon is possibly related to Choristostigma Warren, now 
placed in the Hydriris group of Munroe (1995), but Cheverella lacks the 
setose projection at base of the valve's costa présent in Choristostigma. The 
female has a clearly circumscribed corpus bursae with a short, spine-like 
signum. Cheverella also shares a few characters with members of the Siga 
group. The larva is a borer in stems of the endémie Tournefortia pubescens 
Hook./ (Boraginaceae). 

INTRODUCTION 

During his first expédition to the Galapagos in 1989 at the onset of his inven- 
tory of the Lepidoptera of the archipelago, B. Landry (BL) came across the nicely 
patterned pyraloid treated here, and began to investigate its generic affinities. The 
double praecinctorium pointed to a Spilomelinae, and along the years since 1989 the 
species was shown to several colleagues (James Hayden, Eugène Munroe, Matthias 
Nuss, Michael Shaffer, Aima Solis), who concluded that it was undescribed and that it 
did not belong to any described genus, while various proposais were made with regard 
to phylogenetic affinity to other gênera. In 1983, Eugène Munroe had, in fact, 
examined a spécimen preserved in the Muséum of Comparative Zoology, Cambridge, 
Massachusetts, and added a label saying 'New Pyraustine', a group which at the time 
included the Spilomelinae. 

Here this new taxon is finally described. Its récognition will allow for an 
évaluation of its conservation status and to plan future studies on its ecology. 



Manuscript accepted 28.07.201 1 



640 



B. LANDRY ET AL. 



With more than 3700 species described, the Spilomelinae is one of the two 
largest subfamilies of the Pyraloidea, a group including either one (Pyralidae) or two 
(Crambidae and Pyralidae) families, depending on authors. It is included in the mono- 
phyletic group of pyraloid subfamilies here referred to as Crambiformes (the 
Crambidae, or Crambinina), the other group of subfamilies being known as Pyralidae 
s. str., or Pyralinina, or Pyraliformes. Munroe (1995) treated the subfamily as a tribe of 
Pyraustinae and recorded 1437 species in the Neotropical Région. The Spilomelinae 
are believed to be polyphyletic (Minet. 1982; Solis & Maes, 2003), but this has never 
been tested. The moths are characterized by the absence of chaetosemata, a bilobed 
praecinctorium. a projecting fornix tympani (tympanic frame), a pointed spinula, the 
absence of a gnathos, and the absence of a rhomboidal signum on the bursa copulatrix 
of the female genitalia (Minet, 1982; Nuss et al, 201 1). Other diagnostic characters are 
the loss of the subcostal hamus (frenulum hook) on the maie forewing and the maie 
valva with a clasper in some other conformation than in Pyraustinae (Munroe, 1976: 
8). When Cheverella is compared with the matrix and characters used by Solis & Maes 
(2003) in their phylogenetic analysis of the crambiforme subfamilies, the genus scores 
as a représentative of the Spilomelinae for ail 1 7 characters used. although the shape 
of the médian ridge of the tegumen is here complète from base to apex, not u-shaped 
at base. 

In the Galapagos Islands, there are 42 species of Spilomelinae currently known, 
of which nine are undescribed and under investigation (BL, unpublished). 

MATERIAL AND METHODS 

Spécimens were collected mostly by BL at light during five expéditions to ail 
of the major islands of the Galapagos in 1989, 1992, 2002, 2004, and 2005. The 1989 
material was deposited at the Canadian National Collection of Insects, Ottawa, Canada 
(CNC) while most of the rest is deposited in the Muséum d'histoire naturelle, Geneva 
(MHNG). Many valuable spécimens, including the single reared spécimen came from 
co-author L. Roque- Albelo while he worked at the Charles Darwin Research Station, 
on Santa Cruz Island, Galapagos (CDRS). Additional material came from the 
California Academy of Sciences, San Francisco, California, USA (CAS), the Muséum 
of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA 
(MCZ), and the Natural History Muséum, London, U.K. (BMNH). 

The terminology of the tympanal organs follows Landry (1995). That of geni- 
talia follows Solis & Maes (2003). The manner of giving the label data of the holotype 
and paratypes is presented in Landry (2006) as are the methods used for spécimen 
collecting. 

DESCRIPTIONS 

Cheverella Landry gen. n. Figs 1-8 

Type species: Cheverella galapagensis sp. n. 
Gender: féminine. 

DlAGNOSIS: Cheverella can be separated from the other gênera of Spilomelinae 
by two apomorphies of the maie genitalia, i.e. the reduced uncus and the présence of 



NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 64 1 




FIGS 1-2 

Cheverella galapagensis Landry, sp. n. (1) Holotype (CNC). (2) Darker coloured fcmale 
paratype (MHNG). 

setose pads on anterodorsal extensions of the valva mediobasally, and the clearly 
circumscribed corpus bursae with a short, spine-like signum in the female. Choristo - 
stigma Warren, a mainly North American genus with nine species, also has a reduced 
uncus and setose pads on a transtilla or extensions of the valva as in Cheverella (C. ele - 
gantalis Warren and C. plumbosignalis (Fernald) examined), but its valva is more 
narrow and it has a short, setose projection at base of costa. The female genitalia of 
Choristostigma (C. plumbosignalis (Fernald) examined) differ more strongly in that 
the wide ductus bursae has a large colliculum, the papillae anales are elongate rather 
than triangular, and the signum is very large, oval, and with multiple spines that are 
4-5 X longer than their basai width and irregularly distributed. 

DESCRIPTION: Head (Fig. 3) with antennae filiform, slightly thinner in female, 
reaching almost 2/3 of length of forewing, with dense, short ciliation ventrally, with 
single short seta arising from scale-coated dorsal edge near middle of first 12 flagel- 
lomere approximately, with last flagellomere terminating into distinct sensillum stylo- 
conicum; with ocelli; without chaetosemata; maxillary palpus very short; labial palpus 
curved upward at about half right angle, reaching slightly above eye. 

Forewing rather narrow, 2.4 X longer than largest width. Frenulum simple in 
maie, with 2 acanthae in female. Retinaculum a short bunch of scales below cubital 
stem; maie without frenulum hook. Wing venation (n=l) (Fig. 4): Forewing Se, RI, 
and R2 free, latter from before upper angle of cell; R3 and R4 stalked for most of 
length, stemming from upper angle of cell; R5, M 1-3, and CuAl-2 veins free; Ml and 
M2 well separated at base; M3 from lower angle of cell; 1A+2A clear; 3 A faintly 
indicated. Hindwing with Sc+Rl connected with Rs from 1/2 to 3/4; M 1-3, and 
CuAl-2 veins free; Ml and M2 well separated at base; M2, M3 and CuAl stemming 
from lower angle of cell; anal veins clearly distinct. Abdomen: Maie intersegmental 
membrane VIII-IX without associated sclerites or hair-like scales. Sternum VIII 
broadly sclerotized at base, with short médian extension and long, thin latéral 
extensions reaching apex; tergum VIII with broad sclerotized band along whole 
segment medially. Female segment VII well sclerotized, narrower and longer than 
preceding segment, with tergum a large quadrangular plate. Tympanal organs (n=6) 
(Fig. 5): Tergo-sternal sclerite with broadly rounded, deeply concave ventral margin. 



642 



B. LANDRY ET AL. 



Tympanum plane almost at right angle from sternal plane. Tympanic frame slightly 
projecting ventrad of margin of segment. Tympanic crest short, situated slightly 
anterad of middle. Tympanic drum short, slightly longer than wide, extending anterad 
to base of bridge. Transverse ridge slightly concave medially, without tympanic 
pockets. or unapparent, blending with surface of sternum. Tympanic bridge about 1/3 
length of drums. Praecinctorium only slightly bilobed. 

Maie genitalia (n=3) (Figs 6, 7). Tegumen with wide médian ridge, narrowly ex- 
tended laterally at apex, widened at base and narrowly connected with latéral ridges; 
area between médian and latéral ridges more thinly sclerotized, slightly bulged and 
with scale sockets, as opposed to bare ridges. Uncus reduced, mostly thinly sclerotised, 
rounded, apically setose, occasionally with very short médian dépression dorsally; ven- 
tral margin more thickly sclerotized, occasionally with very tiny point medially. Short 
arms of gnathos (sensu Solis & Metz, 2011) fused with narrow apicolateral ridges of 
tegumen, narrowly triangular, not Connecting medially. Dorsal articulation of valva 
with vinculum of adjacent type (see Solis & Maes, 2003). Costa of valva with medially 
directed projection posterad of dorsal articulation of valva with vinculum; projections 
not connected medially and supporting rounded setose pads ventrally. Valva short, nar- 
rowing to half basai width near middle, apically rounded, with digit-like, medio- 
dorsally recurved projection (sella) medially between pair of ridges, with shallow 
rounded cavity ventrad of sella, with long, abundant setation on basai part of sella 
dorsally. Juxta a thin, elongate plate with lobed ventral and apical margins, with short 
wing-like extensions laterally before middle. Vinculum shorter than tegumen + uncus, 
narrow, with anterior end curved upward, apically blunt in latéral view. Phallus short, 
stout, without pronounced coecum pénis; vesica with bunch of about 20 short, slender 
cornuti. 

Female genitalia (n=3) (Fig. 8). Papillae anales simple, rounded, setose, uncon- 
nected dorsally, with straight sclerotized band at base. Apophyses posteriores straight, 
reaching middle of segment VIII. Latter with well sclerotized plates laterally, sparsely 
setose, expanding apicoventrally toward midline, but medially not connected, dorsally 
approximate on distal half and fused on proximal half. Apophyses anteriores slightly 
curved and longer than posterior ones, not quite reaching middle of segment VII. 
Lamella postvaginalis triangular, located at base and between apical ventral extensions 
of sclerotized plates of segment VIII. Ostium bursae at bottom of cup-like, thinly 
sclerotized antrum. Ductus bursae with girth about 1/3 width of middle of antrum, 
more or less ridged on distal half, proximal half gradually expanding, without colli- 
culum. Ductus seminalis arising subdistally, at slightly less than 1/3 of length of ductus 
bursae from ostium. Corpus bursae circular, with one small, spine-like signum ven- 
trally near distal end. 

ETYMOLOGY: The name is derived from a fréquent interjection heard in 
Ecuador, chévere, which means great, nice, or cool. The unusual maculation of the 
moth prompted this interjection, or a synonym, to me and others who examined it. This 
type of maculation is found in another species of Galapagos Spilomelinae, but not in 
any other members of this subfamily as far as we know. 

BlOLOGY: The Caterpillar of Cheverella galapagensis is a borer in stems of 
Tournefortia pubescens Hook. / (Boraginaceae). One moth was reared by Lazaro 



NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 643 




FIG. 3 

Cheverella galapagensis Landry, sp. n. Head of paratype female from Santa Cruz Island (CNC). 



Roque-Albelo in 1999, from a plant growing on the Barranco, just behind the Charles 
Darwin Research Station on Santa Cruz Island. This endémie species of Tournefortia 
is known to contain pyrrolizidine alkaloids (Roque-Albelo et al., 2009). Thèse are 
known to protect Utetheisa connerorum Roque-Albelo & Landry (Lepidoptera. 
Arctiidae) (formerly mostly known as U. galapagensis (Wallengren)) moths from 
being consumed by Eustala (Araneidae) spiders (Garrett et al, 2008). Whether or not 
Cheverella also stores pyrrolizidine alkaloids remains to be discovered. 

Remarks: The medially directed projections arising from the costal edge of the 
valvae posterad from the dorsal articulations of the valvae with the vinculum are not 
called a transtilla as the définition of this structure is restricted to 'the [dorsal] 
sclerotisation of the diaphragma' (Solis & Maes, 2003). 

Systematic POSITION: The current classification of Neotropical Spilomelinae 
(Munroe, 1995) recognizes 14 groups of gênera and 51 unplaced gênera. Un - 
fortunately, Munroe did not provide diagnostic characters to support his groups, and 
there is no classification available for any other Spilomelinae fauna. Therefore, we 
examined représentatives of Munroe's generic groups to fmd apparently diagnostic 
character combinations, and we comment on their validity and applicability to the 
newly described genus. Cheverella was found to have affinities with the Hydriris and 



644 



B. LANDRY ETAL. 



the Siga groups of Munroe (1995). With the Hydriris group, Cheverella shares a 
reduced uncus without robust bifid spines and the présence of setose pads in the vici- 
nity of the transtilla in Choristostigma Warren. However, in the other gênera of the 
group, the setose pads are in différent positions (on the costa in Geshna Dyar, on the 
tegumen in Hydriris Meyrick) or absent in Nehydriris Munroe. Most of the members 
of this group share tufts of setae at or around the base of the costa of the valva, but 
thèse setae are lacking in Cheverella and Nehydriris, and the female genitalia vary 
among the three gênera for which they are known. The Diagnosis above explains some 
of the différences between the female genitalia of Cheverella and Choristostigma. 
Those of Geshna show a poorly differentiated, elongate corpus bursae without signum, 
while those of Hydriris have a very short ductus bursae and an elongate corpus bursae 
with an appendix bursae and with two large signa showing about 12 long, thin and 
curved projections. The labial palpi are variable. They are upturned in Hydriris and 
Geshna but porrect with downturned apical meron in Choristostigma, so the upwardly 
directed palpi of Cheverella fit in this range. Hence, there is no clear indication that the 
Hydriris group is monophyletic as presently constituted. 

The Siga group of Munroe (1995) comprises large, robust-bodied Neotropical 
moths. Our concept of the group is here informed by our addition of Loxomorpha 
Amsel and Maracayia Amsel, because they share the same structural characters and 
known larval habits, despite their much smaller size. So defined, the Siga group varies 
greatly in maculation, but the forewing postmedial line is usually roundly concave on 
the anal fold. The labial palpi are short or obliquely ascending with a short apical 
meron. The maie genitalia are robust with a moderately inflated sacculus and well- 
sclerotized costa, and the saccus is absent or weakly developed. The sella is aciculate 
in most member taxa (as in the Hydriris group), but it is quadrate in some (Cirrho- 
cephalina Munroe) or with a basai process (Laniifera Hampson). The uncus is bifid 
with a short stalk or none at ail (with the two uncus arms arising from the tegumen 
separately), and the apices of the uncus are armed with robust, bifid chaetae. In the 
female genitalia, where known, the ostium bursae is strongly sclerotized and shaped 
like a funnel or pitcher plant, and the corpus bursae lacks a signum. Known larvae 
(Laniifera, Beebea Schaus, Loxomorpha, Maracayia) are ail borers in Cactaceae. 
Among thèse characters, Cheverella shares the straight and obliquely ascending labial 
palpi, robust genitalia with inflated sacculus, and the distally quadrate sella of some 
members. The breadth of the ostium bursae is similar, but it is only weakly sclerotized 
in Cheverella. The absence of an uncus, biffd or otherwise, would be explained as 
derived from the reduced bilobate condition seen in Zeuzerobotys Munroe. Although 
Cheverella is much smaller than most, it is comparable in size to Loxomorpha species. 
The black and white wing coloration is shared with Zeuzerobotys. 

The following characters are shared by the two groups and Cheverella, so 
although they do not favor placement in either group, they exclude Cheverella from 
many other spilomeline groups. The valve costa is straight or slightly concave, and the 
apex of the valve is rounded but attenuate (narrower than the valve width across the 
costa). The valve sella originates variably from the costal half of the valve or near 
middle of the valve. The hindwing maculation is nearly absent, and the praecinctorium 
is weakly (not strongly) bilobed. 



NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 645 



(4) Cheverella galapagensis Landry, sp. n. Wing venation of paratype maie (MHNG). (5) 
Cheverella galapagensis Landry, sp. n. Tympanal organs of paratype maie, without praecinc- 
torium (MHNG). 

Other characters were examined (tympanal organs, venation) but were found to 
be either invariant or so variable as to be uninformative of relationships at the level of 
generic group. The diagnostic characters in thèse groups (and others) should be evalu- 
ated in a phylogenetic context to détermine their influence and to help uncover the 
relationships of Cheverella. 

The biogeographic relationships of Cheverella are ambiguous but suggest 
testable hypothèses. The Hydriris group includes one Neotropical member (Nehydriris 
of southern Brazil) and the cosmopolitan Hydriris; the other members range to the 
southern Nearctic. As far as known, the Siga group is exclusively Neotropical to 
southern Nearctic, with members distributed in many subregions. Among thèse, 
Beebea guglielmi Schaus is endémie to the Galapagos Islands, so despite the great 
différence in body size and maculation, the possibility of a close relationship should be 
considered. If Cheverella belongs to the Siga group, two hypothèses suggest them- 
selves. A close relationship with Beebea would support rapid evolutionary divergence 
in body size and wing maculation, which might be expected on the Galapagos. 
Conversely, a closer relationship with a mainland taxon would suggest more than one 
dispersai event to the islands or a more complex biogeographic scénario. If Cheverella 
is more related to some other spilomeline group, similarly intriguing hypothèses may 
présent themselves. Cladistic and biogeographic analyses of Spilomelinae are greatly 
needed for choosing among thèse alternative hypothèses. 

Cheverella galapagensis Landry, sp. n. Figs 1-8 

MATERIAL EXAMINED: Holotype maie: 1- 'ECUADOR | GALÂPAGOS | Santa Crùz | 
Los Gemelos | 31.1.1989, M[ercury]V[apour]L[ight] | B. Landry' [printed in black ink on white 
card stock, with 'ECUADOR' sideways on left]; 2- 'HOLOTYPE | Cheverella | galapagensis | 
B. Landry' [hand-written in black ink on red card stock]. Deposited in the CNC. 




FIGS 4-5 



646 



B. LANDRY ET AL. 



PARATYPES: 26 â, 49 9, 1 of unknown sex from the Galapagos Islands, Ecuador: - 
Fernandina: 1 â, SW side, crater rim, G[lobal]P[ositioning]S[system]: 1341 m, S 00° 21.910', 
W 091° 34.034', 12.ii.2005, u[ltra-]v[iolet]l[ight] (B. Landry, P. Schmitz). - Isabela, Alcedo: 
1 9, Lado Este, 700 m elev[ation]., 6.iv.l999, uvl-f[?].l[ight] (L. Roque); 1 S, 1 9, NE slope, 
about 400 m up (S) Los Guayabillos camp, GPS: 892 m elev., S 00° 25.208', W 91° 04.765', 
l.iv.2004, uvl (B. Landry, P. Schmitz); 1 S, 1 9 (DNA voucher, Lepidoptera, M. Nuss 2007, no. 
263), lado NE, 400 m [elev.], camp pega-pega, 15.iv.2002, uvl (B. Landry, L. Roque); 2 â, 1 ? , 
570 m elev., 1 1 .x. 1 998, uvl (L. Roque). - Isabela, Sierra Negra: 3 â , 1 1 km N P[uer]to Villamil, 
1 3.iii. 1 989, M[ercury]V[apour]L[ight] (B. Landry); 1 9, same locality, G[lobalP[osi- 
tioning] S [System]: S 00° 87.613' [sic], W 91° 0.903', 9.iv.2004, uvl (P. Schmitz); 1 9, San[to] 
Tomas, 1200 F[ee]t. Alt[itude]., 22.viii. 1906 (F. X. Williams); 1 â, 1 9 , Alemania, xi.1974 (T.J. 
de Vries); 2 9, Corazon Verde, xi-xii.1974 (T.J. de Vries). - Isabela, Volcan Dai*win: 1 6, 300 
m elev., 15.V.1992, MVL (B. Landry); 1 S, 630 m elev., 16.V.1992, MVL (B. Landry); 1 S (dis- 
sected, slide MHNG ENTO 6070), same data except 17.V.1992; 2 S (one dissected, slide MH- 
NG ENTO 3041), 300 m elev., 20.V.1992, MVL (B. Landry). - San Cristobal: 1 9 (dissected, 
slide CNCPYR 366), pampa zone, 1 8.ii. 1989, MVL (B. Landry); 1 9 , La Toma, ca. 5.6 km East 
El Progreso, GPS: 299 m elev., S 00° 55.356', W 089° 31.089', 23.ii.2005, uvl (B. Landry). - 
Santa Cruz: 1 9 , C[harles]D[arwin]R[esearch]S[tation], arid zone, 19.U989, MVL (B. Landry); 
1 9, 4 km N Puerto Ayora, 20.L1989, MVL (B. Landry); 4 9, Media Luna, pampa zone, 
21.U989, MVL (B. Landry); 2 S, 1 9, same data as holotype; 1 6, C[harles] Darwin 
Resfearch]. Sta[tion]., ait. ± 5 m, 2.ii.l970, at u.v. "blacklight" (R. Silberglied); 1 S, Tortuga 
Resjerve]., W S[an]ta Rosa, 6.H.1989, MVL (B. Landry); 1 of sex unknown, Media Luna, pampa 
zone, 8.ii. 1989, MVL (B. Landry); 1 9, Horneman Farm, 220 m, 16.ii.1964 (D. Q. Cavagnaro); 
1 9, same data except 25.iii.1964; 1 9, same data except 30.iii.1964; 3 9, same data except 
5.iv.l964; 2 9, same data except 3.V.1964; 1 9, same data except 7.V.1964; 1 9, 2 km W Bella 
Vista, 27.ii.1989, MVL (B. Landry); 1 S, Academy Bay, Darwin Research Sta[tion]., 27.ii.1964 
(D. Q. Cavagnaro); 1 â, 3 9, [no précise locality], iii.1969, B.M. 1970-172, Réf. No. L.80 (no 
collector); 1 9 , E[stacion]C[ientifica]C[harles]D[arwin]., 6.iii. 1992, uvl (B. Landry); 1 S (dis- 
sected, slide MHNG ENTO 6069), transition zone, recently eut road, GPS: S 00° 42.528', W 90° 
18.849', 12.iii.2004, uvl (B. Landry, P. Schmitz); 1 9, low agriculture zone, GPS: S 00° 42.132', 
W90° 19.156', 13.iii.2004,uvl (B. Landry, P. Schmitz); 1 9 , Finca S[teve]. Devine, 17.iii.1989, 
MVL (B. Landry); 1 â, grassland, 750 m, 6.iv.l964 (D. Q. Cavagnaro); 1 â, vic[inity]. 
"Mirador", W of Media Luna, ait. ± 620 m, 26.V.1970 (Silberglied); 4 9 (one dissected, slide 
MNHG ENTO 6068), Los Gemelos, 27.V.1992, MVL (B. Landry); 2 9, Station Darwin, 
lumière, x.1964 (J. & N. Leleup); 2 9 , Hacienda Schiess., xi.1974, B.M. 1975-7, Réf. No. L (no 
collector); 1 â, CDRS, Barranco, barrenador de tallos [de] Tournefortia pubecens [sic], emergio 
17.xi. 1 999 (L. Roque); 1 9, 80 [referring to note in notebook of ?T. J. de Vries, deposited at 
CDRS]. - Santiago: 1 9, N side, GPS: 527 m elev., S 00° 13.690', W 90° 44.135', 5.iii.2005, 
uvl (P. Schmitz); 1 9 , Aguacate [camp], 520 m elev., 6.iv.l992, MVL (B. Landry); 3 9 , Central 
[camp], 700 m elev., 9.iv.l992, MVL (B. Landry); 1 6 (wings on slide MHNG ENTO 3043), 
1 9, Aguacate, 520 m elev., 12.iv.1992, MVL (B. Landry). Deposited in the BMNH, CAS, 
CDRS, CNC, MCZ, and MHNG. 

ETYMOLOGY: From the name of the archipelago where this species is pre- 
sumably endémie. 

DlAGNOSIS: Currently unique within the genus Cheverella, this species can be 
separated from presumably related taxa as mentioned in the generic diagnosis above. 
In the Galapagos, this species is unique by virtue of its whitish grey colour with deep 
dark brown markings on the forewing, head, and thorax. An undescribed species of 
endémie Agathodes Guenée (Pyralidae, Spilomelinae) has a similar colour and pattern, 
but its wings are narrower and longer, and its hindwing is completely suffused with 
brown, with a light purple shine. 

Description: MALE (n=27) (Fig. 1). Head white with greyish brown as spot 
between antennae, and ventrally behind eyes. Antenna white, with brown on basai 



NEW GENUS AND SPECIES OF SPILOMELINAE FROM XIII GAI U'AGOS ISl.ANDS 



647 




7 8 



FIGS 6-8 

Cheverella galapagensis Landry, sp. n. Maie genitalia of paratype (MHNG). (6) Genitalia 
without phallus. (7) Phallus. (8) Cheverella galapagensis Landry, sp. n. Female genitalia of 
paratype (CNC). 

flagellomeres. Haustellum and maxillary palpus white. Labial palpus mostly white, 
brown laterally on palpomeres II and III. Thorax white with brown spots as shown, 
deeper brown (almost black) at collar, greyish brown on metascutellum. Foreleg with 
greyish brown on dorsal side, white ventrally; midleg white with brown at tips of fémur 
and tibia; hindleg entirely white. Forewing white with deep dark brown spots and Unes 
as shown, sometimes (1 spécimen) with more extensive dark brown suffusion, 
especially between subapical line and outer margin. Forewing length 6.4-9.6 mm 
(holotype 8.7 mm). Hindwing white with more or less strongly marked subapical and 
apical lines, except in anal sector. Abdomen mostly white, with some greyish brown 
dorsally on ail or most segments. 

Maie genitalia: See generic description. 

FEMALE (n=49) (Figs 2, 3, 8). Forewing length 7.7-10.3 mm. Female geni- 
talia: See generic description. 

Remarks: As mentioned above, L. R.-A. reared the only known larva from the 
host plant Tournefortia pubescens Hook. / (Boraginaceae). This taxon is endémie to 
the Galapagos and found on the islands of Fernandina, Floreana, Isabela, Pinzon, San 



648 



B. LANDRY ETAL. 



Cristobal, Santa Cruz, Santiago, and Wolf (McMullen, 1999). Thus, the moth may also 
occur on the islands of Floreana, Pinzon, and Wolf, where adults have not yet been 
collected. 

The latitude and longitude data of the spécimen collected on Sierra Negra, 
Isabela, 1 1 km north of Puerto Villamil, on 9 April 2004 by P. Schmitz were taken in 
décimal degrees shown above in degrees, minutes, and seconds. In degrees with 
décimal minutes thèse data correspond to S 0° 52.568, W 91° 05.418. 

ACKNOWLEDGEMENTS 

We are thankful to the authorities of Parque Nacional Galapagos and those of 
the CDRS for allowing and facilitating the field work of B. Landry and L. Roque, and 
for permits to export spécimens. BL is greatly indebted to Stewart B. Peck, his Ph.D. 
advisor at Carleton University, Ottawa, for taking him on to explore the Galapagos in 
1989 and 1992, and for his inspiring companionship in the field. This fieldwork was 
supported by an operating grant to S. B. Peck from the Natural Sciences and 
Engineering Research Council of Canada for field research on arthropod évolution. 
Other much appreciated field companions during one or more of BL's five expéditions 
to the Galapagos were Novarino Castillo, Charlotte Causton, Joyce Cook, Moraima 
Inca, José Loaiza, Ricardo Palma, L. Roque-Albelo, Patrick Schmitz, Bradley J. 
Sinclair, and Eduardo Vilema. BL is also grateful to the Charles Darwin Foundation 
and the Galapagos Conservation Trust (UK) for providing financial support for his in- 
vestigations at the BMNH in 2000 and in the Galapagos in 2002. In the BMNH we 
would like to acknowledge the great support received from curators Michael Shaffer 
(deceased) and Kevin Tuck. JEH was supported by a Rea Postdoctoral Fellowship 
(CMNH, Pittsburgh) during his study of Spilomelinae. JEH thanks Aima Solis for pro- 
viding a manuscript of a revision of the Polygrammodes group by E.G. Munroe 
(deceased), which partly infonned the discussion of the Siga group. Finally, we thank 
Philippe Wagneur (MHNG) for his photos of the moths shown here, Florence Marteau 
(MHNG) for producing the plates, Louis Marcotte (Gatineau, Canada) for helping with 
GPS data and jargon, pyraloid workers E. Munroe, M. Nuss, M. Shaffer, and Aima 
Solis for their expertise on this species' taxonomic and systematic status, and the 
reviewers for their comments. 

REFERENCES 

GARRETT, S. E., W. E. CONNER & L. ROQUE-ALBELO 2008. Alkaloidal protection of Utetheisa 

galapagensis (Lepidoptera: Arctiidae) against an invertebrate and a vertebrate predator 

in the Galapagos Islands. Galapagos Research 65: 2-6. 
LANDRY, B. 1995. A phylogenetic analysis of the major lineages of the Crambinae and of the 

gênera of Crambini of North America (Lepidoptera: Pyralidae). Memoirs on Entomo- 

logy, International, Gainesville. Vol. 1. 245 pages. 
LANDRY, B. 2006. The Gracillariidae (Lepidoptera, Gracillarioidea) of the Galapagos Islands, 

Ecuador, with notes on some of their relatives. Revue suisse de Zoologie 113: 437^485. 
MCMULLEN, C. K. 1999. Flowering plants of the Galapagos. Comell University Press, Ithaca 

and London. xiv + 370 pp. 
MINET, J. 1982. Les Pyraloidea et leurs principales divisions systématiques. Bulletin de la 

Société entomologique de France 86: 262-280. 



NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 649 



MUNROE, E. G. 1976. Pyraustinae, Pyraustini (pp. 1-78). In: Dominick, R. B. et al. (eds). The 
Moths of America North of Mexico Fasc. 13.2A, Pyraloidea, Pyralidae (part). E. W. 
Classey Limited and the Wedge Entomological Research Foundation. 78 pp. + plates 
1-4& A-H. 

MUNROE, E. G. 1995. Pyraustinae (pp.53-79). In: Heppner, J.B. (éd.), Atlas of Neotropicai 
Lepidoptera, Volume 3. Association for Tropical Lepidoptera and Scientific Publishers, 
Gainesville. 

NUSS, M. B. LANDRY, F. VEGLIANTE, A. TRÀNKNER, R. MALLY, J. HAYDEN, A. SEGERER, H. 

Ll, R. SCHOUTEN, M. A. SOLIS, T. TROFIMOVA, J. DE PRINS & W. SPEIDEL 2011. 
Global Information System on Pyraloidea. Muséum fur Tierkunde, Dresden. 
www.pyraloidea.org. 

ROQUE-ALBELO, L., S. E. GARRETT, W. E. CONNER 2009. Darwin's moth: Utetheisa in the 
Galapagos Islands (pp. 207-222). In: Conner, W.E. (ed.) Tiger Moths and Woolly Bears: 
Behavior, Ecology, and Evolution of the Arctiidae, Chapter 14. Oxford University Press, 
New York. 

SOLIS, M. A. & K. V. N. MAES 2003. Preliminary phylogenetic analysis of the subfamilies of 
Crambidae (Pyraloidea Lepidoptera). Belgian Journal of Entomology 4: 53-95. 

SOLIS, M. A. & M. A. Metz 2011. Maie génital homology based on musculature originating 
from the tegumen in snout moths (Lepidoptera: Pyraloidea). Arthropod Structure & 
Development 40: 435-448. 



Revue suisse de Zoologie 118 (4): 651-657; décembre 201 1 



A new species of Anthrenus Geoffroy, 1762 

(Coleoptera: Dermestidae) from Oman, with a key to related 

species 

Marcin KADEJ 1 & Jiri HÂVA 2 

1 Department of Biodiversity and Evolutionary Taxonomy, Zoological Institute, 
University of Wroclaw, ul. Przybyszewskiego 63/77, 51-148 Wroclaw, Poland. 
E-mail:entomol@biol. uni. wroc.pl. 

2 Private Entomological Laboratory and Collection, Ûnëtice u Prahy 37, 
252 62 Praha-zâpad, Czech Republic. 

E-mail: jh.dermestidae@volny.cz; http://www.dermestidae.wz.cz. 

A new species of Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae) 
from Oman, with a key to related species. - Anthrenus (Anthrenus) ardoi 
sp. n. from Oman is described. The habirus, antennae, and maie genitalia are 
illustrated and compared to related species. The new species is characterized 
by eye with médian side broadly and deeply emarginated at about anterior 
1/3, 11-segmented antenna and broadly oval scales of the dorsum. Elytra 
covered with oval white patch on each side of the suture at its base, with 
three white, transverse, irregularly interrupted patches. The new species is 
most similar to Anthrenus (s. str.) namibicus Hâva, 2000 and A. (s. str.) 
flavipes flavipes LeConte, 1854. Identification features to externally similar 
species of the genus are given. The most distinctive taxonomic characteris- 
tics concern the tibiae, the maie genitalia and the 9 th abdominal sternite are 
also given. 

Keywords: Coleoptera - Dermestidae - Anthrenus - new species - taxonomy 
- morphology - comparative study - key - Oman. 

INTRODUCTION 

Genus Anthrenus is one of the 57 gênera within the family Dermestidae (Hâva 
2007, 2010), and is characterized by the présence of scales covering almost a whole 
body. The scales often create colorful and very useful in identification patterns on the 
dorsum, and on the visible abdominal ventrites I-V. Currently genus includes 220 
species, grouped in 10 subgenera (Hâva 2003, 2008). Most of the species have been 
described from Palearctic and Afrotropical régions. Only five species have been 
recorded from Oman (Hâva 2007, 2010). Two of them are members of the subgenus 
Anthrenops Reitter, 1881 {A. cervenkai Hâva & Herrmann, 2006, A. coloratus Reitter, 
1881), one of the subgenus Anthrenodes Chobaut, 1898 {A. malkini Mroczkowski, 
1980), another one of Anthrenus s. str. {A. flavipes flavipes LeConte, 1854) and the last 
one oïNathrenus Casey, 1900 (A.jakli Hâva, 2001). In the material from Oman studied 
recently we have found spécimens representing a new species of the nominotypical 
subgenus. Its description is given below. 



Manuscript accepted 28 .07 .20 1 1 



652 



M. KADEJ & J. HÂVA 



MATERIAL AND METHODS 

The following abbreviations to measurements were made: 
total length (TL) - linear distance from anterior margin of pronotum to apex of elytra. 
pronotal length (PL) - maximum length measured from anterior margin to posterior 

margin. 

pronotal width (PW) - maximum linear transverse distance. 

elytral length (EL) - linear distance from shoulder to apex of elytron. 

elytral width (EW) - maximum linear transverse distance. 

The following abbreviations are used in the text: 
IZDBET Institute of Zoology, Department of Biodiversity and Evolutionary 

Taxonomy, Wroclaw, Poland. 
JHAC Private Entomological Laboratory & Collection, Jiri Hâva, Prague-west, 

Czech Republic. 
MZLU Lund University, Lund, Sweden. 
NHMW Naturhistorisches Muséum, Wien, Austria. 
MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland. 
OXUM Oxford University Muséum of Natural History, England. 

Locality labels are cited in the original version. Separate labels are indicated by 
a slash ( / ). Author's remarks are in square brackets [ ]. 

Spécimens have been labeled with red printed label of the following text: 
"HOLOTYPE [or PARATYPE, respectively] Anthrenus (Anthrenus) ardo'x n. sp.". 

Morphological structures were cleared in boiling 1 0% KOH solution, rinsed in 
distilled water, mounted in glycerin, and then examined, measured, and illustrated un- 
der a Nikon Eclipse E 600® phase contrast microscope. External structures were 
examined under a Nikon SMZ-800® stereoscopic microscope. The dissected parts were 
placed with glycerin in a plastic microvial and attached to the pin of the spécimen. 
Photos were taken with a Canon 500D® and a Nikon Coolpix 4500®. 

Terminology: The terminology used in this paper follows Beal (1998) and 
Lawrence & Slipirïski (2010), distribution and classification follows the world cata- 
logue ofHâva (2010). 

SYSTEMATIC PART 

Megatominae Leach, 1815 
Anthrenini Casey, 1900 
Anthrenus Geoffroy, 1762 

Anthrenus (Anthrenus) ardoi sp. n. Figs 1-9 

Material studied: Holotype (S): Oman, Ghuzyan rd. Al Khabural-liberi 10.IV.1985 
leg. Paul Ardô (MZLU). Allotype ($): Oman, Yiti Wadi 7.IV.1985 Paul Ardô (MZLU). 

Paratypes (16 exx. not sexed): (2 exx.) the same datas as holotype (1 MZLU, 1 JHAC); 
(8 exx.) the same data as allotype (5 MZLU, 1 MHNG, 1 IZDBTE, 1 JHAC); (5 exx.) Oman, Al 
Sinain, 10.IV.1985, leg. Paul Ardô (4 MZLU, 1 JHAC); (1 ex.) Oman, Nizwa, 12.IV 1985 leg. 
Paul Ardô (MZLU). 

Etymology: The epithet is a patronym honoring Dr. Paul Ardô (NHMW), who 
collected the type séries material. 



NEW SPECIES OF ANTHRENUS FROM OMAN 



653 




Figs 1-4 

Anthrenus (Anthrenus) ardoi. (1) Habitus, dorsal aspect (holotype, maie). (2) Habitus, dorsal 
aspect (allotype, female). (3) Visible abdominal ventrites I-V (holotype, maie). (4) Visible 
abdominal ventrites I-V (allotype, female). 



Diagnosis: The new species can be differentiated from some of the species 
occurring in Oman by the: 

1) The total number of antennomeres: in A. ardoi antennae with eleven antenno- 
meres; in A. cervenkai Hâva & Herrmann, 2006 and A. coloratus Reitter, 1881 
with nine antennomeres; in A. malkini Mroczkowski, 1980 with ten antenno- 
meres. 

2) The morphology of eye: in A. ardoi eye with médian side broadly and deeply 
emarginated at about anterior 1/3; in A. cervenkai, A. coloratus and A.jakli eye 
oval. 

Due to variation in the dorsal setal pattern, the new species dorsal appearance 
resembles some forms of Anthrenus (s. str.) namibicus Hâva, 2000 and A. (s. str.) 
flavipes flavipes LeConte, 1854 and can be easily confused with them. The distinction 
between thèse two species can be done based on the following characteristics: 

1) In A. ardoi and A. flavipes flavipes tibial spines on first pair of legs absent; in 
A. namibicus présent. 

2) In A. ardoi claws of third pair of legs with one small denticle, in A. flavipes 
flavipes denticle not présent; in A. namibicus three denticle présent. 

3) In A. ardoi apex of paramers slightly curved in to middle, paramers with apex 
and médian lobe slender and narrower, médian lobe with distinct extension in 
1/2 of its length (Fig. 6); in A. flavipes flavipes and A. namibicus, although 
maie genitalia are similar in gênerai shape to A. ardoi, their parameres are 
broader; while in A. namibicus Hâva, 2000 parameres are distinctly curved 
toward the middle, setae présent only on apex of the parameres, médian lobe 
with distinct extension in 3/5 of its length; in A. flavipes flavipes parameres are 
curved to ward the middle, setae présent on apex and in the inner margin of 
parameres, médian lobe without distinct extension. 



654 



M. KADEJ & J. HÂVA 



4) In A. ardoi 9 th abdominal sternite without distinct waist, with the same width 
throughout its length, the apex with cordate, deep indentation, sclerotization on 
the sides and only in the mid section of the upper part - extending to the top and 
underneath; in A. flavipes flavipes 9 th abdominal sternite spatulate, with the 
apex narrow and flat, three times wider posteriorly than the apex width, distinct 
waist in a mid section présent, sclerotization on the sides and the mid section 
only - extending underneath; in A. namibicus 9 th abdominal sternite without 
distinct waist, with the same width throughout its length, the apex almost flat 
with slight and shallow indentation, sclerotization on the sides and only in the 
mid section of the upper part - extending to the top and underneath. 

Description 

Holotype maie. Body. measurements (mm): TL 2.0 PL 0.6 PW 1.3 EL 1.4 EW 
1.7 SL 1.1 SW 1.5. Body convex, rounded, integument brown, fmely punctured, 
covered by broadly oval scales. 

Scales: two times as long as width, their surface mostly with 14 complète, linear 
ribs; the apex of the scale is truncated or rounded and an apical lappet is not présent 
(Figs 8-9). 

Head: is characterized by large convex eyes. Eye with médian side broadly and 
deeply emarginated at about anterior 1/3. Médian ocellus présent on the frons. 
Antennae with 11 antennomeres, with 3-segmented antennal club (Fig. 5); anten- 
nomeres dark-brown. Antenna occupies whole cavity of antennal fossa. Antennal club 
in both sexes occupies less than half length of the antenna, and is distinctly longer than 
length of two basai segments combined. Antennal fossa is completely open along 
latéral margin of the pronotum and occupies 1/3 of the length of latéral margin. Dorsal 
margin of antennal cavity not visible from above. Dorsal surface covered with white, 
light- and dark brown scales (Figs 1-2) as follows: pronotum with mixed and irregular 
light-dark brown and white scales; small white patch near latéral angles enclosing light 
brown patch. 

Elytra: covered with one oval white patch on each side of the suture at its base, 
with three white, trans verse, irregularly interrupted patches: first in basai third, second 
close the middle, and third in apical fourth or fifth. Dark brown patches are présent 
near humeri, in the central part of dise close to suture, and in basai third near the latéral 
margin. The remaining areas between bands are covered with light brown scales mixed 
with single white scales. 

Ventral surface: with ail scales white except for visible ventrites II-V covered 
with mix of yellowish (light brown) and white scales; first abdominal ventrite has stria 
(Figs 3-4). " 

Legs: brown and covered with white scales on dorsal surface. Tibiae without 
tibial spines. Tarsus with two slightly curved claws. Claws of the third pair of legs with 
one small denticle in the half of its length. 

Maie genitalia: as in figure 6. Parameres are deeply U-shaped, covered with 
few short setae on the latéral margins as well as in the central and inner areas. Médian 
lobe in latéral view straight, with a distal end of aedeagus pointing up; in the frontal 



NEW SPECIES OF ANTHRENVS FROM OMAN 



655 




Figs 5-9 

Anthrenus (Anthrenus) ardoi (holotype). (5) Maie antenna. (6) Maie genitalia. (7) 9 th abdominal 
sternite. (8-9) Scales. 

view, wider posteriorly, with narrow apodemes which occupying 1/3 of médian lobe 
length (Fig. 6). 

9 th abdominal sternite as in figure 7. Pygidium with sub-basal, transverse, dark, 
carina-like line. 

Observed VARIATIONS: The dorsal patterns vary in color intensity (either darker 
or lighter). White patches occupies bigger or smaller areas on the pronotum and elytra 
(Figs 1-2). Body measurements for allotype (female): TL 1.9 PL 0.6 PW 1.2 EL 1.4 
EW 1.5 SL 1.0 SW 1.4; for paratypes varying from (mm): TL 2.7-3.3 PL 0.6-0.7 PW 
1.4-1.6 EL 2.1-2.4 EW 1.9-2.4. 

Short key to the related species and to the known Anthrenus species from 
Oman: 

1. Antenna with 10-11 antennomeres 2. 

la. Antenna with 9 antennomeres 3. 

2. Antenna with 10 antennomeres . A. (Anthrenodes) malkini Mroczkowski, 1980 
2a. Antenna with 1 1 antennomeres 4. 

3. Antennal club with two antennomeres, maie antennomere 9 at least 4x 
longer than antennomere 8 A. {Anthrenops) coloratus Reitter, 1881 

3a. Antennal club with three antennomeres, maie antennomere 9 at least 

2.5x longer than antennomere 7 

A. {Anthrenops) cervenkai Hâva & Herrmann, 2006 

4. Inner margin of eye emarginated 5. 

4a. Inner margin of eye oval A. (Nathrenus) jakli Hâva, 2001 



656 



M. KADEJ & J. HÂVA 



5. First pair of legs with tibial spines A. (s. str.) namibicus Hâva, 2000e 

5a. First pair of legs without tibial spines 6. 

6. Ventrites II-V covered with mix of yellowish (light brown) and white 
scales; 9 th abdominal segment without distinct waist, with same width 
throughout its length, apex of segment with cordate, deep indentation; 
apex of paramers slightly curved in to middle, paramers with apex and 
médian lobe slender and narrower, médian lobe with distinct extension 

in 1/2 of its length A. (s. str.) ardoi sp. n. 

6a. Ventrites I-V covered with white and light-brown scales; light-brown 
scales cover posterior margin of ventrites I-V and middle section of ven- 
trite V; 9 th abdominal sternite spatulate, with the apex narrow and flat, 
three times wider posteriorly than the apex width, distinct waist in a mid 
section présent; parameres curved toward middle, setae présent on apex 
and in inner margin of parameres, médian lobe without distinct extension 
A. (s. str.) flavipes flavipes LeConte, 1854 

DISCUSSION: Many of the species of the genus Anthrenus are similar externally. 
However, there are some species that have varying scales color and patterns (A. fia - 
vipes flavipes LeConte, 1854, A. lepidus LeConte, 1854, A. pimpinellae pimpinellae 
(F., 1775), A scrophulariae scrophulahae (L., 1758) ,A. verbasci (L., 1767); compare 
with Beal 1998, Hinton 1945, Kadej 2005a, b). Thus spécimens of particular species 
can be often more similar to other distinct species than to typical form. As a resuit iden- 
tification should be based on the gênerai morphology of the maie genitalia and détails 
in structure of médian lobe, shape of the 9 th abdominal segment and respectively galea 
with lacinia in ail cases. 

ACKNOWLEDGEMENTS 

We would like to thank to Prof. L. Borowiec (Zoological Institute, Wroclaw 
University, Poland) for help in improving this manuscript, to R. Danielsson (MZLU) 
for the loan of material and J. Cooter (OXUM) for help with translation of the 
manuscript. This work was supported by funding (project no 1244/M/IZ/201 1) from 
the Institute of Zoology, University of Wroclaw, Poland. 

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Dermestidae). Transactions of the American Entomological Society 124: 271-332. 
CASEY, T. L. 1900. Review of the American Corylophidae, Cryptophagidae, Tritomidae and 

Dermestidae with other studies. Journal New York Entomological Society 8: 51-172. 
CHOBAUT, A. 1898. Description de quelques espèces et variétés nouvelles de Coléoptères 

algériens. Revue d'Entomologie Caen 17: 74-88. 
FABRICIUS, J. C. 1775. Systema Entomologie, sistens Insectorum classes, ordines, gênera, 

species, adiectis synonymis, locis, descriptionibus, observationibus. Flensburgi et 

Lipsiae: Officina Libraria Kortii, xxxii + 832 pp. 
GEOFFROY, E. L. 1762. Histoire abrégée des Insectes qui se trouvent aux environs de Paris, dans 

laquelle ces animaux sont rangés suivant un ordre méthodique. 1. Paris: Durand, xxviii 

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HÂVA, J. 2000. New intcresting Dermestidae (Coleoptera) from the world with descriptions of 

ten new species. Verôffentlichungen Naturkundemuseum Erfurt 19: 161-171. 
HÂVA, J. 2001. Anthrenus (Nathrenus) jakli sp. n. (Coleoptera: Dennestidac) from the Sultanate 

of Oman, and distributional notes on some other Anthrenus species. Acta Societatis 

Zoologicae Bohemîae 65: 1-3. 
HÂVA, J. 2003. World Catalogue of the Dermestidae (Coleoptera). Studie a Zprâvy Oblastniho 

Muzea Praha-vychod v Brandyse nad Labem a Staré Boleslavi, Supplementum 1 : 1 - 1 96. 
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Palaearctic Coleoptera. Volume 4. Elateroidea, Derodontoidea, Bostrichoidea, Lymexy- 

loidea, Cleroidea and Cucujoidea. Apollo Books, Stenstrup, 935 pp. 
HÂVA, J. 2008. Description of a New Subgenus Setapeacockia subgen. nov. of the genus 

Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae: Anthrenini) from Central Asia. 

Litvijas Entomologos 45: 43-45. 
HÂVA, J. 2010. Dermestidae of the World (Coleoptera). An interactive manual (opened in 2004), 

available from: http://www.dermestidae.wz.cz (last modification 18.01.2010) 
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Yémen - Part I. Mitteilungen des Internationalen Entomologischen Vereins 31:1-10. 
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London: Order of the Trustées ofthe British Muséum, viii + 443 pp. 
KADEJ, M. 2005a. Morphological variability of pimpinellae-group (Coleoptera: Dermestidae: 

Anthrenus) with a key to identification. Part 1 . Polskie Pismo Entomologiczne 74: 

117-135. 

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scrophulariae-group with a key to identification. Polskie Pismo Entomologiczne 74: 
509-521. 

LAWRENCE, J. F. & SlipiNSKI A. 2010. 6.1. Dermestidae Latreille, 1804 (pp. 198-206). In: 
LESCHEN, R. A. B., BEUTEL, R. G., & LAWRENCE, J. F. (eds). Coleoptera, beetles. 
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partim). Handbook of Zoology, a natural history of the phyla of the animal kingdom. 
Volume IV. Arthropoda: Insecta. Part 38 (KRISTENSEN, N. P. & BEUTEL, R. G., eds). 
Walter de Gruyter, Berlin, Germany. 

LeCONTE, J. L. 1854. Synopsis of the Dermestidae of the United States. Proceedings of the 
Academy of the Natural Sciences of Philadelphia 7: 106-1 13. 

LlNNAEUS, C. von 1758. Systema Naturae per régna tria naturae, secundum classes, ordines, 
gênera, species, cum characteribus, differentiis, synonymis, locis. Editio décima, refor- 
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gênera, species, cum characteribus, differentiis, synonymis, locis. Editio décima tertia, 
ad Editionen duodecimam reformatam. Tom L, Pars II. Holmiae: Laurentii Salvii, pp. 
533-1327 + 37 pp. 

MROCZKOWSKI, M. 1980. Insects of Saudi Arabia. Coleoptera: Fam. Dermestidae. Part 2, 
Description of three new species. Fauna Saudi Arabia 2: 124-126. 

REITTER, E. 1881. Bestimmungs-Tabellen der europàischen Coleopteren. III Heft. I. Auflage. 
Enthaltend die Familien: Scaphidiidae, Lathridiidae und Dermestidae. Verhandlungen 
der Kaiserlich-Kôniglichen Zoologisch-Botanischen Gesellschaft in Wien 30: 41-94. 



Revue suisse de Zoologie 118 (4): 659-666; décembre 201 1 



First record of the subfamily Pycnocheiridiinae from 

South America, with the description of Leptocheiridium pfeiferae 

gen. n., sp. n. (Arachnida:Pseudoscorpiones: Cheiridiidae) 

Volker MAHNERT* & Jurgen SCHMIDL** 
* Muséum d'histoire naturelle, Ville de Genève, case postale 6434, 1211 Geneva 6. 

Switzerland. E-mail: volker.mahnert@wanadoo.fr 
** Friedrich-Alexander-Universitât Erlangen-Nùrnberg, Department Biologie, AG 

Ôkologie, Staudtstr. 5, 91058 Erlangen, Germany. 

E-mail: jschmidl@biologie.uni-erlangen.de 

First record of the subfamily Pycnocheiridiinae from South America, 
with the description of Leptocheiridium pfeiferae gen. n., sp. n. 
(Arachnida: Pseudoscorpiones: Cheiridiidae). - The cheiridiid subfamily 
Pycnocheiridiinae is recorded for the first time from South America. 
Leptocheiridium gen. n. and its type species pfeiferae sp. n. are described 
and illustrated. The new genus shares with Pycnocheiridium Beier, 1964 
(type genus of the Pycnocheiridiinae Beier, 1964) the same morphology of 
walking legs, eight trichobothria on the fixed chelal finger, only two on the 
movable finger, and the présence of strongly clavate vestitural setae. 
Leptocheiridium gen. n. is well characterized by the shape of its slender 
pedipalps, the présence of three sub-equal setae in the rallum, the présence 
of five setae on the cheliceral hand, the morphology of the female galea, and 
the présence of a well-developed transverse furrow on the carapace. 

Keywords: Taxonomy - ecology - neotropical région - Ecuador - bark 
dweller. 

INTRODUCTION 

Beier (1964) erected, within the family Cheiridiidae, the subfamily Pycnochei - 
ridiinae Beier, 1964 for the monotypical genus Pycnocheiridium Beier, 1964 (with its 
type species mirum Beier. 1964). This genus. takes an intermediate position between 
Cheiridiidae Hansen, 1893 and Pseudochiridiidae Chamberlin, 1923 in having clearly 
separated fémur and patella on walking legs VU, femur+patella of legs III and IV with 
a distinct, nearly vertical suture (Judson, 1992), eight trichobothria on fixed chelal 
finger, and 5 setae on the cheliceral hand (pseudochiridiid characters), whilst having 
normal, unmodified coxae IV also in females, normal-shaped tergites (not chevron- 
shaped), a rallum with three setae, of which the anterior one is larger and dentate 
(Judson, 1992), and only two trichobothria on the movable chelal finger (cheiridiid 
characters). Only a few spécimens of Pycnocheiridium mirum are recorded from South 
Africa (Eastern Cape, Kwazulu-Natal). Nothing is known on its biology (one female 
was "sifted, (from) very humid rain forest, very thick humus"), and it "appears to be 
one of the rarest South African species (of pseudoscorpions)" (Judson, 1992). 



Manuscript accepted 16.09.201 1 



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V. MAHNERT & J. SCHMIDL 



Harvey (1992) placed Cheiridiidae (without mentioning the subfamily 
Pycnocheiridiinae), together with four other families, in the superfamily Garypoidea. 
Murienne et al. (2008) re-established the superfamily Cheiridioidea, including Cheiri- 
diidae. Pseudochiridiidae and Sternophoridae, by using molecular séquence data. 
However. their analysis is based on a very low number of gênera and species (three) 
studied. The phylogeny of Cheiridioidea and its validity as a superfamily has also been 
discussed by Judson (2000). 

In the course of the Worldwide research project (MACAG - Monitoring of 
Arthropods along Climate and Altitude Gradients: see Schmidl, 2009) conducted by 
the junior author (JS), 339 samples from tree bark were collected in 2007, 2008 and 
2009 at 1000 m (warm rainforest), 2000 m (cloud forest) and 3000 m (Paramo tran- 
sition forest) in the Podocarpus National Park in Ecuador, to analyze species compo- 
sitions of arthropods in regard to patterns of diversity between trees, plots and altitu- 
dinal levels, and for the évaluation of the monitoring performance (application) of the 
corticolous arthropod fauna. Samples were taken using the barkspray method (protocol 
according to Schmidl, 2007). A defined area of a living, unbroken tree trunk bark is 
sprayed with a pyrethroid insecticide spray. Pyrethroids are contact poisons which are 
highly spécifie to arthropods and block voltage-sensitive sodium channels in the nerve 
membrane. Within a few minutes the arthropods lose the ability of coordinated move- 
ments and tumble down from the substratum. Pyrethroids show an immédiate effect on 
ail arthropods. but are relatively harmless for higher vertebrates. Before spraying, a 
plastic sheet is tightly pinned to the bottom of the sampled trunk to collect ail affected 
arthropods, which after 1 5 minutes are brushed into a small, tree-wise labelled plastic 
bag (Whirl Paks) containing 70% ethanol. 

Among the 287 pseudoscorpion spécimens, representing seven families and 15 
morphospecies, collected during this field survey, we found three spécimens of a genus 
that we place with hésitation in the cheiridiid subfamily Pycnocheiridiinae. A new 
genus and species are described and figured below. It is well differentiated from 
Pycnocheiridium, the only other known genus in this subfamily. 

DESCRIPTION 

Leptocheiridium gen. n. 

DlAGNOSlS: Member of Cheiridiidae, 'subfamily Pycnocheiridiinae. Tégument 
normally sclerotized, vestitural setae of carapace, tergites and pedipalps strongly broa- 
dened, leaf-like. Carapace subtriangular, with one pair of eyes, two distinct transverse 
furrows, the subbasal one flattened, metazone narrow. Most tergites divided, with 
unmodified latéral borders. Manducatory process rounded, with two marginal setae. 
Sternites IV to X divided, XI undivided; anal cone with 2 dorsal and 2 acute ventral 
setae and located between tergite XI and sternite XI; maie genitalia of typical cheiri- 
diid morphology (latéral sacs long, well developed, atrium of ejaculatory canal well 
developed. médian génital sac short), female genitalia consisting of three major 
cribrate plates, the latéral ones more sclerotized than the larger médian one. Pleural 
membrane striate, partly covered with granula. Chelicera with 5 setae on hand; galea 
short, distinct apical rami présent only in female, subgaleal seta near galea base, rallum 



LEPTOCHEIRIDIUM GEN. N. FROM ECUADOR 



661 



with three setae, the distal one with a few fine denticles on anterior margin. Pedipalps 
coarsely granular, hand nearly parallel-sided; teeth on both fingers mostly acute and 
contiguous; fixed finger with 8 trichobothria (ail situated in basai half of finger), 
movable finger with 2 trichobothria (b and /); long venom duct présent in both fingers. 
Coxae IV much wider than coxae I. Leg I: fémur distinctly longer than patella and with 
articulation, tarsal segments fused, without visible suture. Leg IV: fémur much shorter 
than patella, suture nearly vertical, tarsal segments fused, without visible suture, no 
tactile tarsal seta; undivided arolia shorter than smooth claws. 

TYPE SPECms: Leptocheiridium pfeiferae sp. n. 

ETYMOLOGY: The genus name is a combination of Cheiridium with the Greek 
adjective "leptos" (thin, slender), referring to the slender pedipalps. 

AFFINITIES: Leptocheiridium gen. n. is distinguished from Pycnocheiridium by 
its slender pedipalps, the distinct transverse furrows on its carapace, the présence of 
5 setae on its cheliceral hand, the female galea with 6 apical/subapical rami, and the 
three sub-equal setae of its rallum (in Pycnocheiridium the anterior seta is distinctly 
larger: Judson, 1992). 

REMARKS: As currently defmed, the subfamily Cheiridiinae of the Cheiridiidae. 
is characterized by having a reduced number of trichobothria on fixed (at most 7) and 
movable (at most 2) chelal fingers, by the fused fémur and patella of leg I and the at 
least partly fused femur+patella of leg IV; coxa of leg IV also in females unmodified, 
not enlarged; carapace with two distinct transverse furrows, with depressed metazone 
(except in Apocheiridium), and two eyes; cheliceral hand with 4 setae; first seta of 
rallum enlarged. 

The subfamily Pycnocheiridiinae differs from the Cheiridiinae by having fémur 
and patella of leg I well separated and articulated, femur+patella of leg IV with dis- 
tinct, nearly vertical suture, and eight trichobothria on the fixed chelal finger. 
Considering thèse différences, the Pycnocheiridiinae may even deserve familiar rank. 

The new genus Leptocheiridium gen. n. confirms the main subfamiliar 
characters, but differs by the following: number of setae on cheliceral hand (4 vs 5) and 
morphology of rallum (3 setae, the first one not enlarged), which are probably of 
generic, but not of subfamiliar importance. 

Leptocheiridium pfeiferae sp. n. Figs 1-10 

HOLOTYPE: MHNG, without registration number; 6\ Ecuador, Prov. Zamora, env. 
Estaciôn Cientifica San Francisco (ECSF), 2000m asl, cloud forest, on tree bark by bark 
spraying; 31.VIII.2009, lg. T. Pfeifer (tree # EL6B05). 

PARATYPES: MHNG, without registration number; 1 9 1 deutonymph; same locality and 
method, 29.VIII.2009; lg. T. Pfeifer (tree # EQ5B07). 

Remark: The three spécimens were collected on two différent trees several 
hundred mètres apart. Tree EL6B05 had a dimension at breast height (dbh = diameter 
at 130 cm height) of 26 cm, a médium coverage of climbing plants, a dense cover of 
epiphytes (mosses, lichens, algae), and a médium rough bark. Tree EQ5B07 had a dbh 
of 23 cm, differing from the previous tree by a very smooth bark. Both trees had an 
intact, unbroken bark. 



662 



V. MAHNERT & J. SCHMIDL 




FIGS 1-7 

Lepîocheiridium pfeiferae gen. n., sp. n., holotype. (1) Carapace, with granules enlarged. (2) Left 
chelicera S , with rallum and galea of 6 and 9 (paratype) enlarged. (3) Left pedipalp. (4) Tricho - 
bothrial pattern. (5) Tergal seta, enlarged. (6) Left leg I. (7) Right leg IV. Scale unit 0.1 mm. 



LEPTOCHEIRIDIUM G EN. N. FROM ECUADOR 



663 




FIGS 8-10 

Leptocheiridium pfeiferae gen. n., sp. n. (8) Sternites II/III of maie. (9) Sternites II/III of female, 
with cribrate plates. (10) Trichobothrial pattern of deutonymph. 

DESCRIPTION OF ADULTS: Pedipalps, and carapace brown, tergites yellowish 
brown, legs including coxae reddish brown. Carapace (Fig. 1) subtriangular. broader 
than long (length/breadth = 0.9), with two trans verse furrows, médian furrow broad 
and laterally distad directed, granular, the subbasal one distinct only laterally; two 
distinct eyes with flat lenses; anterior margin straight, without epistome; coarsely gra- 
nular; with 42-46 strongly clavate, leaf-like setae (4 along anterior, 7-9 along posterior 
margin). Ail tergites visible dorsally, I-X divided, coarsely granular, setae leaf-like 
(Fig. 5), chaetotaxy of half-tergites: 5-6/6/6/5-6/6/5-6/5-6/4-5/5/3-4, on IV-X one seta 
anterolaterally placed, tergite XI with 6 setae (no tactile setae). Manducatory process 
scaly, with 2 marginal setae; pedipalpal coxa coarsely granular, with 8-9 clavate setae, 
0-2 smooth setae and one distal tactile seta, coxa I with 6-8 setae (2 slightly clavate), 
II with 8-13 smooth setae, III with 15-20, IV with 32-35 smooth setae; génital opercu- 
lum of â with 28 long setae (Fig. 8), 2/2 short entrance setae, génital organs similar to 
that of other cheiridiid species (e. g. Dumitresco & Orghidan, 1981; Judson, 1992); 
génital operculum of 9 (Fig. 9) with 15 marginal and discal setae, medial cribrate plate 
apparently divided into one big and one small part, latéral cribrate plates small (Fig. 9); 



664 



V. MAHNERT & J. SCHMIDL 



sternites IV-X divided, the anterior ones scaly, the posterior ones ctenoid-scaly 
sculptured, some lyrifïssures présent in front of and between setae, XI undivided, 
ctenoid-scaly; setae on anterior sternites smooth, acute, on posterior ones slightly 
clavate (except latéral setae and those of XI), chaetotaxy of half-sternites: 6-8, supra- 
stigmal setae/7-9+1 suprastigmal seta/8-9/7-8/6-7/5-7/5-6/4, latéral seta on V-X short. 
Anal cone situated between tergite and sternite XI, with 2 smooth dorsal and 2 (3 in 
holotype) ventral setae. Pleural membrane shortly granular-striate. 

Chelicera (Fig. 2) with 5 long acute setae, b distinctly shorter than sb, es as long 
as sb; fixed finger with 3 retrorse teeth and 2-3 subapical granula, lamina exterior thin; 
movable finger of S with a few thin, thorn-like projections on latéral side, one also at 
base of galea, with a tooth-like subapical lobe, subgaleal seta reaching clearly beyond 
tip of galea in S , reaching almost to tip of galea in 9 , galea of cT short (Fig. 2) and 
rounded, with 2 tiny rounded apical tubercles, galea of 9 long, with 6 short sub- 
apical/apical rami; serrula exterior with 12 lamellae; rallum (Fig. 2) with 3 setae, 
anterior margin of distal one with 2-3 denticles. 

Pedipalp (Fig. 3) coarsely granular, vestitural setae leaf-like, chelal finger with 
fine curved setae; trochanter with small dorsal hump, 2.0-2.1 times longer than broad; 
fémur distally slightly broadened, with a distinct pedicel, 4.3 times longer than broad; 
patella 3.3 times longer than broad; hand parallel-sided, with pedicel 2.7 times (â) or 
2.3 times ( 9 ) longer than broad and 1 .6 times ( S ) or 1 .5 times ( 9 ) longer than finger; 
chela with pedicel 4.4 times (â) or 3.9 times (9), without pedicel 4.2 times (â) or 3.6 
times ( 9 ) longer than broad; teeth of chelal fmgers acute, slightly retrorse (in distal 
third), then becoming flattened and rounded in basai part, fixed finger with 33 (â) or 
29 (9) teeth, movable finger with 27 (â) or 21 (9) teeth; long venom duct présent in 
both fingers, nodus ramosus situated in middle of finger; 10 trichobothria (Fig. 4), ail 
8 on fixed finger situated in basai half, 2 on movable finger situated in basai third (b 
and t, following Harvey, 1990); sensillum distal to b. 

Leg I (Fig. 6): fémur and patella articulated, fémur 1.6 times (6) or 1.4 times 
(9) longer than patella, fémur 2.4-2.5 times longer than deep, patella 1.6 times, tibia 
2.3-2.6 times, tarsus 4.3-4.4 times longer than deep, tarsus 1.4-1.5 times longer than 
tibia; leg IV (Fig. 7): fémur much shorter than patella, distinct suture nearly vertical, 
femur+patella 4.4-4.6 times longer than deep, tibia 4.9 (â) or 4.5 ( 9 ) times, tarsus 5.2 
(cT ) or 5.6 ( 9 ) times longer than deep, no tactile setae présent; undivided arolia shorter 
than smooth claws. 

MEASUREMENTS (in mm) â (9): Total length 1.4; carapace length/breadth 
0.43/0.49 (0.44/0.51); pedipalp length/breadth: trochanter 0.23/0.1 1 (0.24/0.12), fémur 
0.44/0.10 (0.47/0.11), patella 0.40/0.12 (0.44/0.13), hand with pedicel 0.36/0.13 
(0.37/0.12), length of pedicel 0.03, finger length 0.22 (0.24), chela length with pedicel 
0.58 (0.62); leg I length/depth: fémur 0.19/0.08 (0.19/0.08), patella 0.12/0.08 
(0.14/0.08), tibia 0.16/0.05 (0.16/0.07), tarsus 0.22/0.05 (0.23/0.05); leg IV 
length/depth: femur+patella 0.40/0.09 (0.39/0.09), tibia 0.31/0.06 (0.32/0.07), tarsus 
0.28/0.05 (0.31/0.06). 

Description of deutonymph: Carapace 0.80 times broader than long, co - 
arsely granular, medial transverse furrow distinct, only latéral dépressions of subbasal 



LEPTOCHEIRJDIUM GEN. N. FROM ECUADOR 



665 




FIGS 11-12 

Photographs of tree EL6B05 (locality of holotype) (11) and of tree EQ5B07 (locality of para- 
types) (12). 

furrow distinct, metazone unsclerotized; 2 flattened eyes; 25 leaf-like setae, 4 along 
anterior and 6 along posterior margin; tergites divided, smooth, half-tergites with 
3 leaf-like marginal setae, tergite XI with 4 setae; manducatory process with 2 setae, 
pedipalpal coxa with 5 leaf-like setae, coxa I/II with 3, III with 5, IV with 7 smooth 
acute setae, half-sternites mostly with 3 acute setae, sternite XI with 4 setae. Chelicera 
with 5 setae on hand, galea with 4 apical/subapical rami, serrula exterior with 1 0(?) 
lamellae. Pedipalp: fémur 3.5 times longer than broad (0.26 mm/0.07 mm); patella 2.9 
times longer than broad (0.24/0.08); hand with pedicel 2.2 times (0.23/0.11) longer 
than broad and 1.36 times longer than hand with pedicel, finger length 0.17 mm; chela 
with pedicel 3.8 times, without pedicel 3.6 times longer than broad; fixed finger with 
23, movable finger with 21 teeth. Six trichobothria on fixed finger (3 anti- and 3 
paraxial ones), movable finger with one trichobthrium (Fig. 10). 

ETYMOLOGY: This species is dedicated to Miss Tina Pfeifer (Fiirth) for collec- 
ting and processing the pseudoscorpions in the 2009 sampling campaign. She invested 
much time for taxonomic pre-sorting and photographie documentation of this 
collection in the course of her "Zulassungsarbeit" at the University of Erlangen- 
Nuremberg. 



666 



V. MAHNERT & J. SCHMIDL 



ACKNOWLEDGEMENTS 

The junior author is grateful to our coopération partners in Ecuador, the 
Research Unit 816 of the Deutsche Forschungsgemeinschaft (DFG), the Ministerio del 
Ambiente Ecuador (Direccion Régional Loja - Zamora Chinchipe) for granting the 
export permits, and especially the ECSF station managers Félix Matt and Jôrg 
Zeilinger for practically facilitating our stays and research at the ECSF research 
station. Spécial thanks go to Maurice Leponce (RBINS Brussels) for the congenial 
project partnership and to David Prôtzel (Erlangen) for an excellent performance in the 
2009 sampling campaign. Sincère thanks are addressed to Dr Mark S. Harvey (Perth) 
and Dr Mark L. I. Judson (Paris) for their helpful comments on the taxonomic status 
of this new taxon. 

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(Arachnida, Chelonethi, Cheiridioidea) from Dominican amber. Zootaxa 1393: 45-51. 
MURIENNE, J., HARVEY, M. S. & GIRIBET, G. 2008. First molecular phylogeny of the major 

clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and 

Evolution 49: 170-184. 

SCHMIDL, J. 2007. Barkspray - Technical manual. Unpublished report, University Erlangen- 
Nuremberg, 2 pp. 

SCHMIDL, J. 2009. MACAG - monitoring of arthropods along climate and altitude gradients - 
a multiscale approach for evaluating patterns and responses (p. 192). In: HOLZHEU, S. & 
B. THIES, B. (eds). Dimensions of ecology*- From global change to molecular ecology. 
Book of Abstracts, 39th Annual Conférence of the Ecological Society of Germany, 
Austria and Switzerland (Bayreuther Forum Ôkologie 115). Bayreuth, 246 pp. 



Revue suisse de Zoologie 118 (4): 667-693; décembre 201 1 



Four new species from China and Southeast Asia 
(Diptera, Lauxaniidae, Homoneurinae) 

Li SHI 1 ' 2 , Ding YANG 1 & Stephen D. GAIMARI 3 

1 Department of Entomology, China Agricultural University, Beijing 100193, China. 
E-mail: dyangcau@yahoo.com.cn 

2 Collège of Agronomy, Inner Mongolia Agricultural University, 
Huhhot 010019, China. E-mail: li_shi_lauxaniid@yahoo.com.cn 

3 Plant Pest Diagnostics Center, California Department of Food and Agriculture, 
3294 Meadowview Road, Sacramento, CA 95832-1448, USA. 

E-mail: stephen.gaimari@cdfa.ca.gov 

Four new species from China and Southeast Asia (Diptera, Lauxa- 
niidae, Homoneurinae). - Four new species are described and illustrated, 
including Cestrotus quadrimaculatus sp. n., Dioides merzi sp. n., Noo- 
namyia bipunctata sp. n. and Noonamyia flavoscutellata sp. n., and seven 
species are recorded for the first time from Vietnam, Thailand, Philippines 
and/or Malaysia, including Cestrotus flavoscutellatus Meijere, 1910, 
Cestrotus liui Shi, Yang & Gaimari, 2009, Cestrotus heteropterus Shi, Yang 
& Gaimari, 2009, Dioides furcatus Shi & Yang, 2009, Dioides incurvatus 
Shi & Yang, 2009, Noonamyia sabahna Sasakawa & Pong, 1990, Noona - 
myia sasakawai Papp, in Papp, Merz & Fôldvâri, 2006 and Phobeticomyia 
uncinata Shi & Yang, 2009. The species Noonamyia sasakawai is 
redescribed and illustrated completely according to two paratypes and 
several additional spécimens. A key to ail species of the genus Noonamyia 
of the world is presented. 

Keywords: Lauxaniidae - Homoneurinae - new species - key - Southeast 
Asia. 

INTRODUCTION 

The family Lauxaniidae is one of the larger Acalyptratae families. There are 
more than 160 gênera and nearly 2000 described species, distributed Worldwide except 
for Antarctica. The following four gênera, which belong to the subfamily Homo - 
neurinae, Cestrotus Loew, 1862, Dioides Kertész, 1915, Noonamyia Stuckenberg, 
1971 and Phobeticomyia Kertész, 1915 have conspicuous wing patterns. The genus 
Cestrotus includes 1 1 described species in the Oriental Région and 1 3 in the Afro - 
tropical Région; the genus Dioides includes 6 described species in the Oriental Région 
and 1 in the Australasian/Oceanian Région; the gênera Noonamyia and Phobeticomyia 
are represented by 12 and 6 described species respectively, distributed exclusively in 
the Oriental Région (Shi, Yang & Gaimari, 2009; Shi, Li & Yang, 2009a, 2009b; Shi 
& Yang, 2009). 



Manuscript accepted 26.09.201 1 



668 



L. SHI ETAL. 



In the présent paper, four new species are described and illustrated, namely 
Cestrotus quadrimaculatus sp. n., Dioides merzi sp. n., Noonamyia bipunctata sp. n. 
and Noonamyia flavoscutellata sp. n., and seven species are recorded for the first time 
from Vietnam, Thailand, Philippines and/or Malaysia: Cestrotus liui Shi, Yang & 
Gaimari, 2009, Cestrotus heteropterus Shi, Yang & Gaimari, 2009, Dioides furcatus 
Shi, Li & Yang, 2009, Dioides incurvatus Shi, Li & Yang, 2009, Noonamyia sabahna 
Sasakawa & Pong, 1990, Noonamyia sasakawai Papp, in Papp, Merz & Fôldvâri, 2006 
and Phobeticomyia uncinata Shi, Li & Yang, 2009. The species Noonamyia sasakawai 
is redescribed and illustrated completely based on two paratypes and additional spé- 
cimens, and a key to separate ail species of the genus Noonamyia is presented. 

MATERIAL AND METHODS 

The spécimens examined are deposited in the following collections: 
BMNH The Natural History Muséum, London, UK 
BPBM Bernice P. Bishop Muséum, Honolulu, Hawai'i, USA 
CASC California Academy of Sciences, San Francisco, California, USA 
CAUC Entomological Muséum of China Agricultural University, Beijing, China 
CNCI Canadian National Collection of Insects, Ottawa, Ontario, Canada 
CSCA California State Collection of Arthropods, Sacramento, California, USA 
HNHM Hungarian Natural History Muséum, Budapest, Hungary 
MHNG Muséum d'histoire naturelle, Genève, Switzerland 
MZSP Museu de Zoologia, Universidade de Sào Paulo, Sào Paulo, Brazil 
NHMW Naturhistorisches Muséum Wien, Vienna, Austria 
NMSA KwaZulu-Natal Muséum, Pietermaritzburg, South Africa 
QSBG Queen Sirikit Botanic Garden, Chiangmai, Thailand 
USNM National Muséum of Natural History, Washington, DC, USA 
ZISP Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia 

Terminology follows Me Alpine (1981) and Shewell (1987). Genitalia prépara- 
tions were made by removing and macerating the apical portion of the abdomen in cold 
saturated NaOH for 2-6 hours. After examination, genitalia were transferred to fresh 
glycerin and stored in a microvial pinned below the spécimen. 

The following abbreviations are used: a - anterior seta, aer - acrostichal seta, ad 
- anterior dorsal seta, anepst - anepisternal seta, app - apical posterior seta, apv - apical 
ventral seta, av - anterior ventral seta, de - dorsocentral seta, kepst - katepisternal seta, 
oc - ocellar seta, or - fronto-orbital seta, pd - posterior dorsal seta, prsc - prescutellar 
seta, pv - posterior ventral seta, sa - supra-alar seta. 

SYSTEMATIC PART 

Cestrotus quadrimaculatus sp. n. Figs 1-5, Plates Al, A3, Bl 

MATERIAL: HOLOTYPE 6 (MHNG): Thailand, Kamphaeng Phet Province, KhlongLan 
District, nr KhlongLan Watf, 280 m, 16.07.51N/99.16.41E, 11-12.XII.2003, P. Schwendinger. 
PARATYPES: THAILAND. - Chaiyaphum Province, Pa Hin Ngam NP, deciduous, 
15°39.966'N 101°27.198'E, 357 m, 19-23.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T1352) (ld, QSBG). - Pa Hin Ngam NP, deciduous, 15°40.232'N 
101°26.942'E, 398 m, 5-12.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 



NEW SPECIES OF DIPTERA FROM ASIA 669 





FIGS 1-5 

Cestrotus quadrimaculaius sp. n., maie paratype. (1) Epandrium and protandrium, latéral view. 
(2) Protandrium, anterior view. (3) Epandrium, posterior view. (4) Aedeagal complex, ventral 
view. (5) Aedeagal complex, latéral view. Scale 0. 1 mm. 

(T1345) (26 6, 2 9 9 , CSCA). - Pa Hin Ngam NP, Dry dipterocarp, 15°38.099'N 101°23.921'E, 
698 m, ll-18.viii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T445) (\6, 
BPBM). - Pa Hin Ngam NP, Dry evergreen forest at waterfall, 15°34.802'N 101°25.99'E, 
430 m, 22-28.xi.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1038) (\S, 
19, CSCA). - Pa Hin Ngam NP, Dry evergreen forest at waterfall, 15°34.802*N 101°25.99'E, 
430 m, 16-22.xi.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1035) (16", 
3 9 9, CSCA). - Pa Hin Ngam NP, Dry evergreen next to creek, 15°40.569'N 101°26.705'E, 
461 m, 19-23.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1353) (3c? o*, 
1 9 , QSBG). - Pa Hin Ngam NP, ecotone between mix deciduous and dipterocarp forest. 
15°38.132*N 101°23.922'E, 698 m, 7-13.ii.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1646) (1 S , QSBG). - Pa Hin Ngam NP, Ecotone between mixed deciduous and dry 
dipterocarp forest, 15°34.913 r N 101°25.658'E, 444 m, 16-22.xi.2006, ex. Malaise trap, coll: 
Katae Sa-nog & Buakaw Adnafai (T1034) (le?, 19, CNCI). - Pa Hin Ngam NP, Ecotone 
between mixed deciduous and dry dipterocarp forest, 15°34.913 r N 101°25.658'E, 444 m, 
10-16.xi.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1031) (1 9, USNM). 



670 



L. SHI ET AL. 




Plate A 

Head, anterior view and thorax, dorsal view. (1), (3) Cestrotus quadrimaculatus sp. n. (2), (4) 
Dioides merzi sp. n. 

- Pa Hin Ngam NP, Mixed deciduous forest, 15°34.686'N 101°26.082'E, 419 m, 22-28.xi.2006, 
ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1039) (16, CSCA). - Pa Hin Ngam 
NP, Nature trail at Lan Hin Nor, 15°37.615'N 101°23.436'E, 668 m, 15-21.iii.2007, ex. Malaise 
trap, coll: Katae Sa-nog & Buakaw Adnafai (T2331) (1 9, USNM). - Pa Hin Ngam NP, Thung 
Dok Kra Jeow in Dry evergreen, 15°38.438'N 101°23.576'E, 780 m, 7-13.L2007, ex. Malaise 
trap, coll: Katae Sa-nog & Buakaw Adnafai (T1457) (1 cî, 1 ? , BMNH). - Tat Tone NP, By the 
stream, 15°58.771'N 102°2.397*E, 305 m, 19-26.i.2007, ex. Malaise trap, coll: Tawit Jaruphan & 
Orawan Budsawong (T1564) (le?, QSBG). - Tat Tone NP, Chaiyapoom forest fire station, 
16°0.738'N 102°1.342'E, 195 m, 26.xii.2006-2.i.2007, ex. Malaise trap, coll: Tawit Jaruphan & 
Orawan Budsawong (T1377) (1 6, USNM). - Tat Tone NP, Dry Dipterocarp Forest, 15°59.037*N 
102°2.103'E, 250 m, 25-27.vi.2006, ex. Malaise trap, coll: M. Sharkey (T10) (1 ?, BPBM). -Tat 
Tone NP, Lum pa tao head water/dry evergreen, 15°58.486'N 102°2.239'E, 270 m, 19-26.ii.2007, 
ex. Malaise trap, coll: Tawit Jaruphan & Orawan Budsawong (T1733) (\â , QSBG). - Tat Tone 
NP, Phu hang sing, 15°58.723'N 102°2.231'E, 290 m, 26.i.-2.ii.2007, ex. Malaise trap, coll: 
Tawit Jaruphan & Orawan Budsawong (T1569) (lâ, QSBG). - Tat Tone NP, Streamside at Tat 
Fah waterfall, 15°56.463'N 102°5.953'E, 242 m, 26.iii.-2.iv.2007, ex. Malaise trap, coll: Tawit 
Jaruphan&Orawan Budsawong (T2355) (266, 1 9, BPBM). - Tat Tone NP, Streamside at Tat 
Fah waterfall, 15°56.463'N 102°5.953'E, 242 m, 5-12.iii.2007, ex. Malaise trap, coll: Tawit 
Jaruphan&Orawan Budsawong (T2346) (29 9, QSBG). - Chiang Mai Province, Doi Inthanon 
NP, Vachirathan Fall, 18°32.31'N 98°36.048'E, 700 m, 15-22.iv.2007, ex. Malaise trap, coll: Y. 
Areeluck (T1842) (19, CSCA).- Kamphaeng Phet Province, data same as holotype (16, 
CAUC). - Khonkaen Province, Nam Pong NP, office, 16°37.341'N 102°34.467'E, 324 m, 
19-26.vii.2006, ex. Malaise trap, coll: Khamphol Jaidee (T109) (\â, QSBG). - Nam Pong NP, 
office, 16°37.377'N 102°34.454'E, 344 m, 19-26.vii.2006, ex. Malaise trap, coll: Khamphol 
Jaidee (Tlll) (16, QSBG). - Loei Province, Phu Kradueng NP, Forest protection unit Loei .5 



NEW SPECIES OF DIPTERA FROM ASIA 



671 



(Phakbung), 16°50.54*N 101°41.663'E, 406 m, 25.ii.-l.iii.2007, ex. Malaise trap, coll: Sonkgran 
Kamtue (T1505) (2 c? c?, CSCA; 2 9 9, QSBG). - Phu Kradueng NP, Forest protection unit Loei 
.5 (Phakbung), 16°50.54*N 1 1 °4 1 .663'E, 406 m, 7-13.ii.2007, ex. Malaise trap, coll: Sutin 
Khonglasae (T1496) (le?, 1 9, QSBG; 29 9, CSCA). - Phetchabun Province, Khao Kho NP, 
Mix deciduous, 16°39.589'N 101°8.185'E, 168 m, 5-12.i.2007, ex. Malaise trap, coll: Somchai 
Chachumnan & Saink Singtong (T1392) (le?, USNM). - Khao Kho NP, Mix deciduous, 
16°39.589'N 101°8.185'E, 168 m, 19-26.i.2007, ex. Malaise trap, coll: Somchai Chachumnan & 
Saink Singtong (T1398) (le?, QSBG). - Khao Kho NP, Mix deciduous near office, 16°39.587'N 
101°8.134'E, 220 m, 26.ii.2007, ex. Malaise trap, coll: Somchai Chachumnan & Saink Singtong 
(T1608) (1 9, CNCI).- Khao Kho NP, office, 16°39.479'N 101°8.105'E, 220 m, 12-19.vii.2006, 
ex. Malaise trap, coll: Somchai Chatchumnan & Sa-ink Singtong (T166) (1 c?, USNM). - Khao 
Kho NP, View point at Khla Stream, 16°39.12'N 101°7.8rE, 246 m, 12-19.iii.2007, ex. Malaise 
trap, coll: Somchai Chachumnan & Saink Singtong (T2413) (3 9 9, QSBG; 19, BPBM). - 
Sakon Nakhon Province, Phu Phan NP, Nam Hom Waterfall # Sao Hi, 17°7.34*N 104°0.788'E, 
344 m, 25-31.iii.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T2377) (le?, 1 9, USNM). 
- Phu Phan NP, North of well, 17°3.543*N 103°58.452'E, 312 m, 23-30.i.2007, ex. Malaise trap, 
coll: Sailom Tongboonchai (T1525) (2c? c?, 19, USNM). - Phu Phan NP, West of well, 
17°3.521'N 103°58.45*E, 322 m, 5-ll.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai 
(T1517) (2c? cî, 1 9 , CNCI). - Phu Phan NP, West of well, 17°3.521'N 103°58.45*E, 322 m, 17- 
23.Ï.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1523) (1 c?, 1 9 , NHMW). - Phu Phan 
NP, West of well, 17°3.521'N 103°58.45'E, 322 m, 23-30.i.2007, ex. Malaise trap, coll: Sailom 
Tongboonchai (T1526) (1 9, QSBG). - Ubon Ratchathani Province, Pha Taem NP, Saeng 
Chan Waterfall, 15°31.985'N 105°35.774'E, 155 m, 12-20.iii.2007, ex. Malaise trap, coll: 
Porntip Tonsu & Bunlu Sapsiri (T2146) (19, QSBG). VIETNAM. - Tay Ninth Province, Nui 
Ba Den, 300 m, 11.22N/106.11E, 20-21.VIII.2003, P. Schwendinger (19, MHNG; le?, CAUC). 

ETYMOLOGY: Latin, quadri-, prefix meaning four + maculatus, meaning macu- 
late or with spots, referring to the face having a pair of pale brown subbasal spots and 
a pair of blackish brown round subapical spots; a masculine adjective. 

DiAGNOSis: Face with a pair of pale brown subbasal spots and a pair of blackish 
brown round subapical spots. Antennal lst flagellomere entirely yellow. Palpus yellow. 
Mesonotum with a pair of grayish brown médian stripes, a pair of grayish brown sub- 
lateral stripes along upper margin of sa and a pair of brownish gray latéral stripes 
behind suture. Scutellum yellow with dense yellow pruinosity. Ail legs yellow, femo- 
ra with a grayish black apicoventral spot, tibiae with a subbasal and apical black ring 
and tarsomeres 3-5 brown. Wing with a wide, brown, subapical stripe from costal 
margin to hind margin; a hyaline elliptical spot on M] présent between r-m and dm-cu; 
r-m and dm-cu brown. Abdomen brown to black with dense gray pruinosity, some 
tergites with yellow laterally. 



Description 

Length: body 3.5-4.0 mm (maie), 4.0 mm (female); wing 3.4-3.7 mm (maie), 
3.7 mm (female). 

Head (Plates A 1 , A3) pale yellow. Frons as long as wide and parallel-sided, with 
dense pale yellow pruinosity; with a pair of black velvety triangular spots, covering 
base of anterior or and a médian hump slightly projecting anteriorly; ocellar triangle 
grayish black, with 3 rows of short setulae on grayish médian stripe behind ocellar 
triangle; oc strong and longer than anterior or\ anterior or reclinate, shorter than 
posterior or. Face covered by sparse white pruinosity, with a pair of pale brown sub- 
basal spots, a pair of blackish brown subapical spots and three dark spots on ventral 



672 



L. SHI ETAL. 



margin. Gena about 1/4 height of eye, with 1 long seta and a black spot. Antennal scape 
and pedicel grayish black, lst flagellomere entirely yellow, nearly 1.8 times as long as 
high; arista brown, long plumose, with longest setula as long as height of lst flagel- 
lomere. A pale brown spot présent between antenna and eye. Proboscis yellow, except 
pale brown at tip, with yellowish and blackish setulae; palpus yellow, with blackish 
setulae. 

Thorax (Plate A3) black, with dense grayish and yellowish pruinosity. 
Mesonotum with a pair of grayish brown médian stripes, a pair of grayish brown sub- 
lateral stripes along upper margin of sa, a pair of brownish gray latéral stripes behind 
suture and a black rectangular spot on postpronotum; 0+2 de; acr in irregular 8 rows; 
prsc longer than lst postsutural de. 1 anepst, 2 kepst. Scutellum yellow, with dense 
yellow pruinosity. Halter pale yellow. 

Legs yellow, ail femora with sparse grayish pruinosity and a grayish black api- 
coventral spot, ail tibiae with a subbasal and apical black ring and tarsomeres 3-5 
brown. Fore fémur with 4-5 pv and 5-6 pd, ctenidium with 12 short setulae; fore tibia 
with 1 preapical ad and 1 short apv. Mid fémur with 5 a; mid tibia with 1 preapical ad 
and 2 strong apv. hind tibia with 1 short preapical ad and 1 short apv. 

Wing (Plate Bl) hyaline, with brown irregular spots; a wide brown subapical 
stripe extending from costal margin to hind margin; an hyaline elliptical spot on M] 
présent between r-m and dm-cu; r-m and dm-cu brown; subcostal cell hyaline, with a 
pale brown round apical spot, extending nearly to R2+3; Costa with 2nd (between R± 
and R2+3), 3rd (between R 2 +3 and R4+5) and 4th (between R 4+5 and Mj) sections in 
proportion of 1.7 mm : 0.6 mm : 0.3 mm; r-m beyond middle of distal cell; ultimate 
and penultimate sections of in proportion of 1.1 mm: 2.0 mm; ultimate section of 
CuAj about 1/13 of penultimate. 

Abdomen brown to black, with dense gray pruinosity, some tergites with yellow 
laterally. Maie genitalia (Figs 1-5): protandrium circular; epandrium broad in latéral 
view; surstylus with an acuate process, curved upward; hypandrium broad at middle 
and curved backward, hypandrial apodeme small; aedeagus with a pair of triangular 
basai processes, a deep apical incision, subuliform apically in ventral view and curved 
backward in latéral view. 

REMARKS: The new species is similar to Cestrotus pilosus (Hendel) from 
Tanzania. According to the original description by Hendel (1920) and study of the type, 
they share the following characters: face with white pruinosity and brown spots, 
antennal scape and pedicel grayish black, 1 st flagellomere yellow, mesonotum with acr 
in 8 rows, wing broadly brown in apical third. But it can be separated from the latter 
by the pruinose brown stripe behind the ocellar triangle flanked on both sides by solid 
yellow pruinose upper frons, the mesonotum having a pair of grayish brown médian 
stripes and a pair of short brownish gray latéral stripes separated completely, the scu - 
tellum having grayish yellow pruinosity, and the distal margin of the wing being 
hyaline. In Cestrotus pilosus, the area behind the ocellar triangle is concolorous with 
the upper frons, the mesonotum has four brown stripes confluent on the anterior half 
and isolated on the posterior half, the scutellum is bright yellow pruinose, the distal 
margin of the wing is completely brown with no marginal hyaline area, and other 
détails of the wing differ. 



NEW SPECIES OF DIPTERA FROM ASIA 



673 



Distribution: Thailand, Vietnam. 
Cestrotus flavoscutellatus Meijere, 1910 

Cestrotus flavoscutellatus Meijere, 1910: 142. Type locality: Indonesia (Java). 

MATERIAL: THAILAND. - Loei Province, Phu Ruea NP, Nern Wibaak ditch, 
17°29.907'N 101°20.483'E, 1196 m, 19-26.ii.2007, ex. Malaise trap, coll: Patikhom Tumtip 
(T1711)(lct,QSBG). 

Distribution: China (Guangxi, Hainan, Hunan, Taiwan), Laos, Népal, 
Indonesia (Java), Malaysia (Sabah), Vietnam, new record to Thailand. 

Cestrotus liui Shi, Yang & Gaimari, 2009 

Cestrotus liui Shi, Yang & Gaimari, 2009: 57. Type locality: China (Yunnan). 

MATERIAL: MALAYSIA. - Pahang, Bukit Fraser, Maxwell Trail, 1300 m, 3.43N/ 
101.44E, 12-16.V.2004, P. Schwendinger (2 et et, 19, MHNG; lct, CAUC). THAILAND. - 
Loei Province, Phu Kradueng NP, Savannah in pine forest, 16°53.092'N 101°47.413'E, 1257 m, 
9-16.i.2007, ex. Malaise trap, coll: Thanongsak Srisa-ad (T1226) (1 et, QSBG). 

Distribution: China (Hainan, Yunnan), new records to Malaysia and Thailand. 
Cestrotus heteropterus Shi, Yang & Gaimari, 2009 

Cestrotus heteropterus Shi, Yang & Gaimari, 2009: 54. Type locality: China (Yunnan). 

MATERIAL: MALAYSIA. - Kelantan, Gua Keris, Gua King Kong, South Pahang, 
130 m, 52.1N/102.01E, 8-9.VI.2004, P. Schwendinger (le?, MHNG). THAILAND. - 
Chaiyaphum Province, Pa Hin Ngam NP, Car park at Thung Dok Kra Jeow, 15°38.391'N 
101°23.609'E, 750 m, 7-13.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1456) (3 et et, 19, USNM). - Pa Hin Ngam NP, Car park at Thung Dok Kra Jeow, 
15°38.391'N 101°23.609'E, 750 m, 19-25.i.2007, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T1462) (1 9, QSBG). - Pa Hin Ngam NP, deciduous, 15°39.966'N 
101°27.198'E, 357 m, ll-12.xii.2006, ex. pan trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1344) (1 9, CNCI). - Pa Hin Ngam NP, Dry evergreen next to creek, 15°40.569'N 
101°26.705'E, 461 m, 19-23.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1353) (1 9, CSCA). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dipterocarpus forest, 
15°38.208'N 101°23.556'E, 720 m, 19-2512007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1464) (3 S et , 2 9 9 , QSBG; 1 9 , BPBM). - Pa Hin Ngam NP, Thung Dok Kra Jeow 
in Dipterocarpus forest, 15°38.208'N 101°23.556'E, 720 m, 7-13.i.2007, ex. Malaise trap, coll: 
Katae Sa-nog & Buakaw Adnafai (T1458) (3 â et , QSBG). - Pa Hin Ngam NP, Thung Dok Kra 
Jeow in Dry evergreen, 15°38.438'N 101°23.576'E, 780 m, 3-412007, ex. pan trap, coll: Katae 
Sa-nog & Buakaw Adnafai (T1448) (1 et, CSCA). - Pa Hin Ngam NP, Thung Dok Kra Jeow in 
Dry evergreen, 15°38.438'N 101°23.576'E, 780 m, 4-5.i.2007, ex. pan trap, coll: Katae Sa-nog 
& Buakaw Adnafai (T1449) (1 et, 1 9 , USNM). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry 
evergreen, 15°38.438'N 101°23.576'E, 780 m, 19-25.i.2007, ex. Malaise trap, coll: Katae Sanog 
& Buakaw Adnafai (T1463) (1 et, QSBG). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry ever- 
green, 15°38.438'N 101°23.576'E, 780 m, 2-3.i.2007, ex. pan trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1447) (1 et, 3 9 9, CSCA). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry ever- 
green, 15°38.438*N 101°23.576'E, 780 m, 5-6.i.2007, ex. pan trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1450) (19, QSBG). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry evergreen, 
15°38.438'N 101°23.576'E, 780 m, 7-1312007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1457) (2 et et, 39 9, QSBG). - Tat Tone NP, Dry Dipterocarp Forest, 15°59.037'N 
102°2.103'E, 250 m, 25-27.vi.2006, ex. Malaise trap, coll: M. Sharkey (T10) (1 et, QSBG). - Tat 
Tone NP, Dry dipterocarp forest near swamp at Sab somboon forest unit, 16°1.059'N 
101°58.603'E, 674 m, 26.xi.-3.xii.2006, ex. Malaise trap, coll: Tawit Jaruphan (Tl 146) (19, US- 
NM). - Tat Tone NP, Lum pa tao head water/dry evergreen, 15°58.486'N 102°2.239'E, 270 m, 
19-26. ii. 2007, ex. Malaise trap, coll: Tawit Jaruphan & Orawan Budsawong (T1733) (29 9, 
USNM). - Tat Tone NP, Next to Pa Eang waterfall, 15°57.657'N 101°54.724'E, 301 m, 



674 



L. SHI ETAL. 



19-26.X.2006, ex. Malaise trap, coll: Tawit Jaruphan (T685) (le?, QSBG). - Tat Tone NP, 
Streamside at Tat Fah waterfall, 15°56.463'N 102°5.953*E, 242 m, 5-12.iii.2007, ex. Malaise 
trap, coll: Tawit Jaruphan&Orawan Budsawong (T2346) (le?, USNM). - Chiang Rai Province, 
Doi Inthanon NP, summit marsh, 18°35.361'N 98°29.157'E, 2500 m, 12-19.X.2006, ex. Malaise 
trap, coll: Y. Areeluck (T368) (le?, QSBG). - Mae Si District, Doi Tung, 1300 m, 
20.19.28N/99.49.53E, 17.XII.2003, P. Schwendinger (3 c? c?, MHNG). - Chumphon Province, 
Lang Suan District, Khao Kai Jae. waterf., 80 m, 9.55N/98.57E, 17-18.VII.2002, P. 
Schwendinger (6 c? c? , 2 9 9 , MHNG). - Loei Province, Phu Kradueng NP, Forest protection unit 
Loei .5 (Phakbung), 16°50.493'N 101°41.726'E, 412 m, 7-13.ii.2007, ex. Malaise trap, coll: 
Sutin Khonglasae (T 1 497) (4 c? c? , 4 9 9 , QSBG; 2 c? c? , 1 9 , CSC A; 2 c? c? , 1 9 , USNM; 2 c? c? , 
19,BMNH;2c?c?, 1 9 , BPBM; 2c? c?, 1 9 , CNCI; 2c? c?, 1 9 , MHNG; 2c? c?, 1 9 , MZSP; 2c? c?, 
19, NHMW; 2c?c?, 19, NMSA). - Phu Kradueng NP, Forest protection unit Loei .5 
(Phakbung), 16°50.54*N 101°41.663'E, 406 m, 13-19.ii.2007, ex. Malaise trap, coll: Wuthicahi 
kwanjam (T1499) (4c? c?, 49 9, CSCA). - Phu Kradueng NP, Forest protection unit Loei .5 
(Phakbung), 16°50.54'N 101°41.663'E, 406 m, 13-19.ii.2007, ex. Malaise trap, coll: Wuthicahi 
kwanjam (T1499) (3c? c?, 29 9, QSBG; 2c? c?, CSCA; 19, BMNH; 19, MHNG; 19, MZSP; 
1 9 , NHMW; 1 9 , NMSA). - Phu Kradueng NP, Forest protection unit Loei .5 (Phakbung), 
16°50.54'N 101°41.663'E, 406 m, 7-13.ii.2007, ex. Malaise trap, coll: Sutin Khonglasae (T1496) 
(3 c? S , 1 9 , USNM; 5 c? c? , CSCA; 1 9 , BPBM; 1 9 , CNCI). - Phu Kradueng NP, Hill evergren 
forest at Wang Gwang forest unit, 16°53.362'N 101°47.286'E, 1262 m, 3-9.i.2007, ex. Malaise 
trap, coll: Thanongsak Srisa-ad (T1222) (le?, CSCA). - Phu Kradueng NP, Savannah in pine 
forest, 16°53.092'N 101°47.413'E, 1257 m, 9-16.L2007, ex. Malaise trap, coll: Thanongsak 
Srisa-ad (T1226) (2c? c?, CNCI). - Phu Ruea NP, Nature tail, 17°30.74'N 101°20.65'E, 1353 m, 
5-12.iii.2007, ex. Malaise trap, coll: Patikhom Tumtip (T2305) (1 9, QSBG). - Phetchabun 
Province, Khao Kho NP, Mixed deciduous forest, 16°32.539'N 101°2.483'E, 524 m, 11- 
12.xii.2006, ex. pan trap, coll: Somchai Chachumnan & Saink Singtong (T1177) (1 9, QSBG). 

- Phitsanulok Province, Thung Salaeng Luang NP, Dry Evergreen forest, 16°50.277'N 
100°52.917'E, 486 m, 18-25.iii.2007, ex. Malaise trap, coll: Pongpitak & Pranee & Sathit 
(T2388) (1 9, BPBM). - Sakon Nakhon Province, Phu Phan NP, Behind forest protection unit at 
Huay Wien Prai, 17°6.81'N 104°0.318'E, 318 m, 25.ii.-3.iii.2007, ex. Malaise trap, coll: Sailom 
Tongboonchai (T1693) (1 c?, 1 9 , CSCA). - Phu Phan NP, Behind forest protection unit at Huay 
Wien Prai, 17°6.847'N 104°0.302'E, 376 m, 4-10.ii.2007, ex. Malaise trap, coll: Winlon 
Kongnara (T1685) (le?, BPBM). - Phu Phan NP, Dry evergreen forest near house at 1567 
station, 16°48.628'N 103°53.591'E, 522 m, 22-29.xii.2006, ex. Malaise trap, coll: S. Kongnara 
(T1250) (1 9, QSBG). - Phu Phan NP, Nam Hom Waterfall # Sao Hi, 17°7.34'N 104°0.788'E, 
344 m, 10-17.iii.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T2371) (1 9, CNCI). - Phu 
Phan NP, North of well, 17°3.543'N 103°58.452'E, 312 m, 5-11.L2007, ex. Malaise trap, coll: 
Sailom Tongboonchai (T1516) (19, QSBG). - Phu Phan NP, West of well, 17°3.521'N 
103°58.45'E, 322 m, ll-17.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1520) (le?, 
BPBM). - Phu Phan NP, West of well, 17°3.521'N 103°58.45'E, 322 m, 5-1 l.i.2007, ex. Malaise 
trap, coll: Sailom Tongboonchai (T1517) (2e? e?, 3 9 9 , USNM). - Phu Phan NP, West of well, 
17°3.521'N 103°58.45'E, 322 m, 23-30.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai 
(T1526) (1 9, QSBG). - Phu Phan NP, West of well, 17°3.521'N 103°58.45'E, 322 m, 17- 
23.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1523) (2c? c?, 19, QSBG).- Sunat 
Thanai Province, Khao Sok National Park, 100 m, 8.54.55N/98.31.39E, 12-14.V.2003, 
P. Schwendinger (le?, MHNG). - Trat Province, Ko Chang, 12.03N/102.18E, 17-18.VII.2002, 
P. Schwendinger (19, MHNG). VIETNAM. - Dong Nai Province, NW Tan Phu, 130 m, 
11.25N/107.26E, 26-29.VIII.2003, P. Schwendinger (19, MHNG). - Lam Dong Province, W 
Di Linh, Bo Bia Wat. Fall, 860 m, 1 1.34N/ 108.01E, 03.IX.2003, P. Schwendinger (19, CAUC). 

- Tay Ninh Province, Tay Ninh, Nui Ba Den, 300 m, 11.22N/106.11E, 20-21.VIII.2003, P. 
Schwendinger (8 e? c? , 3 9 9 , MHNG; 1 c? , CAUC). 

Distribution: China (Yunnan), Thailand, new records to Malaysia and 
Vietnam. 

Dioides merzi sp. n. Figs 6-10, Plates A2, A4, B2 

MATERIAL: HOLOTYPE e? (MHNG): Vietnam, Tay Ninh Province, Tay Ninh, Nui Ba 
Den, 300 m, 11.22N/106.11E, 20-21.VIII.2003, P. Schwendinger. PARATYPES: THAILAND. - 



NEW SPECIES OF DIPTERA FROM ASIA 



675 




FIGS 6-10 

Dioides merzi sp. n., maie paratype. (6) Epandrium and protandrium, latéral view. (7) 
Protandrium, anterior view. (8) Epandrium, posterior view. (9) Aedeagal complex, ventral view. 
(10) Aedeagal complex, latéral view. Scale 0.1 mm. 



676 



L. SHI ETAL. 



Chaiyaphum Province, Pa Hin Ngam NP, Car park at Thung Dok Kra Jeow, 15°38.391'N 
101°23.609'E, 750 m, 7-13.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1456) (lcî, 1 9 , USNM). - Pa Hin Ngam NP, deciduous, 15°39.966'N 101°27.198'E, 357 m, 
19-23.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1352) {566, 5? 9, 
CSC A- 26 S, 29 9, BMNH; 5 9 9, USNM). - Pa Hin Ngam NP, deciduous, 15°40.232'N 
101°26.942'E, 398 m, 5-12.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1345) (1 9, USNM). - Pa Hin Ngam NP, Dry dipterocarp, 15°38.099'N 101°23.921'E, 698 m, 

1 l-18.viii.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T445) (1 9, USNM). 
- Pa Hin Ngam NP, Dry evergreen forest (Thepana watrfall), 15°38.884'N 101°25.84'E, 605 m, 
1-7.X.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T658) (\6, QSBG). - Pa 
Hin Ngam NP, Dry evergreen forest at Lan Hin Nor, 15°37.54'N 101°23.313'E, 673 m, 
21-27.iii.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T2335) (1 9, QSBG). - 
Pa Hin Ngam NP, Dry evergreen forest at Lan Hin Nor, 15°37.54'N 101°23.313'E, 673 m, 
9-15.iii.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T2329) (366, 39 9, 
QSBG). - Pa Hin Ngam NP, Dry evergreen forest at Lan Hin Nor, 15°37.54'N 101°23.313'E, 673 
m, 15-21.iii.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T2332) (26 6, 

2 9 9, QSBG; 1 6 , 2 9 9 , MHNG; 29 9, CNCI). - Pa Hin Ngam NP, Dry evergreen forest at 
waterfall, 15°34.802'N 101°25.99'E, 430 m, 16-22.xi.2006, ex. Malaise trap, coll: Katae Sa-nog 
& Buakaw Adnafai (T1035) (66 6, 3 9 9, CSCA). - Pa Hin Ngam NP, Dry evergreen forest at 
waterfall, 15°34.802'N 101°25.99'E, 430 m, 22-28.xi.2006, ex. Malaise trap, coll: Katae Sa-nog 
& Buakaw Adnafai (T1038) (26 6, 2 9 9 , CSCA). - Pa Hin Ngam NP, Dry evergreen forest at 
waterfall, 15°34.802'N 101°25.99'E, 430 m, 10-16.xi.2006, ex. Malaise trap, coll: Katae Sa-nog 
& Buakaw Adnafai (T1032) (1 9, CSCA). - Pa Hin Ngam NP, Dry evergreen next to creek, 
15°40.569'N 101°26.705'E, 461 m, 19-23.xii.2006, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T1353) (266, 39 9, QSBG; 266, 39 9, CSCA). - Pa Hin Ngam NP, 
Ecotone between mixed deciduous and dry dipterocarp forest, 15°34.913'N 101°25.658'E, 
444 m, 10-16.xi.2006, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1031) (16, 
2 9 9, CSCA). - Pa Hin Ngam NP, Ecotone between mixed deciduous and dry dipterocarp 
forest, 15°34.913'N 101°25.658'E, 444 m, 16-22.xi.2006, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T 1 034) (6 6 6 , 8 9 9 , CSCA; 2 6 6 , 2 9 9 , QSBG). - Pa Hin Ngam NP, Mixed 
deciduous forest, 15°34.686'N 101°26.082'E, 419 m, 22-28.xi.2006, ex. Malaise trap, coll: Katae 
Sa-nog & Buakaw Adnafai (T1039) (16,29 9, CSCA). - Pa Hin Ngam NP, Nature trail at Lan 
Hin Nor, 15°37.615'N 101°23.436'E, 668 m, 15-21.iii.2007, ex. Malaise trap, coll: Katae Sa-nog 
& Buakaw Adnafai (T233 1) (2 6 6 , 4 9 9 , QSBG; \6,69 9, CSCA; 1 6 , 6 9 9 , USNM). - Pa 
Hin Ngam NP, Thung Dok Kra Jeow in Dipterocarpus forest, 15°38.208'N 101°23.556'E, 720 m, 
7-13.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1458) (36 6, 19, 
CSCA). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dipterocarpus forest, 15°38.208'N 
101°23.556'E, 720 m, 13-1912007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai 
(T1461) (1 6, 1 9 , USNM). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dipterocarpus forest, 
15°38.208'N 101°23.556'E, 720 m, l-7.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1455) (16, NMSA). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dipterocarpus 
forest, 15°38.208'N 101°23.556'E, 720 m, 19-25.i.2007, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T1464) (266,39 9, QSBG; 266,29 9, NMSA; 26 6, 1 9, CASC). - Pa 
Hin Ngam NP, Thung Dok Kra Jeow in Dry evergreen, 15°38.438'N 101°23.576'E, 780 m, 
19-25.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1463) (\6, QSBG). - 
Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry evergreen, 15°38.438'N 101°23.576'E, 780 m, 
7-13.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw Adnafai (T1457) (266, 39 9, 
MZSP; 26 6, 29 9, ZISP). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry evergreen, 
15°38.438'N 101°23.576'E, 780 m, 13-19.i.2007, ex. Malaise trap, coll: Katae Sa-nog & Buakaw 
Adnafai (T1460) (3 6 6 , 3 9 9 , QSBG). - Pa Hin Ngam NP, Thung Dok Kra Jeow in Dry ever- 
green, 15°38.438'N 101°23.576'E, 780 m, 7-13.L2007, ex. Malaise trap, coll: Katae Sa-nog & 
Buakaw Adnafai (T1457) (5 6 6 , 4 9 9 , BPBM; 3 9 9, QSBG; 39 9, USNM). - Tat Tone NP, 
By the creek at Takfah waterfall, 15°56.463'N 102°5.953'E, 242 m, 26.ix.-3.x.2006, ex. Malaise 
trap, coll: Tawit Jaruphan & Orawan Budsawong (T875) (1 9, MZSP). - Tat Tone NP, By the 
stream, 15°58.538'N 102°2.153'E, 280 m, 12-19.i.2007, ex. Malaise trap, coll: Tawit Jaruphan & 
Orawan Budsawong (T1562) (16, QSBG). - Tat Tone NP, By the stream, 15°58.771'N 
102°2.397'E, 305 m, 12-19.i.2007, ex. Malaise trap, coll: Tawit Jaruphan & Orawan Budsawong 




Plate B 

Wing. (1) Cestrotus quadrimaculatus sp. n. (2) Dioides merzi sp. n. (3) Noonamyia bipunctata 
sp. n. (4) Noonamyia flavoscutellata sp. n. 



678 



L. SHI ETAL. 



(T1561) (1 ?,ZISP). -Tat Tone NP,Chaiyapoom forest fire station, 16°0.738'N 102°1.342'E, 195 
m, 19-23.xii.2006, ex. Malaise trap, coll: Tawit Jaruphan & Orawan Budsawong (T1374) (lcî, 
USNM). - Tat Tone NP, Chaiyapoom forest fire station, 16°0.738'N 102°1.342'E, 195 m, 
26. xii. 2006-2. i. 2007, ex. Malaise trap, coll: Tawit Jaruphan & Orawan Budsawong (T1377) (1 6 , 
QSBG). - Tat Tone NP, Dry Dipterocarp Forest, 15°59.037'N 102°2.103'E, 250 m, 
21-28.vi.2006, ex. Malaise trap, coll: Vinlon Khongnara (T19) (1 9 , BPBM). - Tat Tone NP, Dry 
Dipterocarp Forest, 15°59.037'N 102°2.103'E, 250 m, 28.vi.2006, ex. Malaise trap, coll: Pong 
Sandow (T18) (1 9, ZISP). - Tat Tone NP, Dry Dipterocarp Forest, 15°59.037'N 102°2.103'E, 
250 m, 25-27. vi.2006, ex. Malaise trap, coll: M. Sharkey (T10) (1 9 , CASC). - Tat Tone NP, Dry 
dipterocarp forest near swamp at Sab somboon forest unit, 16°1.059'N 101°58.603'E, 674 m, 
26.xi.-3.xii.2006, ex. Malaise trap, coll: Tawit Jaruphan (T1146) (19, QSBG). - Tat Tone NP, 
Dry dipterocarp forest near swamp at Sab somboon forest unit, 16°1.059'N 101°58.603'E, 674 
m, 12-19.xi.2006, ex. Malaise trap, coll: Tawit Jaruphan (T1140) (1 9, QSBG). - Tat Tone NP, 
Nursery near head water, 15°58.344'N 102°2.169'E, 257 m, 5-12.viii.2006, ex. Malaise trap, 
coll: Tawit Jaruphan & Orawan Budsawong (T547) (1 9, QSBG). - Tat Tone NP, Phu hang sing 
, 15°58.723'N 102°2.231*E, 290 m, 26.i.2.ii.2007, ex. Malaise trap, coll: Tawit Jaruphan & 
Orawan Budsawong (T1569) (1 9, CASC). - Tat Tone NP, Staff house at Takfah waterfall, 
15°56.461'N 102°5.955'E, 242 m, 19-26.ix.2006, ex. Malaise trap, coll: Tawit Jaruphan & 
Orawan Budsawong (T873) (1 9, BPBM). - Chiang Mai Province, Doi Inthanon NP, summit 
marsh, 18°35.361'N 98°29.157'E, 2500 m, 12-19.X.2006, ex. Malaise trap, coll: Y. Areeluck 
(T368) (1 9, QSBG). - Loei Province, Phu Kradueng NP, Forest protection unit Loei .5 
(Phakbung), 16°50.54'N 101°41.663'E, 406 m, 7-13.ii.2007, ex. Malaise trap, coll: Sutin 
Khonglasae (T1496) (le?, QSBG). - Phu Kradueng NP, Koke Hin Ngam, 16°51.817'N 
101°50.704'E, 270 m, 9-16.viii.2006, ex. Malaise trap, coll: Sutin Khonglasae (T483) (\6, 
QSBG). - Phu Kradueng NP, Mixed deciduous forest north of Na Noy Forest Unit, 16°48.17'N 
101°47.666'E, 276 m, 2-8.xii.2006, ex. Malaise trap, coll: Suthin Gong-lasae (T1084) (1 9, 
BPBM). - Phu Kradueng NP, Mixed deciduous forest south of Na Noy Forest Unit, 16°49.099'N 
101°47.624*E, 275 m, 14-20.xi.2006, ex. Malaise trap, coll: Suthin Gong-lasae (T1074) (26 6, 

2 9 9, CSCA; le?, 19, USNM). - Phu Kradueng NP, Mixed deciduous forest south of Na Noy 
Forest Unit, 16°49.099'N 101°47.624'E, 275 m, 18-19.xi.2006, ex. pan trap, coll: Suthin Gong- 
lasae (T1070) (1 9, QSBG). - Phu Kradueng NP, Mixed deciduous forest south of Na Noy Forest 
Unit, 16°49.099'N 101°47.624'E, 275 m, 20-26.xi.2006, ex. Malaise trap, coll: Daorueng 
Sinhpreecha (T1077) (\6, CSCA). - Phu Kradueng NP, Road to Ta Krong waterfall of Na Noy 
Forest Unit, 16°48.913'N 101°47.634'E, 265 m, 14-20.xi.2006, ex. Malaise trap, coll: Suthin 
Gong-lasae (T1073) (36 6, USNM). - Phu Ruea NP, Nern Pitsawong, 17°29.676*N 
101°21.093'E, 1168 m, 5-12.xi.2006, ex. Malaise trap, coll: Patikhom Tumtip (T1117) (IcT, 
BPBM). - Phu Ruea NP, Nern Wibaak ditch, 17°29.907'N 101°20.483'E, 1196 m, 26.viii.- 
2.ix.2006, ex. Malaise trap, coll: Nukoonchai Jaroenchai (T533) (le, CNCI). - Phu Ruea NP, 
Nern Wibaak ditch, 17°29.907'N 101°20.483'E, 1196 m, 19-26.ii.2007, ex. Malaise trap, coll: 
Patikhom Tumtip (T1711) (\6, QSBG). - Phu Ruea NP, Reservior, 17°28.826'N 101°21.33'E, 
931 m, 5-12.L2007, ex. Malaise trap, coll: Patikhom Tumtip (T1538) (\6, NHMW). - 
Phetchabun Province, Khao Kho NP, Mix deciduous, 16°39.572'N 101°8.194'E, 171 m, 
12-19.i.2007, ex. Malaise trap, coll: Somchai Chachumnan & Saink Singtong (T1394) (\6, 

3 9 9, CSCA). - Khao Kho NP, Mixed deciduous forest at Ta Phol river, 16°32.539'N 
101°2.483'E, 242 m, 26.xi.-2.xii.2006, ex. Malaise trap, coll: Somchai Chachumnan & Saink 
Singhtong (T978) (\6, MHNG). - Khao Kho NP, Mixed deciduous forest at Ta Phol river, 
16°32.539'N 101°2.483'E, 242 m, 10-ll.xi.2006, ex. pan trap, coll: Somchai Chachumnan & 
Saink Singhtong (T966) (1 9, NHMW). - Khao Kho NP, Mixed deciduous forest at Ta Phol 
river, 16°32.539'N 101°2.483'E, 242 m, 12-19.xi.2006, ex. Malaise trap, coll: Somchai 
Chachumnan & Saink Singhtong (T972) (1 9, USNM). - Khao Kho NP, Mixed deciduous forest 
at Ta Phol river, 16°32.539'N 101°2.483'E, 242 m, 9-10.xi.2006, ex. pan trap, coll: Somchai 
Chachumnan & Saink Singhtong (T965) (16, 1 9, CASC). - Khao Kho NP, Mixed deciduous 
forest at Ta Phol river, 16°32.539"N 101°2.483'E, 242 m, 19-26.xi.2006, ex. Malaise trap, coll: 
Somchai Chachumnan & Saink Singhtong (T975) (16, QSBG). - Khao Kho NP, Mixed deci - 
duous forest at Ta Phol river, 16°32.561'N 101°2.479'E, 242 m, 12-19.xi.2006, ex. Malaise trap, 
coll: Somchai Chachumnan & Saink Singhtong (T971) (\6, 19, CSCA). - Khao Kho NP, 
Mixed deciduous forest at Ta Phol river, 16°32.561'N 101°2.479'E, 242 m, 5-12.xi.2006, ex. 



NEW SPECIES OF DIPTERA FROM ASIA 



679 



Malaise trap, coll: Somchai Chachumnan & Saink Singhtong (T968) (2c? c?, USNM). - Khao 
Kho NP, Nursery, 16°52.573'N 101°8.077'E, 520 m, 26.x.-2.xi.2006, ex. Malaise trap, coll: 
Somchai Chatchumnan and Sa-ink Singtong (T812) (le?, USNM). - Khao Kho NP, Nursery, 
16°52.581'N 101°8.06'E, 520 m, 12-19.X.2006, ex. Malaise trap, coll: Somchai Chatchumnan 
and Sa-ink Singtong (T808) (le?, QSBG). - Khao Kho NP, Savana at nursery, 16°52.568'N 
101°8.104'E, 520 m, 6-7. x. 2006, ex. pan trap, coll: Somchai Chatchumnan and Sa-ink Singtong 
(T797) (3 9 9, USNM). - Khao Kho NP, Thanthip waterfall, 16°39.087'N 101°7.777'E, 210 m, 

5- 12. ix. 2006, ex. Malaise trap, coll: Somchai Chachumnan & Saink Singtong (T589) (1 9, 
BMNH). - Nam Nao NP, Tham Pra Laad ForestUnit, 16°44.963'N 101°27.833*E, 71 1 m,4-5.vi- 
ii.2006, ex. pan trap, coll: Noopean Hongyothi (T417) (le?, CSCA). - Nam Nao NP, Tham Pra 
Laad Forest Unit, 16°44.986'N 101°27.874'E, 711 m, 31.vii.-7.viii.2006, ex. Malaise trap, coll: 
Noopean Hongyothi (T422) (19, USNM). - Nam Nao NP, Tham Pra Laad Forest Unit, 
16°44.999'N 101°27.804'E, 715 m, 21-28.viii.2006, ex. Malaise trap, coll: Leng Janteab (T429) 
(le?, QSBG). - Phitsanulok Province, Thung Salaeng Luang NP, Moist evergreen, 1 6°50.64 1 'N 
100°52.894'E, 557 m, 25.viii.-l.ix.2006, ex. Malaise trap, coll: Pongpitak Pranee (T572) (le?, 
CSCA). - Sakon Nakhon Province, Phu Pha Yon NP, Channel, 16°55.639'N 104°10.748'E, 295 
m, 17-23. vii. 2006, ex. Malaise trap, coll: Manop Ngoyjansri & Châtrée Cheaukamjan (T295) 
(19, CSCA). - Phu Pha Yon NP, Réservoir, 16°55.655'N 104°10.658'E, 280 m, 23-29.vii.2006, 
ex. Malaise trap, coll: Manop Ngoyjansri & Châtrée Cheaukamjan (T299) (19, QSBG). - Phu 
Phan NP, Behind forest protection unit at Huay Wien Prai, 17°6.81'N 104°0.318'E, 318 m, 25. ii.- 
3.iii.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1693) (1 c?, 3 9 9, QSBG). - Phu Phan 
NP, Behind forest protection unit at Huay Wien Prai, 17°6.847'N 104°0.302'E, 376 m, 
17-25.ii.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1691) (le?, 1 9 9, CSCA; 29 9, 
QSBG). - Phu Phan NP, Behind forest protection unit at Huay Wien Prai, 17°6.863*N 
104°0.327'E, 387 m, 10-17.ii.2007, ex. Malaise trap, coll: Winlon Kongnara (T1689) H 9, 
BPBM). - Phu Phan NP, Behind Huay Wien Prai Forest Unit, 17°6.863'N 104°0.327'E, 387 m, 
8-15.viii.2006, ex. Malaise trap, coll: Vinlon Khongnara (T508) (le?, QSBG). - Phu Phan NP, 
Behind Huay Wien Prai Forest Unit, 17°6.863'N 104°0.327'E, 387 m, 2-8.viii.2006, ex. Malaise 
trap, coll: Vinlon Khongnara (T505) (1 9, CNCI). - Phu Phan NP, Behind Huay Wien Prai Forest 
Unit, 17°6.863'N 104°0.327*E, 387 m, 2-8.viii.2006, ex. Malaise trap, coll: Vinlon Khongnara 
(T505) (le?, QSBG). - Phu Phan NP, car park at Kam hom waterfall, 17°7.411'N 104°1.016'E, 
347 m, 10-16.ix.2006, ex. Malaise trap, coll: Sailom Tongboonchai (T611) (1 9, QSBG). - Phu 
Phan NP, Creek at northend of Huay Nam Pung Forest Unit, 16°55.789'N 103°54.158'E, 305 m, 

6- 13.xi.2006, ex. Malaise trap, coll: Sailom Tongboonchai (T1096) (19, MHNG). - Phu Phan 
NP, Kam Hom waterfall at Haew Sin chai, 17°7.415'N 104°1.179'E, 347 m, 10-16.ix.2006, ex. 
Malaise trap, coll: Sailom Tongboonchai (T613) (19, QSBG). - Phu Phan NP, Nam Hom 
Waterfall # Sao Hi, 17°7.34'N 104°0.788'E, 344 m, 25-31.iii.2007, ex. Malaise trap, coll: Sailom 
Tongboonchai (T2377) (1 9, QSBG). - Phu Phan NP, North of well, 17°3.543'N 103°58.452'E, 
312 m, 1 l-17.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1519) (3c? c?, 3 9 9 , CNCI; 
1 c?, 5 9 *, QSBG; 3 c? c?, MHNG). - Phu Phan NP, North of well, 17°3.543'N 103°58.452'E, 312 
m, 23-30.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1525) (2c? c?, 59 9, USNM; 
2c? c?, 29 9, QSBG; 2c? c?, 29 9, CSCA). - Phu Phan NP, North of well, 17°3.543'N 
103°58.452'E, 312 m, 5-1 l.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai (T1516) (14c? c?, 
69 9, USNM; 3c? c?, 39 9, CSCA; 3c? c?, 29 9, QSBG). - Phu Phan NP, West of well, 
17°3.521'N 103°58.45'E, 322 m, 23-30.i.2007, ex. Malaise trap, coll: Sailom Tongboonchai 
(T1526) (3c? c?, 119 9, QSBG; 26 6, 49 9, MHNG; 26 6, 29 9, NHMW; le?, MZSP; \ 6, 
ZISP). - Phu Phan NP, West of well, 17°3.521'N 103°58.45'E, 322 m, 17-23.È2007, ex. Malaise 
trap, coll: Sailom Tongboonchai (T1523) (8c? c?, 159 9, USNM; 3e? c?, 39 9, CSCA; 36 6, 
39 9, QSBG). - Phu Phan NP, West of well, 17°3.521'N 103°58.45'E, 322 m, 5-ll.i.2007, ex. 
Malaise trap, coll: Sailom Tongboonchai (T1517) (4 c? c?, 2 9 9, QSBG; 4e? c?, 2 9 9, BPBM; 
4c? c?, 29 9, CSCA; 4c? c?, 29 9, CNCI; 4c? e?, 29 9, USNM; 26 6, 29 9, BMNH; 2c? c?, 
29 9, MHNG; 2e? c?, 29 9, NHMW; 2e? c?, 29 9, NMSA; 2e? c?, 19, CASC; 26 6, 19, 
MZSP; 2 c? c?, 1 9 , ZISP). - Ubon Ratchathani Province, Pha Taem NP, East of Thung Luang 
in Dipterocarpus forest, 15°39.989'N 105°30.468'E, 238 m, 14-21. i.2007, ex. Malaise trap, coll: 
Thongkam & Pakdee (T1481) (ld, BMNH). - Pha Taem NP, Don Huay Can, 15°40.016'N 
105°30.502'E, 246 m, ll-18.xii.2006, ex. Malaise trap, coll: Thongcome & Pakdee (T1202) 
(2c? c?, 29 9, QSBG). - Pha Taem NP, Don Huay Can, 15°40.016*N 105°30.502*E, 246 m, 



680 



L. SHI ET AL. 



25.xii.2006- 1 .i.2007, ex. Malaise trap, coll: Thongcome & Pakdee (T1208) (1 9, QSBG). - Pha 
Taem NP, Don Huay Sa-nhom. 15°27.435'N 105°34.838'E, 238 m, 23.v.-30.v.2007, ex. Malaise 
trap. coll: Soravvit Mingman (T2193) (19, QSBG). - Pha Taem NP, First tier of Huay Sa Nhom 
waterfall, 15°27.407'N 105°34.867'E, 230 m, 4-ll.xi.2006, ex. Malaise trap, coll: Sorawit and 
Thongdee (T1053) (le?, QSBG). - Pha Taem NP. First tier of Huay Sa Nhom waterfall, 
15°27.407'N 105°34.867'E, 230 m. 25.xi.-2.xii.2006, ex. Malaise trap, coll: Sorawit and 
Thongdee (T1062) (le?, CSCA). - Pha Taem NP, Foot of Phu Kra jeaw, 15°39.989'N 
105°30.468'E. 238 m, 4-ll.xii.2006, ex. Malaise trap. coll: Thongcome & Pakdee (T1200) 
(3c? e?, 19, QSBG). - Pha Taem NP, Huay Pok waterfall, 15°37.321'N 105°36.982'E, 419 m, 
27.x.-3.xi.2006, ex. Malaise trap, coll: Pornthip Tonsu (T730) (1 9, NMSA). - Pha Taem NP, 
Huay Pok waterfall. 15°37.321'N 105°36.982'E, 419 m, 4-1 l.iv.2007, ex. Malaise trap, coll: 
Bunîu Sapsiri (T2165) (le?, CNCI). - Pha Taem NP. Huay Pok waterfall, 15°37.321*N 
105°36.982*E, 419 m, 13-20.X.2006, ex. Malaise trap, coll: Pornthip Tonsu (T724) (1 9, QSBG). 

- Pha Taem NP. Irrigation area west of Huay Pok forest unit, 15°37.321'N 105°36.982'E, 419 m, 
6-13.X.2006, ex. Malaise trap, coll: unknown (T719) (1 e?, 2 9 9, QSBG). - Pha Taem NP, Kua 
nang née. 15°24.258'N 105°30.855'E, 193 m, 18-25.viii.2006, ex. Malaise trap. coll: Bunlu 
Subsiri (T469)(lc?.QSBG). -Pha Taem NP. Kua nane née, 15°24.258'N 105°30.855'E, 193 m, 
5-12.viii.2006, ex. Malaise trap, coll: Bunlu Subsiri (T463)(lc?, 19, CNCI; le?, 1 9, MHNG). 

- Pha Taem NP. Pah mhon, 15°24.304'N 105°31.258'E, 230 m, 5-12.viii.2006, ex. Malaise trap, 
coll: Bunlu Subsiri (T461) (3 9 9, QSBG). - Pha Taem NP, Pah mhon. 15°24.304'N 
105°31.258'E, 230 m, 5-12.viii.2006, ex. Malaise trap, coll: Bunlu Subsiri (T461) (1 c?, QSBG). 

- Pha Taem NP, Phu Samui foothill, 15°40.021'N 105°30.448'E, 240 m, 2-9.vi.2007, ex. Malaise 
trap. coll: Tongcam & Banlu (T2207) (le?, 19. QSBG). - Pha Taem NP, Rong Hi Noy, 
15°40.021'N 105°30.448'E. 240 m. 1-7.1.2007. ex. Malaise trap. coll: Thongkam & Pakdee 
(T1476) (1 9. QSBG). - Pha Taem NP, west of HuayPok substation, 15°37.212'N 105°36.903'E, 
438 m, 18-25.iv.2007, ex. Malaise trap, coll: Bunlu Sapsiri (T2170) (19, CNCI). - Pha Taem 
NP, Wild flower field 1, 15°27.336'N 105°34.87'E, 232 m, 4-1 l.xi.2006, ex. Malaise trap, coll: 
Sorawit and Thongdee (T1054) (1 e?, USNM). VIETNAM. - Lam Dong Province, 7 km W Di 
Linh, 860 m. 1 L34N 108.01E. 03.IX.2003. P. Schwendinger (le?, CAUC). - Tay Ninh 
Province, data same as holotype (6 e? e? , 2 9 9 , MFTNG; 1 c? , 3 9 9 , CAUC). 

ETYMOLOGY: The new species is named after the Entomologist Bernhard Merz 
who is our good friend and gives us so many kind and generous helps. 

DiAGNOSlS: Frons velvety black except fronto-orbital plate shining black. Face 
with dense grayish white pruinosity, a black velvety basai spot between antennae, and 
a silvery white trapeziform pruinose spot on spherical médian convexity. Antennal lst 
flagellomere yellow except dorsal margin and tip brown black. A silvery white 
pruinose spot présent between base of antenna and eye. Palpus black. Mesonotum 
covered by dense yellow pruinosity except two latéral margins and sutures with grayish 
white pruinosity, and a pair of grayish médian stripes extending to posterior margin and 
confluent with a large black velvety spot. Wing with three white spots présent between 
R t and M { , a white spot respectively in cells dm and cua l5 a white luniform spot 
beyond dm-cu and a white stripe on dm-cu: subcostal cell brown. Abdominal tergites 
3-6 with dense grayish pruinosity along latéral margins. 

Description 

Length: Body 3.7-4.3 mm (maie), 3.7-4.2 mm (female); wing 3.3-3.8 mm. 

Head (Plates A2, A4) black. Frons velvety black except fronto-orbital plate 
shining black, w ider than long and parallel-sided: a silvery white triangular pruinose 
spot close to inner and outer vertical setae: ocellar triangle black; oc strong and longer 
than posterior or; anterior or reclinate, shorter than posterior or. Face with dense 
grayish white pruinosity, a black velvety basai spot between antennae, a silvery white 



NEW SPECIES OF DIPTERA FROM ASIA 



681 



trapeziform pruinose spot on spherical médian convexity; parafacial with dense silvery 
white pruinosity. Gena with dense silvery white pruinosity and 1 long seta. Antennal 
scape and pedicel black; lst flagellomere coniform, yellow except dorsal margin and 
tip brown black, 1.7 times as long as high; arista brown, long plumose, with longest 
serula longer than height of lst flagellomere. A silvery white pruinose spot présent 
between base of antenna and eye. Proboscis blackish brown, with yellowish and 
blackish setulae; palpus black, with blackish setulae. 

Thorax (Plate A4) brownish black. Mesonotum with dense yellow pruinosity, 
except two latéral margins and sutures with grayish white pruinosity, and with a pair 
of grayish médian stripes extending to posterior margin and confluent with a large 
black velvety spot, a pair of grayish spots on each side of a large black velvety semi- 
circular spot; 0+3 de, anterior de hair-like, slightly longer than acr in latéral view, and 
close to second de; acr in 6 rows; prsc weak hair-like. Anepisternum and katepi - 
sternum with dense grayish white pruinosity; 1 anepst, 1 kepst. Scutellum brown with 
a large black velvety spot on basai 2/3 and a grayish white spot on each side of large 
black spot; distal margin silvery pruinose to black, sometimes with ground color of 
margin yellowish. Halter yellow. 

Legs: femora blackish brown with sparse grayish white pruinosity; tibiae 
yellow, silvery shining on dorsal side, with an apical black ring; and ail tarsomeres 
yellow except 3-5 pale brown. Fore fémur with 3 pv and 6 pd; fore tibia with 1 pre- 
apical ad and 1 short apv. Mid fémur with 5 a; mid tibia with 1 preapical ad and 2 
strong apv. Hind fémur with 2 ad, hind tibia with 1 short preapical ad and 1 short apv. 

Wing (Plate B2) grayish with irregular whitish spots; three white spots présent 
between Rj and M l5 a white spot respectively in cells dm and cua l5 a white luniform 
spot beyond dm-cu and a white stripe on dm-cu; subcostal cell brown; Costa with 2nd 
(between Rj and R2+3), 3rd (between R 2+ 3 and R^) and 4th (between R 4 ^ 5 and Mj) 
sections in proportion of 1.4 mm: 0.7 mm: 0.3 mm; r-m beyond middle of cell dm; 
ultimate and penultimate sections of Mj in proportion of 1.2 mm: 1.3 mm; ultimate 
section of CuAj about 1/6 of penultimate. 

Abdomen blackish brown, tergites 3-6 each with dense grayish pruinosity along 
latéral margins. Maie genitalia (Figs 6-10): protandrium circular with two small ventral 
processes; epandrium broad; surstylus with a claviform anterior ventral process and a 
incision on inner side in posterior view; hypandrium Y-shaped; gonopod degenerate; 
aedeagus with a pair of triangular basai processes, constricted apically with a deep 
incision in ventral view but round and broad apically in latéral view. 

Remarks: The new species is unique in the genus by the following characters: 
frons velvety black except fronto-orbital plate shining black, face dense grayish white 
pruinose, a black velvety basai spot présent between antennae and a silvery white 
trapeziform pruinose spot on the spherical médian convexity, and a différent wing 
pattern. 

Distribution: Thailand, Vietnam. 
Dioides furcatus Shi, Li & Yang, 2009 

Dioides furcatus Shi, Li & Yang, 2009: 95. Type locality: China (Hainan) (CAUC). 

MATERIAL: THAILAND. - Nakhon Si Thammarat Province, Khao Luang National 
Park, 380 m, 8.22.45N/99.44.22E, 17. VII. 2005, P. Schwendinger ( 1 9 , MHNG). - Phetchabun 



682 



L. SHI ETAL. 



Province, Khao Kho NP, Nursery, 16°52.573'N 101°8.077'E, 520 m, 19-26.X.2006, ex. Malaise 
trap, coll: Somchai Chatchumnan and Sa-ink Singtong (T809) (\â, QSBG). VIETNAM. - 
Lam Dong Province, 7 km W Di Linh, 860 m, 1 1 .34N/108.01E, 3. IX. 2003, P. Schwendinger 
(26 MHNG). 

Distribution: China (Hainan), new records to Thailand and Vietnam. 
Dioides incurvatus Shi, Li & Yang, 2009 

Dioides incurvatus Shi, Li & Yang, 2009: 96. Type locality: China (Yunnan) (CAUC). 

MATERIAL: THAILAND. - Chiang Mao Province, Chiang Dao District outside Sua 
Dao cave, 450 m, 19.23.33N/98.55.55E, 27.XII.2007, P. Schwendinger (19, MHNG). 

Distribution: China (Yunnan), new record to Thailand. 
Phobeticomyia uncinata Shi, Li & Yang, 2009 

Phobeticomyia uncinata Shi, Li & Yang, 2009: 62. Type locality: China (Yunnan) (CAUC). 

MATERIAL: THAILAND. - Chiang Mai Province, Doi Inthanon NP, Checkpoint 2, 
18°31.554'N 98°29.94'E, 1700 m, 24.xi.-l.xii.2006, ex. Malaise trap, coll: Y. Areeluck (T1870) 
(1(5, CSCA; \ â, QSBG). - Doi Inthanon NP, Vachiratharn Falls, 18°32.311'N 98°36.048'E, 
700 m, 29.vi.-2.vii.2006, ex. Malaise trap, coll: Y Areeluck (T40) (\6, USNM). -Doi Suthep- 
Pui National Park, What Phrathat, 1050m, 18.48N/98.55E, 30.XII.2005, P. Schwendinger (16, 
MHNG). 

Distribution: China (Yunnan), new record to Thailand. 

Noonamyia bipunctata sp. n. Figs 11-15, Plate B3 

MATERIAL: HOLOTYPE 6 (CAUC): China, Guangxi Province, Jinxiu, Dayaoshan 
National Nature Reserve, Longjunshan, 240-700 m, 27.32N/1 1 1.58E, 29.VII.2005, Yajun Zhu. 
PARATYPE: CHINA. - Guangxi Province, Jinxiu, Dayaoshan National Nature Reserve, 
Longjunshan, 240-700 m, 27.32N/1 1 1.58E, 29.VII.2005, Yajun Zhu (1 9, CAUC). 

ETYMOLOGY: Latin, bi-, prefix meaning two + punctata, meaning spotted, 
referring to the wing with two round hyaline spots in cell r 4+5 ; a féminine adjective. 

DlAGNOSlS: Antennal lst flagellomere yellow, except brown on apical 1/3. 
Palpus brown. Mid and hind femora with a pair of brown latéral spots at tip, tarsomeres 
4-5 pale brown. Wing with a hyaline stripe extending downward from Costa to CuAj 
before vertical level of r-m, separated from a short hyaline stripe in cell cua^ two 
round hyaline spots in cell r 4+5 , a hyaline apical stripe between R 4+5 and Mj; no spots 
on r-m and dm-cu; subcostal cell brown. Abdominal tergites 1-2 yellow with a brown 
transverse médian band on tergite 2, and tergites 3-6 black. 

Description 

Length: Body 3.1 mm, wing 3.0 mm. 

Head yellow. Frons about as long as wide and parallel-sided, with 2 brown 
stripes extending to ocellar triangle; ocellar triangle black; oc and or broken. Face pale 
yellow, with a pair of shining brown spots at ventral corner. Gena about 1/5 height of 
eye. Antenna yellow, except lst flagellomere brown on apical 1/3, 2.0 times longer 
than high; arista blackish brown, plumose, with longest setula slightly longer than 
height of lst flagellomere. Proboscis yellow, with yellowish and blackish setulae; 
palpus pale brown, with blackish setulae. 



NEW SPECIES OF DIPTERA FROM ASIA 



683 




14 15 

FIGS 11-15 

Noonamyia bipunctata sp. n., maie holotype. (11) Epandrium and protandrium, latéral view. (12) 
Protandrium, anterior view. (13) Epandrium, posterior view. (14) Aedeagal complex, ventral 
view. (15) Aedeagal complex, latéral view. Scale 0.1 mm 

Thorax blackish brown except yellow on basai 1/4. Mesonotum with 0+3 de, 
acr in 6 rows. 1 anepst, 1 kepst. Scutellum entirely yellow. 

Legs yellow. Mid and hind femora with a pair of brown latéral spots at tip, 
tarsomeres 4-5 pale brown. Fore fémur with 3 pv and 4 pd, ctenidium with 9 short 
setulae; fore tibia with 1 weak preapical ad and 1 apv. Mid fémur with 4 a; mid tibia 
with 1 preapical ad and 2 strong apv. Hind fémur with a row of weak av; hind tibia 
with 1 weak preapical ad and 1 short apv. 



684 



L. SHI ET AL. 



Wing (Plate B3) with a hyaline stripe extending downward from Costa to CuAj 
before vertical level of r-m, separated from a short hyaline stripe in cell cua^ two 
round hyaline spots in cell r 4+5 , a hyaline apical stripe between R 4+5 and Mj; no spots 
on r-m and dm-cu; subcostal cell brown; Costa with 2nd (between Rj and R2+3), 3rd 
(between R 2+ 3 and R4+5) and 4th (between R 4+5 and Mj) sections in proportion of 
1.5 mm : 0.5 mm : 0.4 mm; r-m beyond middle of distal cell; ultimate and penultimate 
sections of Mj in proportion of 1.0 mm : 1.1 mm; ultimate section of CuAj about 1/5 
of penultimate. Halter pale yellow. 

Abdominal tergites 1-2 yellow, with a brown transverse médian band on tergite 
2; tergites 3-6 black and protandrium and epandrium yellow. Maie genitalia (Figs 
11-15): protandrium semicircular; epandrium U-shaped in ventral view, with long setae 
on ventral margin; surstylus consisting of a short claviform process and a long subuli- 
form process with three teeth and many setulae; hypandrium V-shaped with a pair of 
knife-like processes; gonopod short, wide with two or three teeth and one or two apical 
setae in ventral view; aedeagus curved apically, with a pair of triangular basai pro- 
cesses in ventral view; aedeagal apodeme Y-shaped, longer than aedeagus. 

REMARKS: The new species is similar to Noonamyia lyneborgi Stuckenberg 
from the Philippines (Palawan Island) in the wing with two round hyaline spots in cell 
r 4+5 , abdomen with tergites 1-2 yellow and tergites 3-6 black. But it can be separated 
from the latter by the fore tibia having 1 weak preapical ad, the wing having a hyaline 
stripe extending downward from Costa to CuAj and separated from a short hyaline 
stripe in cell cua } , the surstylus consisting of a short claviform process and a long sub- 
uliform process with three teeth and many setulae. In N. lyneborgi, the fore tibia has 
no preapical ad; the wing has a complète hyaline stripe extending from costal margin 
to posterior margin before the vertical level of r-m; the surstylus has a tiny apical tooth 
(Stuckenberg, 1971; Sasakawa & Pong, 1990). 

Distribution: China (Guangxi). 

Noonamyia flavoscutellata sp. n. Figs 16-20, Plate B4 

MATERIAL: HOLOTYPE 6 (CAUC): China, Guangxi Province, Jinxiu, Dayao 
Mountain, Yinshan, 1000 m, 27.VII.2005, Yajun Zhu. PARATYPES: CHINA. - Guangxi 
Province, Jinxiu, Dayaoshan National Nature Reserve, Longjunshan, 240-700 m, 
27.32N/1 1 1 .58E, 29.VII.2005, Yajun Zhu (1 9 , CAUC). THAILAND. - Chiang Mai Province, 
Doi Inthanon National Park, 750 m, 26.XII.2001, P. Schwendinger (26 6, MHNG). - Doi Pha 
Hom Pok W of Fang, 1500 m, 20.02.43N/99.08*.43E, P. Swendinger (26 6, MHNG). - Doi 
Suthep-Pui National Park, Huay Khok Ma, 1 9.XII.200 1 , P. Schwendinger (4 6 6 , 4 9 9 , MHNG; 
36 6, 3 9 9, USNM). - Doi Suthep-Pui National Park, nr Doi Suthep summit, 1550 m, 
1 8.82N/98.89E, 26.X.2000, B. Merz & P. Schwendinger (366,399, MHNG). - Doi Suthep- 
Pui National Park, 1420 m, 18.50.09N/98.53.47E, 18.XII.2001, P. Schwendinger (16, 19, 
CAUC). - Doi Suthep, Huay Khok Ma, 1300 m, 18.48/N98.55E, 26.XII.2006, P. Schwendinger 
(\6, 19, MHNG). - Doi Suthep-Pui National Park, Palad Trail, 4-800 m, 18.79N/98.93E, 
16.X.2000, B. Merz & P. Schwendinger (\6, MHNG). - Doi Suthep-Pui National Park, nr 
Phuphing Palace, 1250 m, 18.80N/98.90E, 26.X.2000, B. Merz & P. Schwendinger (46 6,19, 
MHNG; 26 6, CSCA). - Doi Suthep-Pui National Park, What Phrathat, 900 m, 18.80N/98.92E, 
1 6.X.2000, B. Merz & P. Schwendinger (266,59 9, MHNG; 29 9, CSCA). - Doi Suthep-Pui 
National Park, What Phrathat, 950m, 18.80N/98.92E, 19.X.2000, B. Merz & P. Schwendinger 
(\6, 29 9, MHNG; le?, 3 9 9, CAUC). - Doi Suthep-Pui National Park, What Phrathat, 
1050m, 18.48N/98.55E, 30.XII.2005, P. Schwendinger (\6, MHNG). - Doi Suthep-Pui 



NEW SPECIES OF DIPTERA FROM ASIA 



685 




19 20 

FIGS 16-20 

Noonamyia flavoscutellata sp. n., maie paratype. (16) Epandrium and protandrium, latéral view. 
(17) Protandrium, anterior view. (18) Epandrium, posterior view. (19) Aedeagal complex, ventral 
view. (20) Aedeagal complex, latéral view. Scale 0.1 mm. 

National Park, What Phrathat, 1100m, 18.80N/98.92E, 19.X.2000, B. Merz & P. Schwendinger 
(26 6, 3 9 9 , MHNG). - Doi Suthep-Pui National Park, What Phrathat, 1200m, 18.80N/98.92E, 
1 .XI.2000, B. Merz & P. Schwendinger (366,299, MHNG). - Doi Suthep, 1150-1 300 m, 20- 
28.XII.2003, P. Schwendinger (3 6 6 , 6 9 9 , MHNG; 1 6 , 1 9 , CAUC; 3 6 6 , 3 9 9 , CSCA). - 
Lamphun Province, Doi Khuntan National Park, 45km SE Chiang Mai, 650m, 18.50.N/99.27E, 



686 



L. SHI ETAL. 



2.XI.2000, B. Merz & P. Schwendinger (1 S, MHNG). VIETNAM. - Kien Giang Province, 
Phu Quoc Island, 250m, 10.23N/104.01E, 14-16.VIII.2003, P. Swendinger (1 Ô, MHNG). 

ETYMOLOGY: Latin, flav-, prefix meaning yellow + scutellata, referring to the 
scutellum; a féminine adjective. 

DiAGNOSlS: Antennal lst flagellomere pale brown. Palpus brown except for 
pale yellow at third. Wing with a short hyaline stripe in cell cua^ an elliptical hyaline 
spot in cell r 4+5 , a semicircular hyaline apical stripe between R 2+ 3 and CuAj, confluent 
with a digitifonn hyaline stripe beyond dm-cu; a narrow pale brown stripe on r-m. 

Description 

Length: Body 2.1-2.8 mm (maie), 2.2-2.8 mm (female); wing 3.0-3.5 mm 
(maie), 3.0-3.3 mm (female). 

Head yellow. Frons as long as wide and parallel-sided, with 2 brown stripes 
extending to ocellar triangle; ocellar triangle black; oc strong, as long as anterior or and 
anterior or shorter than posterior or. Face pale yellow, with a pair of shining brown 
latéral spots at ventral corner. Gena about 1/9 height of eye. Antennal scape brownish 
yellow, pedicel yellow and lst flagellomere pale brown except yellow on base part of 
ventral margin, lst flagellomere nearly 1 .8 times longer than high; arista long plumose, 
black except for brownish base, with longest setula longer than height of lst flagello- 
mere. Proboscis yellow, with yellow and blackish setulae; palpus brown except for pale 
yellow at third, with black setulae. 

Thorax blackish brown, slightly shining with sparse grayish white pruinosity. 
Mesonotum with broad yellow anterior margin; 0+3 de, acr in 6 rows; prsc broken. 
1 anepst, 1 kepst. Scutellum entirely yellow. Halter pale yellow except brown knob. 

Legs yellow, except fore tibia slightly brownish yellow, mid and hind femora 
brown at tip, mid tibia brown on basai 1/2-2/3 and ail tarsomeres 3-5 pale brown. Fore 
fémur with 2 pv and 6 pd, ctenidium with 9-10 short setulae; fore tibia with 1 weak 
preapical ad and 1 apv. Mid fémur with 4 a; mid tibia with 1 preapical ad and 2 strong 
apv. Hind fémur with a row of weak av; hind tibia with 1 weak preapical ad and 1 short 
apv. 

Wing (Plate B4) mostly brown, with a triangular hyaline stripe extending down- 
ward from Costa to Mj, a short hyaline stripe in cell cua^ an elliptical hyaline spot in 
cell r 4+5 , a semicircular hyaline apical stripe between R 2+ 3 and CuAj, confluent with 
a digitiform hyaline stripe beyond dm-cu; a narrow pale brown stripe on r-m and a 
tinged brown triangular stripe over dm-cu, extending to the tip of CuA t ; apical part of 
subcostal cell brown; Costa with 2nd (between Rj and R2+3), 3rd (between R 2 +3 and 
R 4+5 ) and 4th (between R 4 ^ 5 and Mj) sections in proportion of 1.9 mm : 0.6 mm : 
0.5 mm; r-m beyond middle of distal cell; ultimate and penultimate sections of Mj in 
proportion of 1.0 mm : 1.0 mm; ultimate section of CuAj about 1/5 of penultimate. 

Abdomen blackish brown, slightly shining with sparse grayish white; protan- 
drium and epandrium yellow. Maie genitalia (Figs 16-20): protandrium semicircular; 
epandrium U-shaped in ventral view; surstylus short, curved downward; hypandrium 
U-shaped, gonopod transverse broad, undulating ventral margin with 1 apical seta in 
ventral view; aedeagus consisting of a small incision, a pair of triangular basai pro - 
cesses, an arch-like ventral sclerite with apical teeth and a pair of dorsal sclerites with 
acuate apical processes; aedeagal apodeme Y-shaped, longer than aedeagus. 



NEW SPECIES OF DIPTERA FROM ASIA 



687 



REMARKS: The new species is very similar to Noonamyia sasakawai Papp from 
Thailand in the following characters: the wing has an incomplète hyaline stripe before 
the base of r-m, it lacks a round hyaline spot in cell dm, it has a semicircular hyaline 
marginal stripe between the tips of R 4+5 and CuAj, the hind tibia has a short preapical 
ad, the scutellum is entirely yellow or brownish yellow, and the surstylus is acuate 
apically and curved backward. The new species differs from Noonamyia saskawai in 
détails of the genitalia, particularly the epandrium having 3-4 long dorsal setae, and the 
gonopod having an undulating ventral margin and an apical seta. In Noonamyia 
sasakawai, the epandrium has 2 long dorsal setae and the gonopod is comma-like with 
a straight ventral margin and 1-2 subapical setae ventrally. The new species is also 
similar to Noonamyia euphlebia Sasakawa & Pong from Malaysia and Thailand in the 
following characters: legs with fore tibia slightly brownish yellow, mid and hind 
femora brown at tip and wing with an incomplète hyaline stripe before base of r-m. But 
it can be separated from the latter by the mid tibia brown on basai 1/2-2/3, the 
scutellum being yellow, the semicircular hyaline marginal stripe being présent, and the 
epandrium with 3-4 pairs of dorsal setae. In Noonamyia euphlebia, the mid tibia is 
entirely brown, the scutellum is brownish black, the brown area in cell mj almost 
extends to the margin of wing but does not form a marginal stripe around to CuAj, and 
the epandrium has 5-6 pairs of dorsal setae (Sasakawa & Pong, 1990; Sasakawa, 1998). 

Distribution: China (Guangxi), Thailand, Vietnam. 
Noonamyia sabahna Sasakawa & Pong, 1990 

Noonamyia sabahna Sasakawa & Pong, 1990: 32. Type locality: Malaysia (Sabah). 

MATERIAL: PHILIPPINES. - Mindanao, Agusan del Sur. 10 km SE San Francisco, 
12.IX.1959, coll: L.W. Quate (1 (sex unknown), BPBM); Misamis Oriental, Minubana, 1050- 
1200 m, 5-9.IV.1961, coll: H. Torrevillas (1 (sex unknown), BPBM). 

Distribution: Malaysia (Sabah), new record to the Philippines. 

Noonamyia sasakawai Papp, 2006 Figs 21-26, Plates C1-C5 

Noonamyia sasakawai Papp, in Papp, Merz & Fôldvâri, 2006: 185. Type locality: Thailand 
(Doi Pui). 

MATERIAL: PARATYPES: THAILAND. - Nan Province. Tham Sakoen NP, 
19°23'N/100°38'E, No. 17, over and along creek in forest, 26.XI.2003, M. Fôldvâri & A. 
Szappanos (1 d, HNHM). - Doi Phuka NP, No. 18, UV light, 26-27.XI.2003, L. Peregovits, M. 
Fôldvâri, Â. KÔrôsi. A. Szappanos & B. Maklâri-Kis (19, HNHM). ADDITIONAL MATERIALS: 
THAILAND. - Chiang Mai Province. Chiang-dao. 5-1 1 .IV. 1958, T.C. Maa (\6, BPBM). - 
Loei Province, Phu Kradueng NP, Savannah in pine forest, 16°53.092'N 101°47.413*E, 1257 m, 
30-31.xii.2006, ex. pan traps, coll: Sutin Gongla-sae (T1215) (19, QSBG). - Phu Ruea NP, Ma 
Kraow ditch, 17°29.652'N 101°21.020'E, 1167 m, 5-12.xi.2006, ex. Malaise trap, coll: Patikhom 
Tumtip (T1116) (19, QSBG). - Phu Ruea NP, Ma Kraow ditch, 17°29.652'N 101°21.020'E, 
1167 m, 7-8.xi.2006, pan traps, Patikhom Tumtip (T1110) (lô\ 19, USNM). - Phu Ruea NP, 
Ma Kraow ditch, 17°29.652'N 101°21.020'E, 1167 m, 8-9.xi.2006, ex. pan trap, coll: Patikhom 
Tumtip (XI 111) (19, QSBG). - Phu Ruea NP, Ma Kraow ditch, 17°29.652'N 101°21.020'E, 
1167 m, 10-ll.xi.2006, ex. pan trap, coll: Patikhom Tumtip (T1113) (ld, QSBG). - Phu Ruea 
NP, office, 17°28.826'N 101°21.330'E, 860 m, 7-8.vii.2006, ex. pan trap. coll: Patikhom Tamtip 
(T305) (26 6, CSCA). - Phu Ruea NP, Pan Hin Khan Maak ditch, 17°30.042'N 101°20.474'E, 
1219 m, 6-7.ii.2007, ex. pan trap, coll: Patikhom Tumtip (T1699) ( 1 9 , CSCA). - Phu Ruea NP, 
Pan Hin Khan Maak ditch, 17°30.042'N 101°20.474'E. 1219 m, 8-9.ii.2007, ex. pan trap, coll: 



688 



L. SHI ETAL. 




FIGS 21-26 

Noonamyia sasakawai Papp, 2006, maie paratype. (21) Epandrium and protandrium, latéral 
view. (22) Protandrium, anterior view. (23) Epandrium, posterior view. (24) Surstylus, anterior 
view. (25) Aedeagal complex, ventral view. (26) Aedeagal complex, latéral view. Scale 0.1 mm. 

Patikhom Tumtip (T1701) (1(5, CSC A). - Phetchabun Province, Nam Nao NP, Héliport, 
16°43.156'N 101°35.118'E, 890 m, 18-19.xii.2006, ex. pan trap, coll: Noopean Hongyothi & 
Leng Janteab (T1425) (19, QSBG). - Nam Nao NP, Héliport, 16°43.156'N 101°35.1 18'E, 890 
m, 20-21.xii.2006, ex. pan trap, coll: Noopean Hongyothi & Leng Janteab (T1427) (le? 19, 
QSBG). - Nam Nao NP, Tham Pra Laad Forest Unit, 16°44.963'N 101°27.833'E, 71 1 m, 4-5.vi- 



NEW SPECIES OF DIPTERA FROM ASIA 689 




PLATE C 

Noonamyia sasakawai Papp, 2006, maie paratype. (1) Habitus, latéral view. (2) Mesonotum, 
dorsal view. (3) Wing. (4) Head, anterior view. (5) Abdomen, dorsal view. 



ii.2006, ex. pan trap, coll: Noopean Hongyothi & Leng Janteab (T417) (19, QSBG). - 
Phitsanulok Province, Thung Salaeng Luang NP, Dry evergreen, 16°50.217'N 101°52.541'E, 
580 m, ll-12.viii.2006, ex. pan trap, coll: Pongpitak Pranee (T559) (19, CSCA). - Thung 
Salaeng Luang NP, Dry evergreen, 16°50.217'N 101°52.541'E, 580 m, 13-19.viii.2006, ex. pan 
trap, coll: Pongpitak Pranee (T562) (1 9, CSCA). VIETNAM. - Lam Dong Province, - 6 km 
S of Dalat, 1550 m, 12.IX.1960, J.L. Gressitt (1<Î, CSCA). Fyan, 900-1000 m, 11.VII-9.VI- 
11.1961, N.R. Spencer (le?, USNM). - Di Lihn (Djiring), 27.IX-14.X.1960, CM. Yoshimoto 
( 1 6 , BPBM). - Fyan, 1 200 m, 1 1 . VII-9. VII 1. 1 96 1 , N.R. Spencer ( 1 6 , 1 9 , BPBM). 

DiAGNOSiS: Antenna brownish yellow, lst flagellomere brown on apical 1/2. 
Palpus brown except for yellow basai 1/4. Wing with a short hyaline stripe in cell cuaj; 



690 



L. SHI ETAL. 



an elliptical hyaline spot in cell r 4+5 , a semicircular hyaline apical stripe between R 2 +3 
and CuAj; a narrow hyaline stripe on r-m and a pale brown triangular stripe over 
dm-cu, extending to the tip of CuAj. 

Description 

Length: Body 2.5 mm (maie), 2.5 mm (female); wing 2.9 mm (maie), 3.0 mm 
(female). 

Head (Plate C4) yellow. Frons with 2 brown stripes extending from anterior 
margin to ocellar triangle, slightly wider than long and parallel-sided; ocellar triangle 
black; oc strong, as long as posterior or and anterior or about 1.5 times as long as 
posterior or. Face white, with a pair of shining blackish brown latéral spots at ventral 
corner. Gena about 1/9 height of eye. Antenna brownish yellow, lst flagellomere 
brown on apical 1/2, 1.5 times longer than high; arista long plumose, black except for 
brownish base, with longest setula longer than height of lst flagellomere. Proboscis 
yellow, with yellowish and blackish setulae; palpus brown except for yellow basai 1/4, 
with black setulae. 

Thorax (Plate C2) blackish brown with sparse grayish white pruinosity. 
Mesonotum with broad yellow anterior margin; 0+3 de, acr in 6 rows. Scutellum 
entirely brownish yellow. 

Legs pale yellow to yellow except mid and hind tibiae brown at tip, mid tibia 
blackish brown on basai 1/3-1/2 and ail tarsomeres 4-5 pale brown. Fore fémur with 
2 pv and 6 pd. ctenidium with 9-10 short setulae; fore tibia with 1 weak preapical ad 
and 1 apv. Mid fémur with 4 a: mid tibia with 1 preapical ad and 2 strong apv. Hind 
fémur with 3 av on apical 1/2; hind tibia with 1 weak preapical ad and 1 short apv. 

Wing (Plate C3) with a triangular hyaline stripe extending downward from 
Costa to M] before vertical level of r-m and a narrow hyaline stripe in cell cua^ an 
elliptical hyaline spot in cell r 4 ^ 5 , a semicircular hyaline apical stripe between R 2+ 3 and 
CuAj; a narrow hyaline stripe on r-m and a pale brown triangular stripe over dm-cu, 
extending to the tip of CuAj; subcostal cell brown at tip; Costa with 2nd (between Rj 
and R2-3), 3rd (between R 2 _3 and R^) and 4th (between R 4+5 and Mj) sections in 
proportion of 1.7 mm : 0.5 mm : 0.4 mm; r-m beyond middle of distal cell; ultimate 
and penultimate sections of Mj in proportion of 1.2 mm : 1.0 mm; ultimate section of 
CuAj about 1/3 of penultimate. Halter pale yellow except knob brown on basai 1/2. 

Abdomen (Plate C5) black, with sparse grayish white pruinosity; protandrium 
and epandrium brownish yellow. Maie genitalia (Figs 21-26): protandrium semi- 
circular; epandrium with two pairs of long dorsal setae in ventral view; surstylus short, 
curved downward with several setae; hypandrium U-shaped; gonopod comma-like 
with 1-2 long setae in ventral view; aedeagus consisting of a shallow incision, a pair of 
slender ventral sclerites with several small acuate processes, membranous part with 
many tiny teeth and a pair of dorsal sclerites with acuate apical processes; aedeagal 
apodeme Y-shaped, longer than aedeagus. 

Remarks: This species is very similar to Noonamyia flavoscutellata sp. n. and 
the differentiating characteristics of the two species are reflected in the key and in the 
remarks for the latter species. 

Distribution: Thailand. new record to Vietnam. 



NEW SPECIES OF DIPTERA FROM ASIA 



691 



KEY TO SEPARATE SPECIES OF THE GENUS NOONAMYIA IN THE WORLD 

(The following key is based on the key by Sasakawa & Pong (1990) with the 
new species integrated at the appropriate couplets) 

1 a Wing with a complète white stripe before r-m, running vertically through 

whole wing 2 

lb Wing without complète white stripe before r-m 4 

2a Wing with 1-2 small white round spots in cell r 4 ^ 5 3 

2b Wing only with a wide white stripe in cell r 4+5 (see Frey, 1958: fig. 1 1 ). 

Distributed in Philippines N. fascipennis (Frey, 1958) 

3a Wing with one white round spot in cell r 4+5 ; katepisternum pale yellow; 
fore tibia without preapical ad; surstylus consisting of two processes 
(see Sasakawa & Pong, 1990: figs 5 A, 6-7). Distributed in Malaysia 

N. abdominalis Sasakawa & Pong, 1990 

3b Wing with two white round spots in cell r 4+5 ; katepisternum brownish 
black; fore tibia with preapical ad; surstylus unique, with an acuate 
apical tooth (see Stuckenberg, 1971: figs 59, 87). Distributed in 

Philippines N. lyneborgi Stuckenberg, 1971 

4a Wing with a round hyaline spot in cell dm; epandrium with a short 
posterior process and surstylus striate (see Sasakawa & Pong, 1990: 

figs 5C, 8-9). Distributed in Malaysia, Philippines 

N. sabahna Sasakawa & Pong, 1990 

4b Wing without spot in cell dm; epandrium and surstylus not as above 5 

5a Wing without spot in cell r 4+5 , only with two white stripes passed 
through cell r 4+5 (see Frey, 1958: fig. 10). Distributed in Philippines . . . 

N. irregularis (Frey, 1958) 

5b Wing with 1-2 white round or elliptical spots in cell r 4+5 6 

6a Hind tibia without preapical ad 7 

6b Hind tibia with a short preapical ad 9 

7a Wing with two narrow white stripes situated around r-m 8 

7b Wing with a wide white stripe before r-m (see Okadome, 1982: fig. 1). 

Distributed in Philippines N. chujoi Okadome, 1982 

8a Wing with a white U-shaped spot surrounding r-m and a white triangu- 
lar spot in cell m} before vertical level of a white round spot in cell r 4+5 

(see Stuckenberg, 1971: fig. 58). Distributed in Philippines 

N. palawanensis Stuckenberg, 1 97 1 

8b Wing without U-shaped spot surrounding r-m, a white triangular spot in 
cell iri] at same vertical level with a white round spot in cell r 4+5 (see 

Sasakawa & Pong, 1990: fig. 5D). Distributed in Malaysia 

N. pleuralis Sasakawa & Pong, 1990 

9a Wing without white or hyaline spot or stripe on r-m and dm-cu, two 
white spots beyond r-m in cell r 4+5 (Plate B3). Distributed in Southern 

China N. bipunctata sp. n. 

9b Wing with white or hyaline spot or stripe on r-m and dm-cu, one white 

spot beyond r-m in cell r 4+5 10 



692 



L. SHI ETAL. 



10a Wing with a semicircular hyaline marginal stripe between tips of R 4+5 

and CuAj; scutellum entirely yellow or brownish yellow 11 

10b Wing without semicircular hyaline marginal stripe between tips of R 4+5 

and CuAj; scutellum at least black or brown on basai 1/2 13 

1 1 a Cross vein r-m with a pale brown spoon-like spot; surstylus with two 
subuliform processes; gonopod with two acuate furcated apical pro- 
cesses in ventral view (see Shi & Yang, 2009: figs 1, 6). Distributed in 
Southern China N. bisubulata Shi & Yang, 2009 

11b Cross vein r-m without pale brown spoon-like spot; surstylus acuate 

apically and curved backward; gonopod not as above 12 

12a Epandrium with 3-4 long dorsal setae; gonopod with undulating ventral 
margin and an apical seta in ventral view (Figs 18, 19). Distributed in 
Southern China, Thailand, Vietnam N. flavoscutellata sp. n. 

12b Epandrium with 2 long dorsal setae; gonopod comma-like with straight 
ventral margin and 1-2 subapical setae in ventral view (Figs 23, 25). 
Distributed in Thailand, Vietnam N. sasakawai Papp, 2006 

13a Wing with a narrow hyaline stripe on r-m; gonopod acinaciform and 
membranous area of aedeagus with small teeth in ventral view (see 
Sasakawa & Pong, 1990: fig. 5B; Sasakawa, 1998: fig. 2B). Distributed 
in Laos, Malaysia, Thailand, Vietnam . . N. euphlebia Sasakawa & Pong, 1990 

13b Wing with a pale brown umbrella-like spot on r-m; gonopod spoon-like 
and membranous area of aedeagus without small teeth in ventral view 

(see Shi & Yang, 2009: figs 2, 12). Distributed in Southern China 

N. umbrellata Shi & Yang, 2009 



ACKNOWLEDGEMENTS 

The first author appréciâtes Dr. Bernhard Merz (Geneva, Switzerland) and 
Dr. Lâszlô Papp (Budapest, Hungary) for their loan and donation of spécimens, 
supplying literature and giving short-period funding support for scientific research in 
their two muséums. Thanks are also extended to Dr. Neal Evenhuis (Honolulu, 
Hawai'i, USA) for loan and donation of spécimens, and Brian Brown and Mike 
Sharkey for providing spécimens from their TIGER project (NSF DEB-0542864), in 
coordination with Wichai Srisuka (QSBG), who is also thanked very much for 
allowing for the distribution of Thai spécimens. Also, sincère thanks to Dr. Bernhard 
Merz for reviewing the manuscript and giving precious comments, to Dr. Gâbor 
Lengyel (Budapest, Hungary) for providing type spécimens, Dr. Mitsuhiro Sasakawa 
(Hirakata, Japan) and Dr. Martin Hauser (Sacramento, USA) for their kind help. We are 
grateful to Dr. L. Peregovits, Dr. M. Fôldvâri, Dr. Â. Kôrôsi, Dr. A. Szappanos & 
Dr. B. Maklâri-Kis (Budapest, Hungary), Dr. P. Schwendinger (Geneva, Switzerland), 
Dr. Yajun Zhu (Shanghai, China), and numerous others from the TIGER project, for 
collecting spécimens. 



NEW SPECIES OF DIPTERA FROM ASIA 



693 



REFERENCES 

FREY, R. 1958. Studien iibcr ostasiatische Dipteren VI. Nothybidac, Micropezidae, 

Opomyzidae. Notulae entomologicae 38: 37-51. 
HENDEL, F. 1920. Neue Cestrotus-Arten des ungarischen Nationalmuseums (Dipter., 

Lauxaniidae). Verhandhmgen der zoologisch-botanischen Gesellschaft in Wien 70: 

74-80. 

MCALPINE, J. F. 1981. Morphology and Terminology — Adults (pp. 9-63). In: McALPINE, J. F, 

PETERSON, B. V., SHEWELL, G. E. TESKEY, H.J.,VOCKEROTH, J. R., & WOOD, D. M. 

(Coords), Manual of Nearctic Diptera. Volume 1. Research Branch, Agriculture Canada, 

Ottawa, Monograph 27: 674 pp. 
MEIJERE, J. C. H. de. 1910. Studien ùber sùdostasiatische Dipteren IV. Die neue Dipterenfauna 

van Krakatau. Tijdschrift voor Entomologie 53: 58-194. 
OKADOME, T. 1982. A new species of the genus Noonamyia Stuckenberg from the Philippines 

(Diptera: Lauxaniidae) (pp. 183-185). In: SATO, M., HORI, Y., OKADOME, T. (eds). 

Spécial issue to the memory of retirement of Emeritus Professor Michio Chujo. 

Association of the Mémorial Issue of Emeritus Professor M. Chujo, Nagoya, Japan. 

185 pp. 

PAPP, L., MERZ, B. & FÔLDVÂRI, M. 2006. Diptera of Thailand: A summary of the families and 

gênera with références to the species représentations. Acta Zoologica Academiae 

Scientiarum Hungaricae 52(2): 97-296. 
SASAKAWA, M. 1998. Oriental Lauxaniidae (Diptera). Part 1. Scientific Report of the Kyoto 

Prefectural University, Human Environment and Agriculture 50: 49-74. 
SASAKAWA, M. & PONG, T. Y. 1990. Lauxaniidae (Diptera) of Malaysia (Part 1). Esakia, Spécial 

Issue No. 1: 123-136. 

SHEWELL, G. E. 1987. Lauxaniidae (pp. 951-964). In: McALPINE, J. F., PETERSON, B.V., 

SHEWELL, G. E., TESKEY, H. J., VOCKEROTH, J. R., & WOOD, D. M. (Coords), Manual 

of Nearctic Diptera. Volume 2. Research Branch, Agriculture Canada, Ottawa, 

Monograph 28: 675-1332. 
SHI, L., YANG, D. 2009. Two new species of the genus Noonamyia from Hainan in China 

(Diptera, Lauxaniidae). Zootaxa 2014: 34-40. 
SHI, L., YANG, D. & GAIMARI, S. D. 2009. Species of the genus Cestrotus Loew from China 

(Diptera, Lauxaniidae). Zootaxa 2009: 41-68. 
SHI, L., Ll, W. L. & YANG, D. 2009a. Five new species of the genus Dioides from China 

(Diptera: Lauxaniidae). Annuals Zoologici 59(1): 93-105. 
SHI, L., Ll, W. L. & YANG, D. 2009b. Two new species of the genus Phobeticomyia from China 

(Diptera, Lauxaniidae). Zootaxa 2090: 57-68. 
STUCKENBERG, B. R. 1971. A review of the Old World gênera of Lauxaniidae (Diptera). Annals 

of the Natal Muséum 20(3): 499-610. 



Revue suisse de Zoologie 118 (4): 695-721; décembre 201 1 



On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphidiinae) 
of the Philippines, II 

Ivan LÔBL 

Muséum d'histoire naturelle, Case postale 6434, CH-121 1 Genève 6. Switzerland. 
E-mail: ivan.lobl@bluewin.ch 

On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphidiinae) of 
the Philippines, II - The Philippine species of Scaphobaeocera and 
Xoîidium are reviewed and a key to the species of Scaphobaeocera is pro- 
vided. The following new species are described: Scaphobaeocera bulbosa 
sp. n.. S. complicans sp. n., S. data sp. n., S. davaoana sp. n., S. episternalis 
sp. n., S. escensa sp. n.. S. excisa sp. n., S. hamata sp. n.. S. monticola 
sp. n., S. montivagans sp. n.. S. orousseti sp. n., S. palawana sp. n., S. pseu- 
dotenella sp. n., S. pubiventris sp. n., S. serpenîis sp. n.. S. watrousi sp. n., 
S. werneri sp. n., and Xotidium tubuliferum sp. n. Scaphobaeocera saba - 
pensis Lôbl is reported for the first time from the Philippines. 

Keyvvords: Coleoptera - Staphylinidae - Scaphidiinae - Scaphisomatini - 
taxonomy - Philippines 

INTRODUCTION 

The présent paper deals with the Philippine species of Scaphobaeocera Csiki, 
1909 and Xotidium Lôbl, 1992, both belonging to the Baeocera group (see Leschen & 
Lôbl, 2005). Members of thèse two gênera have the body laterally compressed and the 
ventrites are highly vaulted. both features correlated with approximate metacoxae. The 
third and more species-rich genus of the group occurring in the Philippines, Baeocera 
Erichson, 1845, will be treated in a separate study. Thèse three gênera share two un- 
reversed synapomorphies: aciculate maxillary palpi and the présence of prothoracic 
corbicula. While Baeocera is almost cosmopolitan in distribution, Scaphobaeocera is 
absent from the New World and occurs mainly in the tropics and subtropics of Asia. 
Members of Scaphobaeocera are commonly found in samples of moist forest litter and 
rotten wood and are according to the so far available observations myxomyceto- 
phagous. Xotidium is a poorly known group of a few tropical species. Data on their host 
préférences are not available. 

Both, Scaphobaeocera and Xotidium, may be distinguished from other Scaphi - 
somatini gênera by keyes given in Lôbl (1992) and Leschen & Lôbl (2005). Most 
species of Scaphobaeocera have elytra iridiscent and each with a fine parasutural stria. 
Eventually also other body parts are iridescent, due to the présence of fine, transver - 
sely striate microsculpture. Thèse features are unknown in other Scaphisomatini. 



Manuscript accepted 14.06.201 1 



696 



L LÔBL 



MATERI AL AND METHODS 

Most of the material dealt with in the présent study was found in samples of 
forest litter and rotten wood, and extracted in winkler-moczarski or berlese devices. 
The spécimens studied are housed in the following institutions: 
FMNH Field Muséum of Natural History, Chicago 
MHNG Muséum d'histoire naturelle, Geneva 
SMNS Staatliches Muséum fur Naturkunde, Stuttgart 

The methods are as in Lôbl (1992). The species of Scaphobaeocera are given 
below in alphabetic order, for convenience and because species groups were not yet 
defined within the genus. 

TAXONOMY 

Scaphobaeocera Csiki, 1909 

This genus comprises 77 species currently recognized as valid. Its range covers 
large parts of tropical and subtropical Asia, extending to warm temperate areas of Far 
East Russia, the Pacific islands, Australia, Africa, Seychelles and the Mascarene archi- 
pelago. Only one species, S. minutissima (Lôbl, 1969), was so far reported from the 
Philippines (Lôbl, 1969, 1972, 1997). It is notable that this species was absent from 
extensive modem collections. It was based on a single spécimen found in an old 
collection that cornes, according to its label data, from Mount Makiling. Scapho- 
baeocera appears particularly diverse on Mount Makiling where eight of the 19 
Philippine species were found. However, this fact may be explained rather by the more 
intensive field work in the easily accessible sites of that mountain than by diversity 
patterns. 

Key to the Philippine species of Scaphobaeocera 

1 Antennomere VII longer than antennomeres IV to VI combined and 

more than 5 times as long as antennomere VIII S. escensa sp. n. 

Antennomere VII much shorter than antennomeres IV to VI combined 

and much less than 5 times as long as antennomere VIII 2 

2 Antennomere XI about 2 or 2.5 times as long as antennomere X and 

about 3 to 5 times as long as wide 3 

Antennomere XI about 1.2 to 1.5 times as long as antennomere X and 
about 2 to 3 times as long as wide 7 

3 Hypomeron lacking stria S. palawana sp. n. 

Hypomeron with longitudinal stria 4 

4 Elytra microsculptured and usually iridescens 5 

Elytra not microsculptured and not iridescent. Aedeagus without spiral 
flagellum 6 

5 Small species 1 .0 mm long. Latéral parts of metaventrite distinctly punc- 
tate. Basolateral parts of abdominal ventrite 1 not microsculptured. 

Aedeagus with flagellum simple, spiral S. minutissima (Lôbl) 

Medium-sized species 1.3-1.4 mm long. Latéral parts of metaventrite 
extremely finely punctate. Entire abdominal ventrite 1 distinctly micro - 
sculptured S. montivagans sp. n. 



SCAPHISOMATINI OF THE PHILIPPINES 



697 



6 Abdominal ventrite 1 with microsculpture consisting of transverse striae. 

Aedeagus with simple internai sac S. watrousi sp. n. 

Abdominal ventrite 1 lacking microsculpture. Aedeagus with complex 

internai sac S. complicans sp. n. 

Elytra with suturai striae extending along basai margins. Punctures 
margining mesocoxal lines coarse and extended laterally along 

mesepimera S. excisa sp. n. 

Elytra with suturai striae not extending along basai margins. Punctures 
margining mesocoxal lines not extending laterally and usually very fine .... 8 

8 Hypomeron with longitudinal stria. Maie with apicomedian surface of 
metasternum completely covered by conspicuously dense patch of 

pubescence S. pubiventris sp. n. 

Hypomeron lacking stria. Maie with pubescence not completely 
covering apicomedian surface of metasternum 9 

9 Elytra with suturai striae shortened, starting clearly posterior pronotal 

lobe S. monticola sp. n. 

Elytra with suturai striae starting near basai margins and usually curved 
along pronotal lobe 10 

1 Thorax and elytra lacking obvious microsculpture, not iridescent 11 

Thorax and/or elytra microsculptured, elytra usually distinctly iridescent ... 1 3 

11 Metepisterna about 0.05 mm wide, with straight suture. Punctures bor- 

dering mesocoxal lines fine S. davaoana sp. n. 

Metepisterna about 0.08-0.14 mm wide, with arcuate suture. Punctures 
bordering mesocoxal lines coarse 12 

12 Elytra with parasutural striae. Antennomere VIII about 2.5 times as long 
as wide. Metepisterna conspicously large, 0.10-0.14 mm wide, with 

suture strongly arcuate S. episternalis sp. n. 

Elytra without parasutural striae. Antennomere VIII conspicuously 
small, only slightly longer than wide. Metepisterna 0.08-0.10 mm wide, 

with suture weekly arcuate S. serpentis sp. n. 

13 Médian lobe of aedeagus without prominent ventral processes or tubercles . 14 
Médian lobe of aedeagus with prominent ventral processes or tubercles .... 17 

14 Aedeagus with flagellum of internai sac sinuate, not forming circles 15 

Aedeagus with flagellum of internai sac forming complète circle 16 

15 Elytra with distinct parasutural striae. Metepisterna very narrow. Fla- 
gellum lacking basai hook S. werneri sp. n. 

Elytra without or with hardly visible parasutural striae. Metepisterna 
fairly wide. Flagellum with basai hook S. hamata sp. n. 

16 Aedeagus with flagellum of internai sac evenly narrow, except at 
abruptly widened base. Parameres shorter than basai bulb of médian lobe 

S. bulbosa sp. n. 

Aedeagus with flagellum of internai sac gradually narrowed apically, 
without abruptly widened base. Parameres longer than basai bulb of 
médian lobe S. sabapensis Lôbl 

17 Aedeagus with ventral processes of médian lobe approximate. Para- 
meres not widened apically S. orousseti sp. n. 



698 



[. LÔBL 



Aedeagus with ventral processes of médian lobe distant. Parameres 

widened apically 18 

18 Aedeagus with flagellum of internai sac strongly sinuate, weakly 

widened basally, without basai hook S. pseudotenella sp. n. 

Aedeagus with flagellum of internai sac weakly sinuate, strongly 
widened basally, with basai hook S. data sp. n. 

Scaphobaeocera bulbosa sp. n. Figs 1, 2 

HOLOTYPE: S, Luzon, Lagunas Prov., Mt. Makiling, 400m, summit road, 19.XI.1995, 
I. Lôbl (MHNG). 

PARATYPES: 1 9 with the same data as the holotype; Lagunas Prov., Mt. Makiling, 
summit rd., 600m, 21.XI.95, I. Lôbl. - 1 9, Lagunas Prov, Mt. Makiling, summit rd., 600m, 
21-22.XI.1995, I. Lôbl. - 1 d, Lagunas Prov, Mt. Makiling, above Mad Springs, 400-700m, 
19-22.XI.1995, J. Kodada leg. -16, Lagunas Prov, Mt. Banahaw ca 1 km Kinabuhayan, 500m, 
26.XI.1995, 1. Lôbl leg (ail MHNG). 

Description: Length 1.05-1.25 mm, width 0.57-0.70 mm, dorsoventral 
diameter 0.59-0.72 mm. Head and body very dark, almost blackish, femora rufous, 
apical abdominal segments and remainder of appendages lighter, ochraceous. Thorax 
and elytra microsculptured, elytra very weakly iridescent. Pronotal and elytral punc- 
tation very fine, distinct at magnification 50x, punctation becoming somewhat coarser 
toward elytral apices. Length ratio of antennomeres as: III 5: IV 8: V 10: VI 9: VII 10: 
VIII 9: IX 12: X 12: XI 15. Segments III to VI equally narrow, segment III about twice 
as long as wide, segment V about 4 times as long as wide. Segments VII to IX each 
about 3 times as long as wide, segment VIII wider than segment VI. Segment XI 
almost 3 times as long as wide. Tip of scutellum exposed. Elytra with parasutural striae 
hardly visible, suturai striae starting at margin of pronotal lobe, slightly curved at base. 
Hypomeron lacking longitudinal stria, with striate microsculpture. Middle part of 
metaventrite flat, with small, shallow impression in middle, densely and finely punc- 
tate, and short pubescence. Sides of metaventrite sparsely, extremely finely punctate, 
with long pubescence. Mesocoxal lines with very fine marginal punctures not 
extending laterally along mesepimera; submesocoxal areas about 0.03-0.05 mm long. 
Metepisterna flat, exposed portion about 0.06 mm wide, parallel-sided, with straight 
suture. Abdomen with fairly distinct striate microsculpture, very finely punctate. Basai 
punctures of ventrite 1 very fine. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 slightly widened, much narrow- 
er than protibiae. Aedeagus (Figs 1, 2) 0.28-0.31 mm long. 

Habitat: Degraded evergreen rain fbrest, in leaf litter and samples of moss and 
epiphytes on logs, under bark. 

Distribution: Philippines: Luzon. 

COMMENTS: This species is in external characters very similar to S. sabapensis 
Lôbl, 1990. It may be distinguished from the latter by the aedeagus with shorter and 
thicker apical part of the médian lobe, shorter parameres, and distinctive shape of the 
flagellum. 

Scaphobaeocera complicans sp. n. Figs 3-6 

HOLOTYPE: â, Luzon, Lagunas Pref, Mt. Makiling, summit rd.. ca 600m, 26.XI.1995, 
I. Lôbl (MHNG). 



SCAPHISOMATINI OF THE PHILIPPINES 



699 




Figs 1-6 

(1,2) Scaphobaeocera bulbosa sp. n., aedeagus in dorsal and latéral views; scale bar = 0.1 mm. 
(3-6) Scaphobaeocera complicans sp. n., aedeagus (3, 4) and internai sac (5, 6) in dorsal and 
latéral views. Scale bars = 0.2 mm for aedeagus, = 0.1 mm for internai sac. 



700 



[. LÔBL 



Paratypes: 7 â, 13 9, with the same data as the holotype. - 1 6, 
Philippines,Mt. Makiling, Lagunas Prov. 4 km SE Los Banos, 12-IV-1977 / berlese 
rotten logs L. E. Watrous. - 1 S with the same data but 9-IV-1977 and berlese rotten 
figs (ail MHNG). 

Description: Length 1.05-1.20 mm, width 0.57-0.68 mm, dorsoventral diam- 
eter 0.65-0.75 mm. Body rufous, apical abdominal segments and appendages light 
ochraceus to yellowish. Thorax, elytra and abdominal ventrite 1 not microsculptured 
and not iridescent. Pronotal and elytral punctation very fine, that on elytra slightly 
coarser than on pronotum. Length ratio of antennomeres as: III 4: IV 8: V 9: VI 7: VII 
10: VIII 6: IX 9: X 10: XI 26. Segment III short and narrow, about twice long as wide. 
Segments IV as narrow as segment III. Segments V and VI slightly wider than segment 
IV, segment V about 3 to 4 times as long as wide. Segment VII about 2.5 times as long 
as wide. Segment VIII twice as long as large. Segments IX and X slightly wider than 
segment VII. Segment XI conspicuously long, about 5 times as long as wide. Tip of 
scutellum exposed. Elytra with very fine parasutural striae, suturai striae starting at 
margin of pronotal lobe, slightly curved at base. Hypomeron with stria. Middle part of 
metaventrite hardly convex, lacking stria or impression, with dense and fairly fine 
punctation except on narrow anterior area, and with fairly long pubescence. Sides of 
metaventrite sparsely and finely punctate, with long pubescence. Mesocoxal lines with 
fine marginal punctures, not extending laterally along mesepimera; submesocoxal 
areas about 0.03 mm long. Metepisterna flat, with exposed portion about 0.06-0.08 mm 
wide, parallel-sided, with straight suture. Abdominal ventrite 1 very finely punctate, 
with basai punctures fine, partly somewhat elongate. Following sternites with punctu- 
late microsculpture. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 moderately widened, narrower 
than protibiae. Aedeagus (Figs 3-6) 0.44-0.53 mm long. 

Habitat: Evergreen rain forest, fungi on large log, in rotten log and on rotten 

figs. 

Distribution: Philippines: Luzon. 

COMMENTS: Several members of Scaphobaeocera have conspicuously long 
apical antennomere, compared to the antennomeres VII and X (S. abnormalis Lôbl, 
1981, S antennalis Lôbl. 1975, S. cyrta Lôbl, 1980, S delicatula Lôbl, 1971, S. dispar 
Lôbl, 1980, S dorsalis Lôbl, 1980, S. japonica (Reitter, 1880), S minutissima (Lôbl, 
1969), S. ponapensis Lôb\, 1981, S. remota Lôbl, 1981, and S stephensoni Lôbl, 1988). 
Scaphobaeocera complicans may be easily distinguished from those species by the 
shape of the internai sac of the aedeagus, in particular by its strongly expanded apical 
part and the bulbous basai part. 

Scaphobaeocera data sp. n. Figs 7-10 

HOLOTYPE: â. Luzon Mount Data Lodge 2200-2300m 22-23.XII.1979, L. Deharveng 
& J. Orousset #100 (MHNG). 

PARATYPES: 26 â, 15 9, with the same data as the holotype; 2 S, with the same data 
but #86. - 1 9,1 9 , with the same data but #162; Mount Data, 9.1.80, J. Orousset #165.-3 «5, 
5 9, Luzon, Mt. Data (2250m) Mountain Prov., 13., 14., 25. and 26. VII.1985, M. Sakai leg. - 
2 S, Luzon Mt. Fangao, (2350m) Mountain Prov., 14.VII.1985, M. Sakai leg. - 2 6, 14 9, 
Pacay (2400m) nr. Sayangan Benguet Prov., il. VII. 1985, M. Sakai leg. (ail MHNG). 



SCAPHISOMATINI OF THE PI III IPPINl S 



701 




FlGS 7-10 

(7-10) Scaphobaeocera data sp. n., aedeagus (7, 9) and internai sac (8, 10) in dorsal and latéral 
views. 

Description: Length 1.15-1.30 mm, width 0.65-0.73 mm, dorsoventral 
diameter 0.70-0.78 mm. Body very dark reddish-brown to almost black, apices of 
elytra usually lighter, apical abdominal segments and appendages ochraceous. Thorax, 
elytra and abdomen microsculptured and iridescent. Pronotal and elytral punctation 
very fine, hardly visible at magnification lOOx. Length ratio of antennomeres as: III 6: 
IV 8: V 10: VI 8: VII 10: VIII 7: IX 10: X 10: XI 16. Segments III and IV equally 
narrow, segment III about 3 times as long as wide. Segments V and VI wider than 
segment IV, each about 3 times as long as wide. Segment VII almost 3 times as long 
as wide. Segment VIII slightly narrower than segment VII, about twice as long as wide. 
Segments IX and X distinctly wider than segment VII, each about twice as long as 
wide. Segment XI about 2.5 times as long as wide. Scutellum entirely concealed. 
Elytra with very fine parasutural striae, suturai striae starting at margin of pronotal 
lobe, slightly curved at base. Hypomeron without stria. Middle part of metaventrite 
flat, with shallow stria, very dense and fine punctation, and very short pubescence. 
Sides of metaventrite sparsely and very finely punctate, with long pubescence. 
Mesocoxal lines with fine marginal punctures, not extending laterally along 
mesepimera; submesocoxal areas about 0.02-0.03 mm long. Metepisterna flat, with 
exposed portion about 0.04-0.06 mm wide, parallel-sided, with straight suture. 



702 



I. LÔBL 



Abdomen with distinct microsculpture consisting of transverse striae. Abdominal 
ventrite 1 very finely punctate, with basai punctures fine, not elongate. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 distinctly widened, narrower 
than protibiae. Aedeagus (Figs 7-10) 0.36-0.40 mm long. 

Habitat: Mountain broad-leaf forest, under bark, in rotten wood and floor 

litter. 

Distribution: Philippines: Luzon. 

COMMENTS: As in S. data, the Asian S. cyrta Lôbl, 1980, S. discreta Lôbl, 1980 
and S. smetanai Lôbl, 1981 possess flagellar basai hook and the médian lobe of the 
aedeagus is lacking ventral processes. The new species may be distinguished from 
them by the convex apicoventral side of the basai bulb. Besides, S. smetanai may be 
separated by the robust basai part of the internai sac, S. discreta by the parameres 
gradually widened apically (in latéral view) and S. cyrta by the médian lobe conspi - 
cuously arcuate, with tip reaching beyond parameres. The Papuan and Australian 
S. papuana Csiki, 1909, S. ornata (Pic, 1956), S. piceoapicalis Lôbl, 1977 and S. 
queenslandica Lôbl, 1986 have also aedeagi comparatively similar to that in S. data. 
The first three of them may be readily distinguished by their distinctive body col- 
oration, S. queenslandica differs drastically by the shape of the parameres. Other Asian 
species that possess an internai sac with a basai hook are those originally described as 
Baeotoxidium Lôbl, 1971, and since transferred to Scaphobaeocera: S. elegans (Lôbl, 
1971), S. gagata (Lôbl, 1971), S. indica (Lôbl, 1979), S. lanka (Lôbl, 1971), S. sia- 
mensis (Lôbl, 1990), and S. yeti (Lôbl, 1992). Thèse species are linked by the internai 
sac of the aedeagus with a basai vesicle containing spine-like structures, and probably 
form a monophyletic group. 

Scaphobaeocea davaoana sp. n. Figs 11, 12 

HOLOTYPE: â, E. slope Mt. McKinley, Davao Prov., MINDANAO IX. 1946 / Elévation 
6400 ft. / CNHM - Philippines Zool. Exped. (1946-47) F. G. Werner leg. (FMNH). 

Description: Length 1.15 mm, width 0.62 mm, dorsoventral diameter 0.55 
mm. Head and body uniformly ochraceous, appendages lighter. Thorax and elytra 
lacking microsculpture, not iridescent. Pronotal and elytral punctation extremely fine, 
barely visible at magnification 160x. Length ratio of antennomeres as: III 6: IV 7: V 8: 
VI 7: VII 11: VIII 5: IX 10: X 10: XI 14. Segment III comparatively short, about 3 
times as long as wide. Segments IV to VI narrow, as wide as segment III. Segment VII 
almost 3 times as long as wide. Segment VIII slightly wider than segment VI, almost 
twice as long as wide. Segments IX and X distinctly larger than segment VII. Segment 
XI not widened apically, about 2.5 times as long as wide. Tip of scutellum exposed. 
Elytra without parasutural striae, suturai striae starting at margin of pronotal lobe, 
hardly curved at base. Hypomeron lacking longitudinal stria. Middle part of meta - 
ventrite flat, lacking stria or impression, coarsely and densely punctate, with long 
pubescence. Sides of metaventrite lacking microsculpture, sparsely, extremely finely 
punctate, with long pubescence. Mesocoxal lines with fine marginal punctures not 
extending laterally along mesepimera; submesocoxal areas about 0.03 mm long. 
Metepisterna flat, exposed portion about 0.05 mm wide, almost parallel-sided, with 
straight suture. Abdomen with striate microsculpture. Tibiae straight. 



SCAPHISOMATINI OF THE PHILIPPINES 



703 




Figs 11-13 

(11, 12) Scaphobaeocera davaoana sp. n., aedeagus in dorsal and latéral views. (13) Scapho - 
baeocera epistemalis sp. n., aedeagus in dorsal view. Scale bars = 0. 1 mm. 

Maie characters. Protarsi with segments 1 to 3 strongly widened, about as wide 
as protibiae. Aedeagus (Figs 11, 12) 0.37 mm long. 

Distribution: Philippines: Mindanao. 

COMMENTS: The species resembles S. nuda Lôbl, 1979 in external characters. 
Both possess not microsculptured thorax and elytra, very fine punctation, fairly similar 
antennae, and large exposed portion of metepisterna. Scaphobaeocera davaoana dif- 
fers from S. nuda by the elytra not darkened at apex and lacking parasutural striae, and, 
more drastically, by its aedeagal characters. The new species may be distinguished 
from its congeners by following characters in combination: médian lobe with robust, 
strongly prominent ventral processes, flagellum very narrow, and parameres wide in 
dorsal view, tapering in latéral view. The shape of the médian lobe is similar to that in 
S. uncata Lôbl, 1990, while the apically widened parameres differ conspicuously. 
Similar ventral processes are présent in B. amicalis Lôbl, 2003, B. stipes Lôbl, 1971, 
B. tenella Lôbl, 1990 and B. uncata Lôbl, 1990. Scaphobaeocera amicalis and B. un- 
cata differ conspicuously by the distal part of the médian lobe strongly arcuate and 
reaching beyond tip of the parameres, B. stipes and B. tenella have the parameres 
widened in apical half and distinctive internai sac. 



704 



[. LÔBL 



Scaphobaeocera episternalis sp. n. Figs 13-16 

HOLOTYPE: 6 , Mindanao, Davao Prov., 25 km W of New Batan 20-22 May 1996, Bolm 
leg. (SMNS). 

PARATYPES: 1 â, with the sama data as the holotype (MHNG). -19, Mindanao, 30 km 
NW of Maramag, 13-17 May 1996 Bagongsilang, 1700m, Bolm leg. (SMNS). 

DESCRIPTION: Length 1.23-1.26 mm, width 0.72-0.75 mm, dorsoventral 
diameter 0.72-0.75 mm. Head, most of body, femora and tibiae light rufous, apices of 
elytra, apex of abdomen and legs lighter, ochraceous, tarsi and antennomeres I to V or 
VI yellowish, antennomers VII to XI brown. Pronotum lacking microsculpture. Elytra 
microsculptured and weakly iridescent. Pronotal and elytral punctation sparse and 
extremely fine, hardly visible at magnification lOOx. Length ratio of antennomeres as: 
III 8: IV 8: V 9: VI 8: VII 11: VIII 8: IX 12: X 12: XI 20. Segments III and IV equal, 
each about 3 times as long as wide. Segments V and VI slightly wider than segment IV, 
segment V 3 times as long as wide. Segment VII almost 3 times as long as wide. 
Segment VIII slightly wider than segment VI, about 2.5 times as long as wide. 
Segments IX and X each distinctly wider than segment VIII. Segment XI parallel- 
sided, distinctly wider than segment X, somewhat more than 3 times as long as wide. 
Tip of scutellum exposed. Elytra with parasutural striae, suturai striae starting at 
margin of pronotal lobe, slightly curved at base. Hypomeron lacking stria. Middle part 
of metaventrite convex, lacking stria or impression, smooth on large central part, with 
dense and fairly coarse punctation at each side of center and with fairly long pubes- 
cence. Sides of metaventrite sparsely and very finely punctate, with long pubescence. 
Mesocoxal lines with fairly coarse marginal punctures not extending laterally along 
mesepimera; submesocoxal areas about 0.02 mm long. Metepisterna flat, with exposed 
portion about 0.10-0.14 mm wide, widest in anterior half, suture broadly arcuate 
toward anterior angles, slightly curved or oblique apically. Abdomen lacking obvious 
microsculpture and with punctation hardly visible. Abdominal ventrite 1 with basai 
punctures coarse, not elongate. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 strongly widened, segment 1 
slightly wider than protibia. Aedeagus (Figs 13-16) 0.57-0.58 mm long. 

Distribution: Philippines: Mindanao. 

COMMENTS: This species may be readily distinguished from its Philippine 
congeners by its large metepisterna. It differs from most other species by the médian 
lobe of the aedeagus that is not bent apically and has no distinct apicoventral processes. 
The internai sac with a complex base and very narrow flagellum reminds that in S. 
tibialis Lôbl, 1984, while the parameres are wider and not arcuate as in the latter 
species. 

Scaphobaeocea escensa sp. n. Figs 17-19 

HOLOTYPE: S, Mindanao 30 km NW of Maramag, 13-17 May 1996, Bagongsilang, 
1700m, Bolm leg. (SMNS). 

PARATYPE: â, with same data as holotype (MHNG). 

Description: Length 1.70 mm, width 1.0 mm, dorsoventral diameter 1.08 mm. 
Head and most of body uniformly very dark brown to blackish, abdomen and appen - 
dages reddish-brown. Pronotum lacking microsculpture, elytra with microsculpture 



SCAPHISOMATINI OF THE PHILIPPINES 



705 




Figs 14-21 



(14-16) Scaphobaeocera episternalis sp. n., aedeagus in latéral view, paramere (15) and internai 
sac (16) enlarged. (17-19) Scaphobaeocera escensa sp. n., antennomeres III to VIII (17), aedea- 
gus in dorsal and latéral views. (20, 21) Scaphobaeocera excisa sp. n., aedeagus in dorsal view 
(20), paramere in ventral view, enlarged. Scale bars = 0. 1 mm. 



706 



[. LÔBL 



hardly visible, not iridescent. Punctation extremely fine on pronotum, hypomera and 
latéral parts of metaventrite. Elytral punctation scattered, punctures much larger those 
on pronotum. Length ratio of antennomeres (Fig. 17) as: III 4: IV 8: V 12: VI 6: VII 
30: VIII 5: IX 26: X 26: XI 26. Segment III comparatively short and wide, about 1.5 
times as long as wide. Segment IV slightly narrower than segment III, about twice as 
long as wide. Segment V hardly wider than segment IV, 3 times as long as wide. 
Segment VI very short, gradually widened apically, wider than preceding segments, 
slightly longer than wide. Segment VII conspicuously long, more than 6 times as long 
as wide. Segment VIII very small, about as wide as segment VII and as wide as long. 
Segments IX and X similar to segment VII. Segment XI slightly widened apically. Tip 
of scutelum exposed. Elytra with suturai striae starting at pronotal lobe, parasutural 
striae présent. Hypomeron not microsculptured, lacking longitudinal stria. Middle part 
of metaventrite flattened, with very short and shallow médian stria, impunctate in cen- 
ter, distinctly punctate and with long pubescence laterally and behind center. 
Mesocoxal lines with very fine marginal punctures not extending laterally along 
mesepimera; submesocoxal areas about 0.02 mm long. Metepisterna flat, about 0.05 
mm wide, parallel-sided, with straight suture. Abdominal ventrite 1 with basai punc- 
tures fairly fine, partly elongate. Protibiae straight, meso and metatibiae weakly 
curved. Ventral side of profemora each with short, arcuate comb situated in level of 
trochanters. 

Maie characters. Protarsi with segment 1 to 3 strongly widened. Aedeagus (Figs 
18, 19) 0.59-0.62 mm long. 

Distribution: Philippines: Mindanao. 

COMMENTS: The very long antennomere VII, in combination with the small an- 
tennomere VIII, is diagnostic for this new species. Scaphobaeocera zdenae Lôbl, 1992 
has also strongly enlarged antennomere VII, but the antennomere VIII twice as long as 
wide. Besides, thèse two species may be easily distinguished by the pronotum and ely- 
tra distinctly microsculptured and iridescent in B. zdenae while they are not iridescent 
in S. es cens a. 

Scaphobaeocera excisa sp. n. Figs 20-23 

HOLOTYPE: 6, Luzon, Baguio, Mt. Santo Thomas, ca 1850m, 14.1.1980, L. Deharveng 
& J. Orousset leg. #191 (MHNG). 

PARATYPES: 3 â, with same data as the holotype; 2 â, Luzon, Baguio, Mt. Santo 
Thomas, ca 2150m, 14.1.1980, L. Deharveng & J..Orousset leg. #198 (ail MHNG). 

Description: Length 0.95-1.05 mm, width 0.60-0.63 mm, dorsoventral 
diameter 0.54-0.55 mm. Head and body ochraceous to blackish, appendages lighter. 
Thorax and elytra lacking microsculpture, not iridescent. Pronotal and elytral punc- 
tation very fine, dense, visible at magnification 32x. Length ratio of antennomeres as: 
III 6: IV 6: V 6: VI 6: VII 10: VIII 5: IX 9: X 9: XI 13. Segments III and IV similar, 
very narrow, each about 4 times as long as wide. Segments V and VI slightly wider 
than segment IV. Segment VII about 2.5 times as long as wide. Segment VIII about 1.3 
times as long as wide. Segments IX and X hardly wider than segment VII. Segment XI 
not widened apically, almost 3 times as long as wide. Tip of scutellum exposed. Elytra 
without parasutural striae, suturai striae curved at base and extending along basai 



SCAPHISOMATINI OF THE PHILIPPINES 



707 



margin to form short basai striae, ending in inner halves of basai elytral width. 
Hypomeron not microsculptured, lacking longitudinal stria. Middle part of meta- 
ventrite flat, lacking stria or impression, coarsely and dcnsely punctate, with short 
pubescence. Sides of metaventrite lacking microsculpture, sparsely, extremely finely 
punctate, with long pubescence. Mesocoxal lines with coarse marginal punctures ex- 
tending laterally along mesepimera; submesocoxal areas about 0.03 mm long. 
Metepisterna slightly convex, exposed portion about 0.05 mm wide, with arcuate 
suture. Abdomen with hardly visible punctulate microsculpture. Abdominal punctation 
fine, except comparatively distinct punctation on propygidium. Basai punctures of 
ventrite 1 partly elongate and coarse. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 slightly widened, much narrower 
than protibiae. Aedeagus (Figs 20-23) 0.35-0.38 mm long. 

HABITAT: Moist ravine, in forest litter. 

Distribution: Philippines: Luzon. 

Comments: The elytra with extended suturai striae and the shape of the para- 
meres are diagnostic for this species. Scaphobaeocera incisa Lôbl, 1990 has also emar- 
ginate parameres, but the emargination is subapical. Besides, it differs conspicuously 
in other aedeagal characters. 

Scaphobaeocera hamata sp. n. Figs 24-26 

HOLOTYPE: â, Philippines: Mt. Makiling, Laguna Prov., 4 km SE Los Banos 12-IV- 
1977 / berlese rotten logs L. E. Watrous (MHNG). 

PARATYPES: 8 â, 3 9, with the same data as the holotype. -3 6\ 1 9, with the same 
data but 9-IV-1977. - 1 ô\ 19, with the same data but 8-IV-1977 / berlese leaf litter; 1 9, with 
the same data but 8-IV-1977 / berlese litter along stream. - 2 â , 1 9, with the same data but 
1 1 -IV- 1977 / berlese débris under bark (ail MHNG). 

DESCRIPTION: Length 1.25-1.30 mm, width 0.66-0.70 mm, dorsoventral dia- 
meter 0.66-0.72 mm. Head and body ochraceous, appendages lighter, elytra sometimes 
slightly darkened near apex. Thorax and elytra microsculptured, elytra and often also 
pronotum weakly iridescent. Pronotal and elytral punctation very fine, dense, visible at 
magnification 32x. Length ratio of antennomeres as: III 5: IV 6: V 9: VI 8: VII 1 1 : VI- 
II 7: IX 11: X 10: XI 15. Segments III and IV similar, narrow, respectively 2.5 and 3 
times as long as wide. Segments V and VI slightly wider than segment IV. Segment VII 
about 3 to 3.5 times as long as wide. Segment VIII about as wide as segment VII and 
almost twice as long as wide. Segments IX and X much wider than segment VII. 
Segment XI not widened apically, about 2.5 times as long as wide. Tip of scutellum ex- 
posed. Elytra without or with hardly visible parasutural striae, suturai striae starting at 
pronotal lobe. Hypomeron not microsculptured, lacking longitudinal stria. Middle part 
of metaventrite flat, with minute impression, densely punctate, pubescence short and 
very dense. Sides of metaventrite microsculptured, sparsely, extremely finely punctate, 
with fairly long pubescence. Mesocoxal lines with fine marginal punctures not 
extending laterally along mesepimera; submesocoxal areas about 0.01-0.02 mm long. 
Metepisterna slightly convex, exposed portion 0.05-0.06 mm wide, narrowed proxi- 
mally, with straight suture. Abdomen with hardly visible striate microsculpture, punc- 
tation very fine. Basai punctures of ventrite 1 not elongate and fine. Tibiae straight. 



708 



I. LÔBL 



Maie characters. Protarsi with segments 1 to 3 strongly widened, almost as wide 
as protibiae. Aedeagus (Figs 24-26) 0.32-0.42 mm long. 

Habitat: Evergreen rain forest, under bark, in rotten wood and leaf litter. 

Distribution: Philippines: Luzon. 

Comments: The aedeagal characters suggest relationship to S. data and S. mon- 
ticola. Scaphobaeocera hamata may be readily distinguished from the former by the 
lighter body color and from the latter by the suturai striae of the elytra reaching elytral 
base. See also comments under S. data. 

Scaphobaeocera monticola sp. n. Figs 27-30 

HOLOTYPE: S, Philippines: Luzon, Baguio, Mt. Santo Thomas, ca 2150m, 14.1.80, L. 
Deharveng & J. Orousset leg. # 198 (MHNG). 

PARATYPES: 1 â, 1 ? , with the same data as the holotype; 1 6, Luzon, Mountain Prov., 
Mount Data Lodge 2250-2300m, 23-24.XII.79, L. Deharveng & J. Orousset (MHNG). 

Description: Length 1.20 mm, width 0.71-0.76 mm, dorsoventral diameter 
0.71-0.75 mm. Body very dark reddish-brown to almost black, apices of elytra usually 
lighter, apical abdominal segments, legs and antennomeres I and II lighter, reddish- 
brown to ochraceous, antennomeres III to XI yellowish. Thorax, elytra and abdomen 
microsculptured and iridescent. Microsculpture on venter of thorax conspicuously 
coarse. Pronotal and elytral punctation very fine, hardly visible at magnification lOOx. 
Length ratio of antennomeres as: III 5: IV 7: V 10: VI 8: VII 10: VIII 6: IX 10: X 10: 
XI 13. Segments III to VI evenly narrow, segment III about 2.5 times as long as wide, 
segments IV well 3 times as long as wide, V about 5 times as long as wide, VI about 4 
times as long as wide. Segment VII about twice as wide as segment VI and almost 3 
times as long as wide. Segment VIII slightly narrower than segment VII, about twice 
as long as wide. Segments IX and X wider than segment VII, each about 2.5 times as 
long as wide. Segment XI about 3 times as long as wide and as wide as segment X. 
Scutellum entirely concealed. Elytra with very fine parasutural striae, suturai striae 
starting posterior margin of pronotal lobe, not curved at base. Hypomeron without 
stria. Middle part of metaventrite flat, slightly impressed apically, without stria, with 
moderately dense, fine punctation and very short pubescence. Sides of metaventrite 
extremely finely punctate, with very short pubescence. Mesocoxal lines with very fine 
marginal punctures not extending laterally along mesepimera; submesocoxal areas 
about 0.02-0.03 mm long. Metepisterna flat, with exposed portion about 0.04-0.05 mm 
wide, parallel-sided, with straight, deep suture. Abdomen with distinct microsculpture 
consisting of transverse striae. Abdominal ventrite 1 extremely finely punctate, with 
basai punctures fine, not elongate. Tibiae straight 

Maie characters. Protarsi with segments 1 to 3 distinctly widened, narrower 
than protibiae. Aedeagus (Figs 27-30) 0.39-0.41 mm long. 

Habitat: Mountain broad-leaf forest, in rotten wood and humus, in moss on 

log. 

Distribution: Philippines: Luzon. 



SCAPHISOMATINI OF THE PHILIPPINES 



709 





Figs 22-30 

(22, 23) Scaphobaeocera excisa sp. n., aedeagus in latéral view (22), internai sac in dorsal view, 
enlarged (23). (24-26) Scaphobaeocera hamata sp. n., aedeagus in dorsal and latéral views, 
internai sac (26) enlarged. (27-30) Scaphobaeocera monticola sp. n., aedeagus in dorsal and 
latéral views, internai sac (29, 30) enlarged. Scale bars = 0.1 mm. 



710 



[. LÔBL 



COMMENTS: This species is characterized by the elytra with shortened suturai 
striae. In addition, its comparatively coarse thoracic microsculpture is diagnostic. The 
aedeagus in 5. monticola is similar to that in S. data, see also comments under the latter 
species. 

Scaphobaeocera montivagans sp. n. 

HOLOTYPE: 6 , Luzon: Philippines Mt. Pangao (2350m) nr. Data Ifugao Pv. 14.VII.1985, 
M. Sakai leg. (MHNG). 

PARATYPE: o\ Luzon: Mount Data 8.1.80. J. Orousset #165 (MHNG). 

Description: Length 1.30-1.40 mm. width 0.76-0.82 mm, dorsoventral 
diameter 0.82-0.87 mm. Head and body dark reddish-brown, apical part of abdominal 
ventrue 1 and following ventrues lighter, reddish to ochraceous, appendages light 
reddish-brown to yellowish. Pronotum and hypomera lacking microsculpture, elytra, 
latéral parts of metaventrite. mesepimera. metepisterna and abdominal ventrites dis- 
tinctly microsculptured. Elytra iridescent. Pronotal and elytral punctation very fine, 
hardly visible at magnification lOOx. Length ratio of antennomeres as: III 5: IV 8: V 
10: VI 7: VU 10: VIO 7: IX 1 1 : X 10: XI 22. Segment III thickened apically, segments 
IV and V slightly narrower than segment III, IV about 4 times as long as wide, V about 
5 times as long as wide. Segment VI distinctly wider than segment V, about twice as 
long as wide. Segment VII about twice as long as wide. Segment VIII slightly narrower 
than segment VII. almost twice as long as wide. Segments IX to XI about as wide as 
segment VII, IX and X each about twice as long as wide, XI about 4.5 times as long as 
wide. Tip of scutellum exposed. Hypomeron with stria. Elytra with very fine para- 
sutural striae. suturai striae starting near base, slightly curved along pronotal lobe. 
Middle part of metaventrite somewhat convex, not impressed apically, without stria, 
with dense, fairly coarse punctation and rather long pubescence. Sides of metaventrite 
extremely finely punctate. with fairly long pubescence. Mesocoxal Unes with very fine 
marginal punctures not extending laterally along mesepimera; submesocoxal areas 
about 0.02-0.03 mm long. Metepisterna flat, with exposed portion about 0.08-0.09 mm 
wide, parallel-sided. with straight. deep suture. Abdomen with distinct microsculpture 
consisting of transverse striae. Abdominal ventrite 1 extremely finely punctate, with 
basai punctures fine, not elongate. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 distinctly widened. narrower 
than protibiae. 

Habitat: Mountain broad-leaf forest. leaf and wood litter. 
DISTRIBUTION: Philippines: Luzon. 

COMMENTS: This species resembles S. complicans from which it may be easily 
distinguished by the microsculpture pattern on the ventral side of body. The aedeagi of 
both maies were lost. Nevertheless, the external characters of 5. montivagans are un- 
ambiguously diagnostic. 

Scaphobaeocera orousseti sp. n. Figs 31-34 

HOLOTYPE: 6. Luzon. Sagada s/résourgence d'Ambasing. 16. XII. 79. L. Deharveng & 
J. Orousset #233 (MHNG). 



SCAPHISOMATINI OF THE PHILIPPINES 



711 



PARATYPES: 1 d, Luzon, Mountain Prov. N. & NE Sagada, 1 5- 19. XI 1.1979, L. 
Deharveng & J. Orousset #39. - 6 d, 2 9, Luzon, Mountain Prov. Sagada, Suyo: Tataya-An, 
20.1.1980, L. Deharveng & J. Orousset # 201 (ail MHNG). 

Description: Length 1.18 mm, width 0.65 mm, dorsoventral diameter 0.68 
mm. Body very dark reddish-brown, apices of elytra, apical abdominal segments and 
appendages light ochraceus to yellowish. Thorax, elytra and abdomen microsculptured, 
pronotal microsculpture barely visible. Pronotum not iridescent, elytra iridescent. 
Pronotal and elytral punctation very fine, distinct at magnification 50x. Length ratio of 
antennomeres as: III 6: IV 8: V 9: VI 8: VII 10: VIII 8: IX 10: X 10: XI 12. Segment 

III about 3 times as long as wide. Segment 4 about as narrow as segment III, segments 
V and VI slightly wider, each about 3 times as long as wide. Segment VII 3 times as 
long as wide. Segment VIII slightly wider than segment VI, about 3 times as long as 
wide. Segments IX to XI distinctly wider than segment VII, segment XI about 2.5 
times as long as wide. Scutellum entirely concealed. Elytra with parasutural striae ob- 
solète, suturai striae starting at margin of pronotal lobe, slightly curved at base. 
Hypomeron without stria. Middle part of metaventrite flattened, with very shallow, 
minute impression in middle, dense and fairly fine punctation except on anterior area, 
and fairly long pubescence. Sides of metaventrite sparsely and very finely punctate, 
with long pubescence. Mesocoxal lines with fine marginal punctures, not extending 
laterally along mesepimera; submesocoxal areas about 0.03 mm long. Metepisterna 
somewhat convex, with exposed portion about 0.06 mm wide, parallel-sided, suture 
deep and straight. Abdominal ventrite 1 very finely punctate, with basai punctures 
fairly coarse and elongate. 

Maie characters. Protarsi with segments 1 to 3 moderately widened, narrower 
than protibiae. Aedeagus (Figs 31-34) 0.60 mm long. 

Habitat: Ravin, in leaf litter and humus. 

Distribution: Philippines: Luzon. 

COMMENTS: This species is similar to S. data in external characters but may be 
distinguished by the elongate punctures at the base of the abdominal ventrite 1 . See al- 
so comments under S. davaoana. 

ETYMOLOGY: The species is named in honour of one of its collectors, Jean 
Orousset, Paris. 

Scaphobaeocera palawana sp. n. Figs 35, 36 

HOLOTYPE: 6, Palawan, Central, Olangoan, 18 km NE San Rafaël, sea level, I. Lôbl, 
5-6.XII.1995 (MHNG). 

Description: Length 1.05 mm, width 0.60 mm, dorsoventral diameter 
0.60 mm. Head and body very dark reddish-brown, apical abdominal segments and 
appendages lighter. Thorax and elytra lacking microsculpture, not iridescent. Pronotal 
and elytral punctation very fine, visible at magnification lOOx. Length ratio of anten- 
nomeres as: III 5: IV 6: V 7: VI 6: VII 9: VIII 6: IX 9: X 10: XI 19. Segments III and 

IV equally narrow, segment IV 3 times as long as wide. Segments V and VI wider than 
segment IV, segment V about 3 times as long as wide. Segments VII and VIII each 
about twice as long as wide, segment VIII distinctly wider than segment VI. Segments 



712 



[. LÔBL 




Figs 31-38 

(31-34) Scaphobaeocera orousseti sp. n., aedeagus in dorsal and latéral views, internai sac 
(32, 33) enlarged. (35, 36) Scaphobaeocera palawana sp. n., aedeagus in dorsal and latéral 
views. (37, 38) Scaphobaeocera pseudotenella sp. n., aedeagus in dorsal and latéral views. Scale 
bars = 0.1 mm for figs 31, 34-38, scale bar = 0.05 mm for figs 32, 33. 



SCAPHISOMATINI OF THF. PHILIPPINES 



713 



IX and X wider than segment VII, each almost twice as long as wide. Segment XI 
somewhat wider and almost 2 times as long as segment X, about 3 times as long as 
wide. Tip of scutellum exposed. Elytra without parasutural striae, suturai striae starting 
at margin of pronotal lobe, slightly curved at base. Hypomeron lacking longitudinal 
stria. Middle part of metaventrite slightly convex, lacking stria or impression, center 
smooth, finely and densely punctate around center, and with short pubescence. Sides 
of metaventrite sparsely, extremely finely punctate. Mesocoxal lines with very fine 
marginal punctures not extending laterally along mesepimera; submesocoxal areas 
about 0.02 mm long. Metepisterna flat, exposed portion about 0.06 mm wide, parallel- 
sided, with straight suture. Abdominal ventrite 1 lacking microsculpture, very finely 
punctate, with basai punctures hardly visible. Following ventrites with punctulate 
microsculpture. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 distinctly widened, narrower 
than protibiae. Aedeagus (Figs 35, 36) 0.38 mm long. 

Habitat: Edge of cultivated area, fungi on log. 

Distribution: Philippines: Palawan. 

Comments: The aedeagal characters of this species suggest relationship with 
S. nuda Lobl, 1979, although the apical part of the médian lobe and the flagellum are 
notably shorter in the new species than in S. nuda. As S. davaoana, the new species 
lacks thoracic and elytral microsculpture and the hypomeral and parasutural striae, and 
has the exposed part of the metepisterna comparatively large. Scaphobaeocera 
palawana and S. davaoana may be readily distinguished by the length of the apical 
antennomere and the présence or absence of obvious abdominal microsculpture. 

Scaphobaeocera pseudotenella sp. n. Figs 37, 38 

Holotype: S, Luzon, Lagunas Prov., Mt. Makiling, 600m, summit rd, 
21. XI. 1995, leg. I. Lôbl (MHNG). 

Paratypes: 1 9 , with the same data as the holotype; 1 9 , Luzon, Lagunas, Mt. 
Banahaw nr school about 1 km from Kinabuhayan, 500m, 26.XI.1995, leg. J. Kodada 
(both MHNG). 

DESCRIPTION: Length 1.15-1.25 mm, width 0.70 mm, dorsoventral diameter 
0.63-0.70 mm. Head and body uniformly reddish-brown, femora lighter, apical 
abdominal segments, tibiae, tarsi and antennae light brown or yellowish. Pronotum 
lacking microsculpture. Elytra, hypomera, latéral parts of metaventrite, mesepisterna, 
mesepimera and abdomen with distinct microsculpture. Elytra iridescent. Pronotal and 
elytral punctation very fine, barely visible at magnification 50x. Length ratio of anten- 
nomeres as: III 4: IV 6: V 8: VI 8: VII 13: VIII 7: IX 12: X 12: XI 15. Segment III 
short, about twice as long as wide. Segments IV to VI narrow, as wide as segment III, 
segment IV about 3 times as long as wide, segments V and VI each about 4 times as 
long as wide. Segment VII about 3 times as long as wide. Segment VIII about twice as 
long as wide. Segments IX to XI each slightly wider than segment VII, XI 3 times as 
long as wide. Scutellum entirely concealed. Elytra with parasutural striae, suturai striae 
starting at margin of pronotal lobe, hardly curved at base. Hypomeron lacking longitu- 



714 



L. LÔBL 



dinal stria. Middle part of metaventrite flat, with médian stria, distinctly, densely punc- 
tate, with short pubescence. Sides of metaventrite with fairly dense, fine punctation and 
fairly long pubescence. Mesocoxal lines with fine marginal punctures not extending 
latéral ly along mesepimera; submesocoxal areas about 0.02 mm long. Metepisterna 
flat, exposed portion about 0.03-0.04 mm wide, parallel-sided, with straight suture. 
Abdominal microsculpture distinct, ventrite 1 iridescent. Basai punctures of ventrite 1 
not elongate. Tibiae straight in female. 

Maie characters. Protarsi with segments 1 to 3 distinctly widened, narrowed 
than protibiae. Protibiae straight, mesotibiae slightly curved, metatibiae somewhat 
sinuate. Aedeagus (Figs 37, 38) 0.42 mm long. 

Habitat: Degraded evergreen rain forest, leaf litter. 

Distribution: Philippines: Luzon. 

COMMENTS: This species shares most of the aedeagal characters with S. tenella 
Lôbl, 1990. However, it differs from the latter species by the shape of the parameral 
apices which are clearly distinctive in latéral view. See also comments under B. 
davaoana. 

Scaphobaeocera pubiventris sp. n. Figs 39, 40 

Holotype: 6, Luzon, Mountain Prov., N. & NE of Sagada 15-19.XII.1979, 
L. Deharveng & J. Orousset #46 (MHNG). 

Paratypes: 4 â, 6 9 with the same data as the holotype (MHNG). 

DESCRIPTION: Length 1.25-1.40 mm, width 0.70-0.75 mm, dorsoventral dia- 
meter 0.71-0.81 mm. Head and most of body uniformly dark rufous to blackish, apex 
of abdomen and legs lighter, ochraceous, antennae conspicuously light, yellowish. 
Pronotum with microsculpture hardly visible, distinctly iridescent in darker spécimens, 
not or barely iridescent in lighter spécimens. Elytra microsculptured and iridescent. 
Pronotal and elytral punctation very fine, irregular, elytra usually with scattered punc- 
tures larger than those on pronotum. Length ratio of antennomeres as: III 5: IV 8: V 9: 
VI 7: VII 12: VIII 7: IX 12: X 12: XI 17. Segment III comparatively short and wide, 
about twice long as wide. Segments IV to VI about as wide as segment III, segment V 
3 times as long as wide. Segment VII about twice as long as wide. Segment VIII about 
1.5 times as long as wide. Segments IX and X hardly larger than segment VII. Segment 
XI not widened apically, about 3 times as* long as wide. Tip of scutellum exposed. 
Elytra with distinct parasutural striae, suturai striae starting at margin of pronotal lobe, 
slightly curved at base. Hypomeron not microsculptured, with longitudinal stria. 
Middle part of metaventrite flat, lacking stria or impression. Sides of metaventrite 
microsculptured, sparsely and finely punctate, with long pubescence. Mesocoxal lines 
with very fine marginal punctures not extending laterally along mesepimera; sub - 
mesocoxal areas about 0.02 mm long. Metepisterna flat, with exposed portion about 
0.04-0.05 mm wide, almost parallel-sided, with suture slightly curved apically. 
Abdomen with striate microsculpture, hardly visible on ventrite 1 in some spécimens. 
Abdominal ventrite 1 with basai punctures fairly coarse, partly elongate. Tibiae 
straight. 



SCAPHISOMATINI OF THE PHILIPPINES 



715 



Maie characters. Protarsi with segments 1 to 3 strongly widened, about as wide 
as protibiae. Middle part of metaventrite, its anterior fourth excepted, completely cov- 
ered by a very dense, flat patch of yellowish setae. Aedeagus (Figs 39, 40) 0.55-0.61 
mm long. 

Female characters. Metaventrite smooth in center, around center densely punc- 
tate, with short pubescence. 

Habitat: In rotten wood under a pine. 

Distribution: Philippines: Luzon. 

Comments: This species has the médian lobe of the aedeagus similar to that in 
S. davaoana, S. amicalis Lôbl, 2003, S. tenella Lôbl, 1990 and S. uncata Lôbl, 1990. 
See also comments under S. davaoana. It differs drastically from thèse species by the 
much wider flagellum. The pubescence covering completely the middle part of the 
maie metaventrite is diagnostic for this new species. 

Scaphobaeocera sabapensis Lôbl, 1990 

MATERIAL EXAMINED: 3 6,2 9, Luzon, Lagunas Prov., Mt. Makiling, 600m, summit 
rd, 21. XI. 1995, leg. I. Lôbl. - 1 $ , same data but 20.XI, 450-500m. -29, Luzon, Lagunas, Mt. 
Banahaw ca 1 km Kinabuhayan, 500m, 26.XI.1995, I. Lôbl. - 1 9, Luzon, Lagunas, Mt. 
Banahaw above Kinabuhayan, 600-700m, trail to Crystalino 24.XI.1995, J. Kodada & B. 
Rygovâ leg. (ail MHNG). 

Habitat: Evergreen rain forest, leaf litter. 

Distribution: Thailand, Philippines: Luzon. 

Comments: The aedeagi of the three Philippine maies have the ventral wall of 
the apical process of the médian lobe almost regularly arcuate and so slightly differ 
from those in the examined Thai spécimens. 

Scaphobaeocera serpentis sp. n. Figs 41-43 

HOLOTYPE: 6, Philippines: Mt. Makiling, Laguna Prov., 4 km SE Los Banos 9-IV-1977 
/ berlese leaf litter L. E. Watrous (MHNG). 

PARATYPES: 1 6 f 2 2 with the same data as holotype; 1 â, with the same data but 
1 1 -IV- 1977 / berlese litter under bark L. E. Watrous (ail MHNG). 

DESCRIPTION: Length 0.95-1.05 mm, width 0.58-0.61 mm, dorsoventral 
diameter 0.53-0.58 mm. Head and most of body uniformly light ochraceous, 
appendages hardly lighter than body. Pronotum and elytra lacking microsculpture. 
Punctation extremely fine on pronotum, elytra, hypomera and latéral parts of meta - 
ventrite. Length ratio of antennomeres as: III 4: IV 6: V 7: VI 5: VII 11: VIII 4: IX 12: 
X. 12: XI 16. Segments III and IV evenly narrow, segment IV about twice as long as 
wide. Segment V slightly wider than segment IV, about 3 times as long as wide. 
Segment VI about twice as long as wide, wider than segment V. Segment VII much 
larger than segment VI, not quite 3 times as long as wide. Segment VIII conspicuously 
small, slightly longer than wide. Segments IX to XI almost even in width, hardly wider 
than segment VII, segment XI about 3 times as long as wide. Tip of scutellum exposed. 
Elytra without parasutural striae, suturai striae starting at margin of pronotal lobe, 
hardly curved at base. Hypomeron not microsculptured, without longitudinal stria. 



716 



I. LÔBL 



Middle part of metaventrite slighly convex, without impression, densely punctate 
around smooth center, with fairly long pubescence. Mesocoxal lines with coarse 
marginal punctures, not extending laterally along mesepimera; submesocoxal areas 
about 0.03 mm long. Metepisterna flat, exposed portion about 0.08-0.10 mm wide, 
with suture notably deep, sulciform, weekly arcuate. Abdominal ventrite 1 very finely 
punctate, lacking obvious microsculpture, with basai punctures coarse, partly elongate. 
Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 strongly widened, about as wide 
as protibiae. Aedeagus (Figs 41-43) 0.25-0.27 mm long. 

Habitat: Evergreen rain forest, leaf litter. 

Distribution: Philippines: Luzon. 

COMMENTS: This species shares with S. epistemalis large metepisterna and 
mesocoxal lines with conspicuous, coarse marginal punctures. Both species may be 
distinguished by charaters given in the key above. In addition, the aedeagus is distinc- 
tive in S. serpentis. 

Scaphobaeocera watrousi sp. n. Figs 44-46 

HOLOTYPE: S, Philippines: Mt. Makiling, Laguna Prov., 4 km SE Los Banos 11 -IV- 
1977 / berlese débris under bark L. E. Watrous (MHNG). 

PARATYPES: 1 9 , with same data as holotype. - 1 S, Philippines: Mt. Makiling, Laguna 
Prov., 4 km SE Los Banos 8-IV-1977 / berlese débris rotten log L. E. Watrous and berlese rotten 
log. - 1 â, 4 9, with same data but berlese leaf litter (ail MHNG); 1 S, Tarragona, Leyte Is., 
Philippine Islds. VIII: 1 1 : 1945 coll. and près, by C. L. Remington (FMNH). 

Description: Length 1.0-1.10 mm, width 0.60-0.65 mm, dorsoventral diame- 
ter 0.63-0.67 mm. Head and most of body uniformly light ochreous, appendages lighter 
than body. Pronotum lacking microsculpture. Basai half of elytra not microsculptured, 
apical half of elytra microsculptured, hardly iridescent. Punctation extremely fine on 
pronotum, elytra, hypomera and latéral parts of metaventrite. Length ratio of anten- 
nomeres as: III 5: IV 6: V 8: VI 5: VII 9: VIII 6: IX 10: X 10 : XI 22. Segments III to 
IV equally narrow, segment IV about 2.5 to 3 times as long as wide. Segment V hard- 
ly wider than segment IV, about 4 times as long as wide. Segment VI about 2 to 3 times 
as long as wide, wider than segment V. Segment VII much wider than segment VI, 
about twice as long as wide. Segment VIII almost as wide as segment VII and not quite 
2 times as long as wide. Segments IX to XI almost equally wide, not or slightly wider 
than segment VII, segment XI about 4 to 5 times as long as wide. Tip of scutellum ex- 
posed. Elytra without parasutural striae, suturai striae starting at margin of pronotal 
lobe, hardly curved at base. Hypomeron not microsculptured, with longitudinal stria. 
Middle part of metaventrite slighly convex, without impression, smooth in center, very 
densely punctate laterally and posterior center, with fairly long pubescence. Mesocoxal 
lines with marginal punctures fine, distinct, not extending laterally along mesepimera; 
submesocoxal areas about 0.02 mm long. Metepisterna flat, exposed portion about 
0.07-0.09 mm wide, narrowed anteriad, with suture straight and deep. Abdomen with 
distinct striate microsculpture. Abdominal ventrite 1 very finely punctate, with basai 
punctures fairly coarse, elongate. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 strongly widened, about as wide 
as protibiae. Aedeagus (Figs 44-46) 0.32-0.34 mm long. 



SCAPHISOMATINI OF THE PHILIPPINES 



717 




Figs 39-46 

(39, 40) Scaphobaeocera pubiventris sp. n., aedeagus in dorsal and latéral views; scale bar = 
0.2 mm. (41-43) Scaphobaeocera serpentis sp. n., aedeagus in dorsal and latéral views, internai 
sac (42) enlarged. Scale bar = 0.1 mm for figs 41, 43; scale bar = 0.05 mm for fig. 42. (44-46) 
Scaphobaeocera watrousi sp. n., aedeagus in dorsal and latéral views, internai sac (45) enlarged. 
Scale bar = 0.1mm for figs 44, 46; scale bar = 0.05 mm for fig. 45. 



718 



[. LÔBL 



Habitat: Evergreen rain forest, rotted wood and leaf litter. 
Distribution: Philippines: Luzon and Leyte. 

COMMENTS: The aedeagus of this species is similar to that in S. palawana. This 
new species may be easily distinguished from the latter by the lighter coloration of the 
body and the présence of hypomeral striae. 

ETYMOLOGY: The species is named in honour of one of its collectors, Larry E. 
Watrous, Ballwin, MO, USA. 

Scaphobaeocera werneri sp. n. Figs 47-51 

HOLOTYPE: â, E. slope Mt. McKinley, Davao Prov., MINDANAO Elev. 3200ft. Lot 
#54. IX, 7-8. 1946; beating / CNHM Philippine Zool. Exped. (1946-47) F. G. Werner leg. 
(FMNH). 

PARATYPES: 3 6 and 2 9, with the same data as the holotype (FMNH, MHNG). 

Description: Length 1.0-1.10 mm, width 0.55-0.62 mm, dorsoventral diameter 
0.58-0.64 mm. Head and most of body uniformly very dark reddish-brown, apex of 
abdomen and appendages light, ochraceous. Pronotum lacking microsculpture, elytra 
microsculptured and distinctly iridescent. Punctation extremely fine on pronotum, 
hypomera, elytra and latéral parts of metaventrite. Length ratio of antennomeres (Fig. 
47) as: III 5: IV 6: V 8: VI 7: VII 10: VIII 7: IX 11: X 10: XI 15. Segment III com- 
paratively short, about 3 times as long as wide. Segments IV to VI narrow, as wide as 
segment III. Segment VII about 2.5 times as long as wide. Segment VIII wider than 
segment VI. Segments IX and X distinctly larger than segment VII. Segment XI not 
widened apically, about 3 times as long as wide. Tip of scutellum exposed. Elytra with 
distinct parasutural striae, suturai striae starting at margin of pronotal lobe, hardly 
curved at base. Hypomeron not microsculptured, lacking longitudinal stria. Middle 
part of metaventrite flat, with or without minute, shallow impression, very densely 
punctate, with short pubescence. Latéral parts of metaventrite sparsely punctate, with 
striate microsculpture hardly visible at lOOx magnification, and fairly long pubescence. 
Mesocoxal lines with fine marginal punctures fine, not extending laterally along 
mesepimera; submesocoxal areas about 0.01 mm long. Metepisterna flat, exposed 
portion about 0.01-0.02 mm wide, almost parallel-sided, with suture slightly curved. 
Abdomen with distinct striate microsculpture. Abdominal ventrite 1 with basai punc- 
tures fairly fine, partly elongate. Tibiae straight. 

Maie characters. Protarsi with segments 1 to 3 strongly widened, about as wide 
as protibiae. Aedeagus (Figs 48-51) 0.29-0.32 mm long. 

Habitat: Found by beating végétation. 

Distribution: Philippines: Mindanao. 

COMMENTS: The aedeagal characters suggest relationship with S. hamata, the 
shape of the base of the flagellum in thèse species is, however, clearly distinctive. This 
species may be distinguished from S. hamata also by its narrow metepisterna. 

ETYMOLOGY: The species is named in honour of its collector, Floyd G. Werner. 

Xotidium Lôbl, 1992 

The genus is distinctive by having two-segmented labial palpi. It comprises five 
species known at présent only from Mauritius, Sri Lanka, Himalaya, and Queensland. 



SCAPHISOMATINI OF THE PHILIPPINES 



719 




Figs 47-55 



(47-51) Scaphobaeocera werneri sp. n., contours of antennomeres VII to XI (47); aedeagus in 
dorsal and latéral views, internai sac (49, 5 1 ) enlarged. Scale bars = 0. 1 mm for figs 47, 48, 50; 
scale bar = 0.05 mm for figs 49, 51. (52-55) Xotidium tubulifenim sp. n., aedeagus in dorsal and 
latéral views (52, 55), apical, tubular part of internai sac (53), apical part of aedeagus with 
extruded basai portion of internai sac (54). Scale bars = 0.2 mm for figs 52, 55; scale bar 0. 1 mm 
for figs 53, 54. 



720 



I. LÔBL 



An additional species is présent within the examinée! material from the Philippines and 
described below. 

Xotidium tubuliferum sp. n. Figs 52-55 

HOLOTYPE: S, Tarragona, Leyte I. Philippine Islds VIII: II: 1945 / Col. & près, by C. L. 
Remington /in rotting burned log (FMNH). 

ADDITIONAL MATERIAL: 1 6 (lacking head and prothorax), with the same data as the 
holotype (MHNG). 

DESCRIPTION: Length 1.60 mm, width 0.98 mm, dorsoventral diameter 0.93 
mm. Body and head uniformly reddish-brown to piceous, femora, tibiae and basai 
abdominal segments hardly lighter, apical abdominal segments, antennae and tibiae 
distinctly lighter, almost yellowish. Length ratio of antennomeres as: III 10: IV 14: V 
16: VI 15: VII 18: VIII 18: IX 15: X 18: XI 22. Segments III to VIII very narrow, 
segments VII and VIII similar, only slightly wider than segment III to VI, segments IX 
to XI much wider. Pronotal and elytral punctation very fine, visible at 50x magnifi- 
cation. Latéral parts of metaventrite and abdomen impunctate. Elytra each with fine 
basai stria joined to suturai and latéral striae; adsutural areas flat. Center of meta- 
ventrite almost flat, with few distinct punctures. Mesocoxal areas 0.07 mm long, 
shortest interval to metacoxae about 0.10 mm. Metepisterna flat, parallel-sided, about 
0.08 mm wide. Abdomen impunctate, with punctulate microsculpture, metacoxal Unes 
impunctate. 

Maie. Segments 1-3 of protarsi slightly widened. Aedeagus (Figs 52-55) 
0.72 mm long. 

Habitat: Rotting burned log. 

Distribution: Philippines: Leyte. 

COMMENTS: This new species resembles X. uniforme Lôbl, 1992 by its compa - 
ratively large size and by the color of the body, in combination with the elytra having 
complète and joined basai and suturai striae. Xotidium mauritianum (Vinson, 1943) is 
almost as large and has also unicolored body, but differs drastically from its congeners 
by the shortened suturai striae and the absence of basai striae (see VINSON, 1943). 
Xotidium tubuliferum may be distinguished easily by its aedeagus lacking sclerotized 
flagellum, and by the long widened apical section of the parameres. 

Both available maies have completely extruded internai sac of the aedeagus. It 
consists of a simple tube narrowed in middle, longer than the entire médian lobe. Very 
fine, spine-like structures are présent in its basai part. 

ACKNOWLEDGEMENTS 

I am particularly indebted to Jan Kodada from Bratislava, Slovakia, Louis 
Deharveng, and Jean Orousset, both Paris, France, late Masataka Satô from Nagoya, 
Japan, and Larry E. Watrous from Ballwin, MO, USA, for the generous gift of material 
from their own collections. Additional material was made available for study thanks to 
late Henry S. Dybas and Alfred F. Newton Jr., both from Chicago, USA, and Wolfgang 
Schawaller, Stuttgart, Germany. 



SCAPHISOMATINI OF THE PHILIPPINES 



721 



REFERENCES 

LESCHEN, R. A. B. & LÔBL, I. 2005. Phylogeny and classification of Scaphisomatini (Staphy- 
linidae: Scaphidiinae) with notes on mycophagy, termitology and functional morpho - 
logy. Coleopterists Society Monographs Patricia Vaurie Séries 3: 1-63. 

LÔBL, [. 1969. Revision der palàarktischen Arten der Tribus Toxidiini (Col. Scaphidiidae). 
Mitteilungen der Schweizerischen entomologischen Gesellschaft 42: 344-350. 

LÔBL, I. 1972. Beitrag zur Kenntnis der Scaphidiidae (Coleoptera) von den Philippinen. 
Mitteilungen der Schweizerischen entomologischen Gesellschaft 45: 79-109. 

LÔBL. [. 1992. The Scaphidiidae (Coleoptera) of the Népal Himalaya. Revue suisse de Zoologie 
99: 471-627. 

LÔBL, I. 1997. Catalogue of the Scaphidiinae (Coleoptera: Staphylinidae). Instrumenta bio - 
diversitatis 1: i-xii + 1-190. 

VlNSON, J. 1943. The Scaphidiidae of Mauritius. The Mauritius Institute Bulletin 2 (3): 177-209. 



Revue suisse de Zoologie 118 (4): 723-758; décembre 201 1 



Taxonomic emendations in the genus Liocheles Sundevall, 1833 
(Scorpiones, Liochelidae) 

Lionel MONOD 

Muséum d'histoire naturelle, Département des arthropodes et d'entolomogie I, 
C. P. 6434, CH-1211 Genève, Switzerland. 
E-mail: lionel.monod@ville-ge.ch 

Taxonomic emendations in the genus Liocheles Sundevall, 1833 
(Scorpiones, Liochelidae). - Examination of an extensive material that 
includes the type séries and recently collected spécimens enable a précise 
reassessment of the status of several taxa belonging to the genus Liocheles 
Sundevall, 1833 (Scorpiones, Liochelidae). Two species previously in syno- 
nymy, i. e. Liocheles boholiensis (Kraepelin, 1914) and Liocheles neocale- 
donicus (Simon, 1877), are revalidated. Liocheles australasiae longimanus 
(Werner, 1939) is elevated to species rank and Liocheles australasiae brevi- 
digitatus Werner, 1936 is synonymized with Liocheles australasiae 
(Fabricius, 1775). L. boholiensis (Kraepelin, 1914), L. neocaledonicus 
(Simon, 1877) and L. longimanus (Werner, 1939) are thoroughly redes- 
cribed, diagnosed and illustrated, and their distribution ranges are accurately 
mapped. 

Keywords: boholiensis - longimanus - neocaledonicus - taxonomy - 
Australasia - Indonesia, Sumatra - New Caledonia - Philippines - Bohol. 

INTRODUCTION 

The genus Liocheles Sundevall, 1833 is currently divided into two species 
groups which are easily distinguishable by the trichobothrial pattern on the retrolateral 
side of the pedipalpal chela, i. e. the Liocheles waigiensis (Gervais, 1843) species 
group and the Liocheles australasiae (Fabricius, 1775) species group (Monod, 2000; 
Monod & Volschenk, 2004). Three species and two sub-species are recognised within 
the australasiae-gvoup and six species within the waigiensis-group. However, an 
ongoing global taxonomic revision of the genus reveals an unsuspected spécifie diver- 
sity. Preliminary results of this work were first presented in an unpublished master 
thesis (Monod, 2000) carried out at the University of Geneva (Switzerland). The 
présent contribution intends to formally présent part of this work. 

Over the years, several authors attempted to revise the genus but were unable to 
clearly assess the taxonomic status and position of most species, and several valid taxa 
are currently considered as junior synonyms owing to the paucity of diagnostic 
characters and to the scarcity of material available in muséum collections, especially 
mature maies that are essential for diagnoses of species. Examination of an extensive 
material, including types, and récent field surveys conducted by the author during 
which fresh spécimens were collected, allowed a précise reassessment and subséquent 



Manuscript accepted 30.09.2011 



724 



L. MONOD 



reinstatement of two species in synonymy, i.e. Liocheles boholiensis Kraepelin, 1914, 
and Liocheles neocaledonicus Simon, 1877. Besides, Liocheles australasiae longima- 
nus (Werner, 1939) is elevated to species rank, while Liocheles australasiae brevidigi- 
tatus Werner, 1936 is synonymized with Liocheles australasiae australasiae 
(Fabricius, 1775). Redescriptions, diagnoses, illustrations of important characters and 
distribution maps are provided for the three re-established taxa. 

Liocheles boholiensis described by Kraepelin in 1914, was placed into the 
synonymy of L. australasiae by L. E. Koch (1977). However, the species clearly 
belongs to the waigiensis group. Unlike in most species of the waigiensis group, the 
anterior area of the carapace is almost completely smooth and devoid of a coarse 
granulation, resembling the carapace of the australasiae group. This feature probably 
led Koch to erroneously considered the species as a junior synonym ofL. australasiae. 
Liocheles neocaledonicus was first described as Lschnurus neocaledonicus by Simon 
( 1877). Kraepelin ( 1 894) synonymized it with Hormurus caudicula L. Koch, 1 867, but 
subsequently (Kraepelin, 1914) restored it as a valid species in the genus Hormurus 
Thorell, 1837. Kopstein (1921) accepted Kraepelin's décision to split//. caudicula into 
several distinct species, including H. neocaledonicus, and Giltay (1931) cited 
Kraepelin (1914) as the référence for taxonomic identification of Hormurus species, 
but contrary to Kopstein he considered the différent species of the waigiensis group as 
subspecies of H. caudicula, listing H. caudicula neocaledonicus from central and sou- 
thern New Caledonia and describing Hormurus caudicula sarasini from the north of 
the island. Following Karsch (1880) who first synonymized Hormurus with Liocheles, 
Takashima (1945) recognized L. neocaledonicus and L. sarasini as subspecies of L. 
caudicula. L. E. Koch (1977) subsequently treated H. caudicula and most of its sub- 
species (including L. caudicula neocaledonicus and L. caudicula sarasini) as junior 
synonyms of Liocheles waigiensis. The présent work confirms that L. neocaledonicus 
is a valid species while L. sarasini is a junior synonym of the former. 

Two subspecies ofL. australasiae, i.e. L. australasiae brevidigitatus andl. aus- 
tralasiae longimanus were described by Werner in 1936 and 1939, respectively. Thèse 
taxa remained widely unknown to subséquent authors before Fet (2000) reported them 
in the Catalog of the scorpions of the world. Significant morphological différences 
from L. australasiae warrant the élévation of L. australasiae longimanus to species 
level. However, L. australasiae brevidigitatus cannot be separated from L. australasiae 
based on morphology and that subspecies is thus abolished. 

MATERIAL AND METHODS 

Examination: Spécimens were examined with a ZEISS Stemi SV8 stereo- 
microscope. Morphological examination is sometimes hampered by mud and soil 
aggregates adhering to the tégument of spécimens and thèse can obscure fine détails. 
Ultrasonic treatment (sonication) was used to render the spécimens free of obstructing 
particles (Nowak et al, 2008). The spécimens were placed for 15 to 90 seconds in a 
digital ultrasonic cleaner filled with soapy water. This step was repeated until a satis- 
tactory degree of cleanliness was achieved. For smaller and older spécimens more 
numerous but shorter rounds of sonication were carried out in order to avoid damage. 
Following sonication, the spécimens were rinsed thoroughly with distilled water. 



Ll OC HELES SCORPIONES, LIOCHELIDAE 



725 



DISSECTION: Mature maie spécimens were dissected using microsurgical 
scissors and forceps in order to study their hemispermatophores. Paraxial organ tissue 
was removed either manually with forceps or chemically with Proteinase K (Qiagen, 
Venlo, The Netherlands) diluted with water to 50%. Hemispermatophores were 
emerged in the solution and then placed in an oven at 45-50°C for 1 5 minutes to an 
hour depending on the size and sclerotization of the structure. Once the soft tissue of 
the paraxial organ was sufficiently digested, hemispermatophore were retrieved from 
the solution and thoroughly rinsed with water. 

TERMINOLOGY follows Vachon (1956, 1963) for cheliceral dentition. Stahnke 
(1970), Stockwell (1989), Sissom (1990) and Hjelle (1990) for pedipalp segmentation, 
Prendini (2000) for pedipalp carinae patterns, Vachon (1974) for trichobothrial 
patterns, Couzjin (1976) for leg segmentation, Prendini (2000) for metasomal carinae, 
and Lamoral (1979) and Monod & Volschenk (2004) for hemispermatophore morpho- 
logy. Measurements follow Stahnke (1970) and were recorded in mm using an occular 
micrometer or a digital caliper. 

Illustration: Drawings were produced using a caméra lucida mounted on the 
stereomicroscope. The sketches were subsequently drawn with ink pens and scanned 
for further processing and editing. Photographs and illustrations were subsequently 
edited in Photoshop CS5 (Adobe Systems, San José, CA, USA) (background removal 
and contrast adjustment) and plates were prepared with Illustrator CS5 (Adobe 
Systems, San José, CA, USA). Colour drawings were produced as digital média based 
on scientific illustrations, photographs of live spécimens served as référence to 
accurately represent the body colours of the animais. 

Mapping: Distribution maps were produced using ArcGIS version 9.3 
(Environmental Systems Research Institute, Redlands, CA, USA), by superimposing 
point locality records on SRTM 90 m (3 arc-second) digital élévation data (Jarvis et al, 
2008) and SRTM 1 km (30 arc-second) global bathymetry dataset (Becker et al, 2009). 
Geographical coordinates for records without GPS data were traced by référence to 
gazetteers, the GEOnet Names Server (http://earth-info.nga.mil/gns/html/index.html), 
or detailed route maps of various earlier expéditions. 

Material LIST: Following Chapman & Grafton (2008), GPS coordinates that 
may be used by potential collectors seeking to conduct mass harvesting in order to 
supply the arachnid pet trade were purposely removed from the list of material. Thèse 
data may be supplied to researchers upon request. 

ACRONYMS: Depositories of material examined are abbreviated as follows: 
AMNH, American Muséum of Natural History (New York, USA); BPBM, Bernice P. 
Bishop Muséum, Honolulu (Hawaii, USA); CAS, Californian Academy of Sciences 
(San Fransisco, USA); LM, Lionel Monod personal collection, to be deposited in the 
collection of the MHNG; MHNG, Muséum d'histoire naturelle (Genève, Switzerland); 
MM, Manchester Muséum, University of Manchester (Manchester, United Kingdom); 
MNHN, Muséum national d'Histoire naturelle (Paris, France); MRHN, Musée Royal 
d'Histoire Naturelle de Belgique (Bruxelles, Belgium); NHML, Natural History 
Muséum (London, United Kingdom); NMB, Naturhistorisches Musem (Basel, 



726 



L. MONOD 



Switzerland); NNHM, National Natuurhistorisch Muséum (Leiden, The Netherlands); 
QM, Queensland Muséum (Brisbane, Australia); USNM, United States National 
Muséum of Natural History, Smithsonian Institution (Washington, DC, USA); WAM, 
Western Australian Muséum (Perth, Australia); ZMAK, Zoologisches Forschungs- 
institut und Muséum Alexander Koenig (Bonn, Germany); ZMH, Zoologisches 
Staatsinstitut, Zoologisches Muséum Hamburg (Hamburg, Germany). Besides, the 
following abbreviations were used: DDEE, Direction du Dévelopement Economique et 
de l'Environnement, Province Nord, Koné, New Caledonia); DRN, Direction des 
Ressources Naturelles, Province Sud, Nouméa, New Caledonia; IAC, Institut Agro- 
nomique Néo-Calédonien, Pocqueureux, La Foa; MIS, misidentification. 

RESULTS 

Family Liochelidae Fet & Bechly, 2001 
Genus Liocheles Sundevall, 1833 

Liocheles australasiae (Fabricius, 1775) 

Liocheles australasiae brevidigitatus Werner, 1936 syn. n. 

Liocheles boholiensis (Kraepelin, 1914) Figs 1-2, Table 1 

Hormurus boholiensis Kraepelin, 1914: 333; Weidner, 1959: 101. 

Hormurus caudicula boholiensis: Giltay, 1931: 12, 13, 18, 19. 

Liocheles caudicula boholiensis: Takashima, 1945: 96. 

Liocheles australasiae: L. E. Koch, 1977: 161 (misidentification). 

Liocheles boholiensis: Monod, 2000: 72-76, pis 22-24, map 8. 

Distribution RANGE: This species is known only from Bohol Island (Central 
Visayas), Philippines. However, it is most likely présent on other nearby Philippine 
islands. 

MATERIAL EXAMINED: ZMH, without registration number; 1 9 syntype (here designated 
lectotype), 1 subadult ? syntype (here designated paralectotype); Philippines, Bohol; X.1863. 

DiAGNOSlS: L. boholiensis can be distinguished from L. waigiensis by the 
following characters: 

(1) prolateral process of pedipalp patella forming 2 distinct spiniform processes 
(bifid), thèse sometimes fused but not forming a large médian spine; (2) ventral inter- 
carinal surface of pedipalp chela granular along prolateral and retrolateral edges only, 
smooth medially; (3) retrolateral ventral intercarinal surface of pedipalp fémur smooth 
or nearly so; (4) ventral intercarinal surface of pedipalp fémur with granulation absent 
or vestigial and limited to the retrolateral edge; (5) ventrosubmedian carinae of meta- 
somal segment I with 1-3 pairs of well developed medial granules; ventrosubmedian 
carinae of metasomal segment II with 1 pair of well developed subposterior spiniform 
granules and 1-3 pairs of well developed medial spiniform granules. 

Description of adult female: Coloration: The poor state of préservation of 
the type spécimens did not allowed to accurately describe the coloration of the animais. 

Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma, pro- 
soma, legs and pedipalp finely punctated. 

Chelicerae (Fig. 1C): Medial and basai teeth of fixed finger fused into a bicusp. 
Movable finger: dorsal margin with four teeth (one subdistal and one basai); distal dor- 
sal tooth smaller than distal ventral tooth; ventral margin smooth. 



LIOCHELES SCORPIONES, LIOCHELIDAE 



727 




Fie. 1 

Liocheles boholiensis Kraepelin, 1914, female holotype (ZMH). (A) Carapace, dorsal aspect. (B) 
Left tarsus IV, ventral aspect. (C) Left chelicera, dorsal aspect. (D) Pectines and génital opercula. 
(E) Metasoma, latéral aspect. Scale Unes, 2 mm (A, E), 1 mm (C, D), 0.5 mm (B). 



728 



L. MONOD 



Table 1 

Meristic data for the types of Liocheles boholiensis Kraepelin. 1914. 



Lectotype. female Paralectotype. 

subadult female 



^ cil updLC 


LCIlglll 


8 8 
o.o 


f\ 7 

O. / 




^YlllCI 1U1 VvlUUJ 


N Q 

J .O 






ruMtrllUl W1UUI 


7.0 


7 £> 


AlClclSUIIlal bCUIIlClll I 


L-CllUlll 


J.A 


Z.Z 




VY 1 Ci L 1 1 


Z..KJ 


1 S 


\A c*tn cr^m 1 1 crm Pti t \ 


Lensth 


-+. J 






W 1U111 


1 (\ 


1 




Height 


1.6 


1.2 


Telson vesicle 


Width 


1.8 


1.3 




ncigiu 


1 Q 


i . j 


Pedipalp 


Fémur length 


8.4 


5.9 




Fémur width 


3.4 


2.3 




Patella length 


8.5 


6.2 




Patella width 


3.4 


2.5 




Chela length 


17.4 


12.0 




Chela width 


7.2 


4.6 




Chela height 


3.6 


2.2 




Chela movable 


8.8 


6.0 




finger length 






Total length 




53.0 


46.0 



Pedipalp: Fémur as long as or shorter than carapace. Chela virtually asetose. 

Chela finger s morphosculpture: Denticulate edge even distally, with a double 
row of primary denticles often fused at the base, with inner accessory denticles (IAD). 
without enlarged granules. Chela fingers linear. without lobes and notches. 

Trichobothriotaxy (Fig. 2): Chela manus without accessory trichobothria: Dr tri- 
chobothrium midpalm or slightly less than midpalm: four V trichobothria: V 3 and V 4 
separate: Esb trichobothrium more distal than Eb group. close to Est: Eb$ trichobo- 
thrium close to Ebj_< Est trichobothrium midpalm or nearly so. Chela fixed finger: db 
trichobothrium on dorsal surface: esb. eb. est. et equidistant (distance est-esb similar to 
distance esb-eb) or est closer to esb: eb trichobothrium at base of finger. behind point 
of articulation between fixed and movable fingers. in line with esb-et axis: esb tricho- 
bothrium in line with est-et axis, in proximal région of fixed finger: two i trichobothria. 
Patella: d 2 trichobothria distal to patellar spur; et séries with three trichobothria; est 
séries with one trichobothrium: em séries with two trichobothria: esb séries with two 
trichobothria: em-esb séries with four trichobothria in a single group or in two groups 
esbj and esb^em^y, eb séries with five trichobothria in two groups ebj and eb : _ 5 or 
ebj/eb 4 _ 5 and eb 2 _f, three v trichobothria. 

Cahnae: Chela manus: dorsal secondary carina obsolète: digital carina distinct, 
costate or granular. stronger than external secondary carina: external secondary carinae 
absent or obsolète: ventroexternal carina granular or crenulate: ventromedian and 
ventrointernal carinae absent or obsolète: internomedian carina présent, granular 
(at least sparsely). Patella: prolateral process présent (distinct), strongly developed. 
forming two distinct spiniform processes (bifid). thèse sometimes fused but not 



LIOCHELES SCORPIONES, LIOCHELIDAE 



729 




Fig. 2 

Liocheles boholiensis Kraepelin, 1914, female holotype (ZMH), pedipalp with trichobothrial 
pattern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) 
Trochanter, fémur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral 
aspect. Scale line, 2 mm. 



730 



L. MONOD 



forming a large médian spine; internodorsal and dorsomedian carinae distinct; dorso- 
external carina absent or obsolète; externomedian carinae granular or costate; 
ventroexternal carina présent and distinct, costate or granular. Fémur: dorsoexternal 
carina présent and distinct, at least as a ridge, usually more distinct in proximal half, 
equally developed than dorsointernal carina; dorsointernal carina présent and distinct; 
internomedian dorsal carina absent, without spines or granules (or with a single basai 
spine); internomedian ventral carina absent, without spines, or at most vestigial, with 
a single spine at proximal extremity; ventromedian carina absent or obsolète; ventro- 
internal carina présent, granular. 

Macrosculpture\ Chela fingers granular (at least sparsely in proximal half). 
Chela fixed finger: area around dbldsbldst trichobothria granular (at least in the most 
proximal part), dbldsbldst trichobothria in a single large smooth dépression. Chela: 
dorsal intercarinal surface entirely and densely granular, with medium-sized spiniform 
granules; retrolateral intercarinal surface granular; ventral intercarinal surface granular 
along prolateral and retrolateral edges only, smooth medially; prolateral intercarinal 
surface at least sparsely granular, granulation less distinct along prolateral ventral edge. 
Patella: dorsal and ventral intercarinal surfaces entirely granular, at least a reticulated 
network of granules présent; retrolateral intercarinal surface at least sparsely granular; 
proximal half of prolateral intercarinal surface at least sparsely granular, distal half 
usually less granular. Fémur: dorsal intercarinal surface densely granular except at 
distal end; dorsal and prolateral intercarinal surfaces with small to medium-sized 
spiniform granules; retrolateral dorsal intercarinal surface at least sparsely granular; 
retrolateral ventral intercarinal surface smooth or nearly so; ventral intercarinal surface 
smooth or with vestigial granulation limited to the retrolateral edge; prolateral inter - 
carinal surface at least sparsely granular. 

Carapace (Fig. 1A): Anterior margin with shallow médian notch. Anterior 
furcated suture/sulcus présent and distinct. Médian ocular tubercle situated anterome- 
dially, small, occupying about 1/9 of carapace width, at least slightly raised; super- 
ciliary carinae présent, without granules. Médian ocelli separated by at least half 
diameter of médian ocellus, at least twice the size of latéral ocelli. Three pairs of latéral 
ocelli; ocelli equal in size, equidistant, close together, almost touching each other. 
Margin behind latéral ocellus without spines. Carapace surfaces with minute spiniform 
granules, evenly and sparsely distributed; anteromedian surface with frontal lobes 
smooth, granular along médian longitudinal sulci and anterior furcated sulci; rest of 
carapace at least sparsely granular. 

Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without 
distinct prominence; posterior margins of pretergite smooth, without spines of 
granules; posttergite entirely smooth or nearly so. Posttergites III- VII with distinct 
reticulated network of ridges and dimples, surface uneven; I-VI with latéral transversal 
sulcus. 

Ventral morphology: Anterodistal tip of coxa III without swelling or bulge. 
Sternum (Fig. 1D) equilateral pentagonal (anterior width slightly greater than posterior 
width); length less than or equal to posterior width. Génital operculum (Fig. 1D) oval 
to semi-oval, as wide as long (or wider than long), approximately same width as 
sternum; sclerites partly fused, médian suture distinct; posterior notch présent, at least 



LIOCHELES SCORPIONES, LIOCHELIDAH 



731 



weakly developed. Pectines (Fig. 1D) short (distal edge not reaching distal edge of 
coxa of leg IV), with fulcrae and three marginal lamellae; six pectinal teeth, long, 
straight, covered with sensory papillae only in distal portion. Stigmata (spiracles) half- 
moon shaped (with a distinct curve), short (less than 1/3 of sternite width). Sternite VII 
without longitudinal carinae. 

Metasoma (Fig. 1E) not flattened laterally. Segments I-IV with dorsomedian 
furrow shallow, weak to absent or only visible on segments E-III; dorsosubmedian 
carinae absent or obsolète; dorsolateral, ventrolateral and paired ventrosubmedian 
carinae présent and distinct on at least some segments. Segment I: width less than or 
equal to height; médian latéral carina présent and distinct; dorsosubmedian carinae 
with posterior spiniform granules weak to absent (not noticeably larger than preceding 
granules); dorsomedian posterior spiniform granules weak to absent; ventral surface 
without posterior spiniform granules; ventrosubmedian carinae with 1-2 pairs of 
moderate to strong subposterior spiniform granules and 1-3 pairs of well developed 
medial granules. Segment II: dorsosubmedian carinae with posterior spiniform 
granules weak to absent (not noticeably larger than preceding granules); dorsomedian 
posterior spiniform granules weak to absent; ventrolateral carinae without posterior 
spiniform granules; ventrosubmedian carinae without posterior spiniform granules, 
with one pair of well developed subposterior spiniform granules and 1-3 pairs of well 
developed medial spiniform granules. Segment III: dorsosubmedian carinae with 
posterior spiniform granules weak to absent (not noticeably larger than preceding 
granules); ventrolateral carinae with ridges indistinct or weak and smooth; ventro- 
submedian carinae with ridges indistinct or weak and smooth (sometimes with a few 
reduced spiniform granules), without subposterior spiniform granules. Segment IV: 
dorsosubmedian carinae with posterior spiniform granules weak to absent (not noti- 
ceably larger than preceding granules); ventrolateral carinae with no ridges visible or 
with weak smooth ridges; ventrosubmedian carinae with no ridges visible or with weak 
smooth ridges (sometimes with few reduced spiniform granules in posterior half), 
without subposterior spiniform granules. Segment V: dorsal surface smooth, without 
smooth shiny dépression in posterior half between ventrolateral carinae; dorsolateral 
carinae absent or obsolète; ventrolateral carinae présent and distinct, smooth or nearly 
so (rarely one pair of vestigial granules posteriorly); ventromedian carina absent or 
obsolète, ridges absent or weak and smooth; anal arch crenulate, at least with few 
reduced teeth. Telson as long as or slightly longer than metasomal segment V; vesicle 
smooth, without granules, unmodified. 

Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae 
often weakly developed). Fémur IV with ventromedian carinae vestigial (only 
expressed distally) or indistinct (only scattered granules). Tibiae I-II: retrolateral 
margins without spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spini- 
form macrosetae; retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV 
(Fig. 1B): two ventrosubmedian rows of secondarily setiform macrosetae; ventro- 
median row of spinules absent or vestigial; basai spinules présent (at least one, usually 
in a short row); terminal ventromedian spinules absent; ungues shorter than telotarsus. 
Telotarsi I-IV: prolateral/retrolateral rows with 3/4, 3/4, 3-4/4 and 4/4 spiniform macro- 
setae respectively. 



732 



L. MONOD 



Liocheles longimanus (Werner, 1939) stat. n. Figs 3-9, Table 2 

Hormurus australasiae longimanus Werner, 1939: 362. 

Liocheles australasiae longimanus: Locket, 1997: 331; Fet, 2000: 397; Monod & Volschenk, 
2004: 686. 

Liocheles longimanus: Monod, 2000: 97-101, pis 36-40, map 12. 

Distribution range and habitat: Indonesia, north-western Sumatra (FIG. 3). 

In humid tropical forests. 

MATERIAL EXAMINED: ZMAK, alte Trockenpràparate LU; 1 S syntype (here designa- 
ted lectotype), 1 9 syntype (here designated paralectotype); Indonesia, Sumatra, Sumatera Utara 
Province, Montes Battak; H. Fruhstorfer. - MNHN, RS 3477; 2 9; Indonesia, Sumatra, 
Kenandam?; III. 1913; Buxton?. - MNHN-RS 3471; 3 â, 5 9, 4 juv.; Indonesia, Sumatra, 
Kenandam?; 1913; Buxton?. - MITNG, Sum-06/11; 1 cT; Indonesia, Sumatra, Sumatera Barat 
Province, Harau Canyon, N of Payakumbuh, 750 m, old secondary forest; 7. VI. 2006; P. 
Schwendinger. -NNHM, without registration number; 1 cT; Indonesia, Sumatra, Sumatera Barat 
Province, Lubuk Sikaping (Sumatra's Westkust), 450 m; 1926; E. Jacobson. - CAS, without 
registration number; 1 â, 1 9.1 subadult d; Indonesia, Sumatra, Sumatera Barat Province, 
Mangani, mine near Kota Tinggi, 700 m; 21.VII.1983; E. S. Ross. - MNHN, RS 7417; 19,1 
juv.; Indonesia. Sumatra. Sumatera Barat Province, Ngalau Kamang Cave, near Bukittinggi; 
20.VII.1979; J. Balazuc. 

DlAGNOSIS: L. longimanus can be distinguished from L. australasiae by the 
following characters: (1) pectinal tooth count slightly higher (L. longimanus: 7-10 in 
maies and 6-8 in females; L. australasiae: 6-1 maies and 5-6 in females); (2) sexual 
dimorphism pronounced, pedipalps more elongated in maies (fémur longer than cara- 
pace); (3) ventral surface of pedipalp chela manus: V3-V4 close together, in proximal 
part of manus; (4) prolateral process of pedipalp patella forming a single large spine; 
(5) patella: dorsal intercarinal surface smooth or nearly so, prolateral edge and proxi- 
mal end weakly granular in some spécimens; retrolateral ventral intercarinal surface 
smooth or nearly so; (6) fémur: dorsal intercarinal surface with granulation absent or 
limited to proximal and retrolateral edges; retrolateral intercarinal surface smooth or 
nearly so; ventral intercarinal surface with granulation absent or vestigial and limited 
to retrolateral edge; (7) carapace entirely smooth or nearly so; (8) metasoma: dorso- 
submedian carinae of segment II with posterior spiniform granules weak to absent (not 
noticeably larger than preceding granules); ventrosubmedian carinae of segment I with 
medial granules weak to absent. 

DESCRIPTION OF ADULT MALE (habitus see Figs 4, 5A, B): Coloration: As in 
figure 4. Pedipalps black to reddish brown. Chelicerae: dorsal surface light brown to 
yellow; fingers slightly infuscated in some spécimens. Carapace dark brown. Tergites 
dark brown, slightly lighter than carapace. Coxapophyses, sternum, génital operculum, 
pectines and sternites light brown to yellow. Metasoma dark brown, slightly lighter 
than carapace. Telson yellow. Legs light brown to yellow (paler than tergites). 

Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma. 
prosoma, legs and pedipalp finely punctated. 

Chelicerae: Medial and basai teeth of fixed finger fused into a bicusp. Movable 
finger: dorsal margin with four teeth (one subdistal and one basai); distal dorsal tooth 
smaller than distal ventral tooth; ventral margin smooth. 

Pedipalp: Fémur slightly longer than carapace, Chela virtually asetose. 



LIOCHELES SCORPIONES, LIOCHELIDAE 



733 




Fig. 3 

Localities of Liocheles longimanus (Wemer, 1939) on Sumatra. 



Chela fingers morphosculpture: Denticulate edge even distally, with double row 
of primary denticles often fused at base, with inner accessory denticles (IAD), without 
enlarged granules. Chela movable finger: suprabasal lobe well developed, gently 
rounded dorsally and lacking a sharp conical tooth, wider than high, not overlapping 
fixed finger; suprabasal lobe and corresponding notch contiguous or at most with a 
reduced gap; basai lobe absent or at most 2-3 small spiniform granules présent. Chela 
fixed finger: suprabasal notch distinct and deep; suprabasal lobe well developed and 
conical; basai lobe absent. 

Trichobothriotaxy (Fig. 6): Chela manus without accessory trichobothria; Dt tri- 
chobothrium midpalm or slightly less than midpalm; four V trichobothria; V 3 and V 4 
close together, in proximal part of manus; Esb trichobothrium basai, in same axis as Eb 
group; Eb 3 trichobothrium close to Ebj_ 2 ; Est trichobothrium midpalm or nearly so. 
Chela fixed finger, db trichobothrium on dorsal surface; esb, eb, est, et equidistant (dis- 
tance est-esb similar to distance esb-eb) or est closer to esb; eb trichobothrium at base 
of finger, behind point of articulation between fixed and movable fingers, above esb- 
et axis; esb trichobothrium below est-et axis, at base of finger, behind point of articu- 
lation between fixed and movable fingers; two i trichobothria. Patella: d? trichobothria 
distal to patellar spur; et séries with three trichobothria; est séries with one tricho - 
bothrium; em séries with two trichobothria; esb séries with two trichobothria; em-esb 
séries with four trichobothria in two groups esbj_ 2 and emj, 2 ; eb séries with five 
trichobothria in two groups ebj and eb 2 _ 5 or ebj/eb 4 _ 5 and eb 2 _ 3 ; three v trichobothria. 

Carinae : Chela manus: dorsal secondary carina obsolète; digital carina distinct, 
costate or granular, stronger than external secondary carina; external secondary carinae 



734 



L. MONOD 



absent or obsolète; ventroexternal carina costate; ventromedian and ventrointernal ca- 
rinae absent or obsolète; internomedian carina présent, granular (at least sparsely or 
faintly). Patella: prolateral process présent (distinct), strongly developed, forming a 
single large spine; internodorsal and dorsomedian carinae distinct; dorsoexternal carina 
absent or obsolète; externomedian carinae granular or costate; ventroexternal carina 
présent and distinct, costate or granular. Fémur: dorsoexternal carina présent and 
distinct, at least as a ridge, usually more distinct in proximal half, less strongly deve- 
loped than dorsointernal carina; dorsointernal carina présent and distinct; interno- 
median dorsal carina absent, without spines or granules (or with a single basai spine); 
internomedian ventral carina vestigial, with two large spines (one proximal and one 
midway); internomedian carinae oriented parallel to dorsointernal and ventrointernal 
carinae; ventromedian carina absent or obsolète; ventrointernal carina présent. 

Macrosculpture : Chela fingers granular (at least sparsely in proximal half). 
Chela fixed finger: area around dbldsbldst trichobothria smooth (trichobothria not in 
distinct dépressions). Chela: dorsal intercarinal surface with vestigial granulation (at 
most low granules présent) limited to prolateral and retrolateral edges, absent from 
médian part; retrolateral intercarinal surface granular; ventral intercarinal surface 
without granules; prolateral intercarinal surface at least sparsely granular, granulation 
less distinct along prolateral ventral edge. Patella: dorsal intercarinal surface smooth or 
nearly so, prolateral edge and proximal end weakly granular in some spécimens; ven- 
tral, retrolateral intercarinal surface smooth or nearly so; proximal half of prolateral 
intercarinal surface at least sparsely granular, distal half usually less granular. Fémur: 
dorsal intercarinal surface with granulation absent or limited to proximal and retro- 
lateral edges, when présent, granulation dense; dorsal and prolateral intercarinal 
surfaces with small to medium-sized spiniform granules; retrolateral intercarinal 
surface smooth or nearly so; ventral intercarinal surface with granulation absent or 
vestigial and limited to retrolateral edge; prolateral intercarinal surface at least sparsely 
granular. 

Carapace. Anterior margin with shallow médian notch. Anterior furcated 
suture/sulcus présent, distinct. Médian ocular tubercle situated anteromedially, small, 
occupying about 1/9 of carapace width, at least slightly raised; superciliary carinae 
présent, without granules. Médian ocelli separated by at least half diameter of médian 
ocellus, at least twice the size of latéral ocelli. Three pairs of latéral ocelli; ocelli equal 
in size, equidistant, close together., almost touching each other. Margin behind latéral 
ocellus without spines. Carapace entirely smooth or nearly so. 

Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without 
distinct prominence; posterior margin of pretergites smooth, without spines of 
granules. Posttergites I-VI smooth medially, latéral areas at least sparsely granular 
posteriorly; VII with anteromedian area smooth, latéral and posteromedian areas 
granular; granular areas of posttergites I-VII with minute spiniform granules, uni - 
formly and sparsely distributed. Posttergites III- VII without reticulated network of 
ridges and dimples, surface even; I-VI with latéral transversal sulcus présent. 

Ventral morphology: Anterodistal tip of coxa III without swelling or bulge. 
Sternum (Fig. 7A) subpentagonal (anterior width approximately equal to or slightly 
less than posterior width); length less than or equal to posterior width. Pectines 



LIOCHELES SCORPIONES, LIOCHELIDAE 



735 




Fig. 4 

Liocheles longimamis (Werner, 1939). Maie, dorsal aspect. Reconstruction based on scientific 
illustrations and photographs of live spécimens. Scale line, 5 mm. 

(Fig. 7A) moderately long (distal edge reaching, but not surpassing distal edge of coxa 
of leg IV), with fulcrae and three marginal lamellae; 7-10 pectinal teeth, long, straight, 
entirely covered by sensory papillae. Stigmata (spiracles) half-moon shaped (with a 
distinct curve), short (less than 1/3 of sternite width). Sternite VII without longitudinal 
carinae. 



736 



L. MONOD 



Metasoma (Fig. 8C): Segments I-V, length similar to or slightly longer than in 
female (Fig. 8D), not flattened laterally. Segments I-IV with médian furrow shallow, 
weak to absent or only visible on segments I-III; dorsosubmedian and dorsolateral 
carinae absent or obsolète; ventrolateral and paired ventrosubmedian carinae présent 
and distinct on at least some segments. Segment I: width less than or equal to height; 
médian latéral carina présent and distinct; dorsosubmedian carinae with posterior 
spiniform granules weak to absent (not noticeably larger than preceding granules); 
dorsomedian posterior spiniform granules weak to absent; ventral surface with 1-2 
pairs of well developed posterior spiniform granules; ventrosubmedian carinae with 
1-2 pairs of moderate to strong subposterior spiniform granules, medial granules weak 
to absent. Segment II: dorsosubmedian carinae with posterior spiniform granules weak 
to absent (not noticeably larger than preceding granules); dorsomedian posterior spini- 
form granules weak to absent; ventrolateral carinae with well developed posterior 
spiniform granules; ventrosubmedian carinae with 2-3 pairs of posterior, one pair of 
subposterior and 1-3 pairs of medial well developed spiniform granules. Segment III: 
dorsosubmedian carinae with posterior spiniform granules moderate to strong 
(distinctly larger than preceding granules); ventrolateral carinae with ridges indistinct 
or weak and smooth; ventrosubmedian carinae with ridges indistinct or weak and 
smooth (sometimes with a few reduced spiniform granules), without subposterior 
spiniform granules. Segment IV: dorsosubmedian carinae with posterior spiniform 
granules moderate to strong (distinctly larger than preceding granules); ventrolateral 
carinae with no ridges visible or with weak smooth ridges; ventrosubmedian carinae 
with no ridges visible or with weak smooth ridges (sometimes with few reduced spini- 
form granules in posterior half), without subposterior spiniform granules. Segment V: 
dorsal surface smooth, with smooth shiny dépression in posterior half between ventro- 
lateral carinae; dorsolateral carinae absent or obsolète; ventrolateral carinae présent 
and distinct, anterior half smooth or nearly so, posterior half with strong conical tooth; 
ventromedian carina expressed only in posterior half, ridges absent or weak and 
smooth; anal arch crenulate, at least with few reduced teeth. Telson as long as or sligh- 
tly longer than metasomal segment V; vesicle smooth, without granules, unmodified. 

Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae of- 
ten weakly developed). Fémur IV with ventromedian carinae vestigial (only expressed 
distally) or indistinct (only scattered granules). Tibiae I-II: retrolateral margins without 
spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spiniform macrosetae; 
retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV (Fig. 8B): two 
ventrosubmedian rows of secondarily setiform macrosetae; ventromedian row of 
spinules absent or vestigial; basai spinules absent; terminal ventromedian spinules ab- 
sent; ungues shorter than telotarsus. Telotarsi I-IV: prolateral/retrolateral rows with 4/4, 
4/4, 4/4 and 4/4 spiniform macrosetae respectively. 

Hemispermatophore (Fig. 9): Distal lamina at least slightly curved, approxima- 
tely same size as basai part or slightly longer; distal crest absent; single lamellar hook, 
basai (in basai 1/3 of distal lamella or below, at least less than mid-length); basai 
extrusion absent; transverse ridge présent and distinct, merging with anterior edge abo- 
ve base of lamellar hook, at approximately same level as base of lamellar hook. 
Lamella thin, folded only proximally and unfolded to flattened distal extremity (tip and 



LIOCHELES SCORPIONES, LIOCHELIDAE 



737 





Fig. 5 

Liocheles longimanus (Werner, 1939). (A) Maie (MNHN-RS 3471), dorsal aspect. (B) Same, 
ventral aspect. (C) Female (MNHN-RS 3477), dorsal aspect. (D) Same, ventral aspect. Scale 
Unes, 5 mm. 



738 



L. MONOD 




FlG. 6 

Liocheles longimanus (Werner, 1939), maie (MNHN-RS 3471), pedipalp with trichobothrial pat- 
tern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) 
Trochanter, fémur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral 
aspect. Scale line, 2 mm. 



LIOCHELES SCORPIONES, LIOCHELIDAE 739 




Fig. 7 

Liocheles longimanus (Werner, 1939), pectines and génital opercula, ventral aspect. (A) Maie 
(MNHN-RS 3471). (B) Female (MNHN-RS 3471). Scale line, 1 mm. 



740 



L. MONOD 




FlG. 8 

Liocheles longimanus (Werner, 1939); maie (MNHN-RS 3471) (A-C), female (MNHN-RS 
3471) (D). (A) Carapace, dorsal aspect. (B) Right tarsus IV, ventral aspect. (C) Metasoma, latéral 
aspect. (D) Posterior part of metasoma, latéral aspect. Scale lines, 2 mm (A, C, D), 0.5 mm (B). 



UOCHELES SCORPIONES, LIOCHELIDAE 



741 




Fig. 9 



Liocheles longimanus (Werner, 1939), maie (CAS), left hemispermatophore. (A) In toto, dorsal 
aspect, Lh (lamellar hook), Tr (transverse ridge). (B) Détail of the capsular région, entai aspect, 
Ld (distal lobe), Ld (distal lobe). (C) Idem, ventral aspect, La (lamella), Lb (basai lobe). Scale 
Unes, 1 mm. 

base approximately of same width); longitudinal 'spine', accessory hook and accessory 
lobe absent; lamellar tip at approximately same level as base of lamellar hook, above 
tip of distal lobe. Distal lobe well developed as a distinct hump, without distinctive 
hook-like shape, without accessory hook, carinae or crest. Basai lobe well developed 
('spoon' shaped), merging with accessory anterior basai lobe; distal edge without 
accessory fold toward ectal part, forming a 90° angle with lamella; basai edge without 
accessory fold (no groove), forming a 90° angle with lamella. 

Description of adult female (habitus see Fig. 5C, D): Same characters as in 
maie except as follows. 

Pedipalp markedly shorter and bulkier than in maie (see Fig. 5). Denticulate 
edge of chela fingers linear or nearly so (without pronounced lobe and notch). 

Mesosoma: Posttergites I-VII entirely smooth or nearly so. 

Ventral Morphology: Génital operculum (Fig. 7B) oval to semi-oval, as wide as 
long (or wider than long), approximately same width as sternum; sclerites partly 



742 



L. MONOD 



Table 2 



Meristic data for adult maies and females of Liocheles longimanus (Werner, 1939). 







maies 






females 






Carapace 


Length 


6.8 


6.4 


6.5 


8.2 


7.2 


6.9 




Anterior width 


4.6 


4.3 


4.5 


5.2 


4.7 


5.0 




Posterior width 


7.5 


7.2 


7.2 


9.0 


7.5 


7.5 


Metasomal 


Length 


2.7 


2.6 


2.7 


2.8 


2.8 


2.7 


segment I 


















Width 


1.7 


1.6 


1.6 


2.1 


1.8 


1.7 


Metasomal 


Length 


4.2 


4.2 


4.1 


4.3 


3.9 


4.0 


segment V 


















Width 


1.2 


1.2 


1.3 


1.6 


1.3 


1.3 




Height 


1.5 


1.5 


1.5 


1.7 


1.5 


1.4 


Telson vesicle 


Width 


1.4 


1.3 


1.5 


1.6 


1.4 


1.4 




Height 


1.4 


1.4 


1.6 


1.8 


1.4 


1.4 


Pedipalps 


Fémur length 


8.3 


7.4 


8.0 


7.8 


6.8 


6.6 




Fémur width 


2.6 


2.9 


2.6 


3.1 


2.6 


2.8 




Patella length 


8.0 


7.6 


7.6 


8.0 


6.8 


6.9 




Patella width 


3.0 


3.1 


3.1 


3.6 


3.1 


3.2 




Chela length 


15.1 


13.6 


15.0 


15.6 


13.6 


13.8 




Chela width 


4.5 


4.6 


4.2 


5.6 


4.9 


4.8 




Chela height 


2.8 


2.7 


2.8 


3.9 


2.6 


2.6 




Chela movable 


6.6 


5.4 


6.5 


7.1 


6.0 


6.4 




finger length 














Total length 




46 


42 


42 


56 


47 


46 



fusedmedian suture distinct; posterior notch présent, at least weakly developed. Six to 
eight pectinal teeth (Fig. 7B), covered with sensory papillae only in distal portion. 

INTRA-SPECIFIC VARIABILITY: As in other scorpions, the number of pectinal 
teeth varies: 7-10 in maies and 6-8 in females. 

Liocheles neocaledonicus (Simon, 1877) Figs 10-16, Table 3 

hchnurus neocaledonicus Simon, 1877: 237, 238. 

Liocheles neocaledonico: Simon, 1887: 113 (misspelling of species name). 

Hormurus caudicula: Kraepelin, 1894: 135 (part)(misidentification); Kraepelin, 1901: 272 

(part)(misidentification). 
Hormurus neocaledonicus: Kraepelin, 1914: 330, 332, 334, 335; Lampe, 1917: 201; Weidner, 

1959: 101. 

Hormurus sarasini Kraepelin, 1914: 332, 335, 336 (considered a subspecies of Hormurus 

caudicula by Giltay, 1931: 13); Weidner, 1959: 101; Forçait, 1961: 49 syn. n. 
Hormurus caudicula neocaledonicus: Giltay, 1931: 13, 18, 19. 
Hormurus caudicula sarasini: Giltay, 1931: 13, 18, 19. 
Liocheles caudicula neocaledonicus: Takashima, 1945: 96. 
Liocheles caudicula sarasini: Takashima, 1945: 96. 

Liocheles waigiensis: L. E. Koch, 1977: 166-172, figs 18, 47, 82, 83, maps 9a, b (part)(mis- 

identification); Kamenz & Prendini, 2008: 11, 43, pl. 134 (misidentification). 
Liocheles neocaledonicus: Monod, 2000: 102-107, pis 41-45, map 13; Monod, 2005: 5. 



LIOCHELES SCORPIONES, LIOCHELIDAE 



743 




FIG. 10 

Localities of Liocheles neocaledonicus (Simon, 1877) on New Caledonia. 



Distribution range and habitat: Endémie to New Caledonia, widely distri- 
buted across the island (Fig. 10). In rocky habitats (scree slopes, boulders) of tropical 
humid forests. 

TYPES: The maie holotype of Ischnurus neocaledonicus Simon, 1877 could not 

be found in the collections of the MNHN, it is considered lost. - ZMH, without regis- 

tration number; 9 paratype of Ischnurus neocaledonicus (examined); New Caledonia; 

20.VIII.1900; Bougier. - NMB, 56-a; â lectotype of Hormurus sarasini Kraepelin, 

1914 (designated by Forcart, 1961: 49) (not examined); New Caledonia, Tchalabel; 

V.1911; F. Sarasin & J. Roux. - NMB, 56-a; 6 6 , 27 9 paralectotypes of Hormurus 

sarasini (not examined); same data as for lectotype. - ZMH, without registration 

number; 1 S, 5 9 and 1 juv. paralectotypes of Hormurus sarasini (examined), 

Tchalabel; 11.1913; F. Sarasin. 

OTHER MATERIAL: MNHN, VA 2660; 1 6 , 1 9 ; without locality data; spécimens exa- 
mined by Vachon in 1981. -AMNH,Acc 37523; 1 6,3 9, 9 juv.; New Caledonia; III-IV. 1939; 
L. Macmillan. - BPBM, without registration number; 1 juv.; J. & M. Sedlacek. - MNHN, 
without registration number; 1 9 ; P. Rougeot. - MNHN, without registration number; 2 9 ; 
14.1.1995, N. & B. Thibaud. - MNHN, RS 0474; 9 <?, 11 9, 21 juv.; collection E. Simon. - 
MNHN, RS 0488; 1 juv.; Germain. - MNHN, RS 0530; 2 9 ; M. Bougier; collection Simon 
no.1998. - MNHN, RS 4424; 19,1 juv.; M. Plessis. - MNHN, RS 8253; 1 9. - MRHN, 
without registration number; 1 9. - NHML, 1924.11. 1.1; 1 9; 1914; P. D. Montague. - QM, S 
17097; 1 6; 17.11.1977; B. Jamieson. - ZMH, without registration number; 19.- MNHN, RS 
8537; 1 juv.; Bogui-Col des Rousettes, Station 26b, 270 m, mesic forest; 4.V.1987; A. Mordan 
& S. Tillier. - MNHN, RS 3240; 3 6, 1 9,1 juv.; Bourail; 1902; H. Méray. - AMNH, without 
registration number; 1 juv.; Col des Rousettes, 490 m, dry forest; 29.V.1987; N. I. Platnick, R. J. 
Raven. - MNHN, RS 8537; 2 9, 2 juv.; Col des Rousettes, 500 m, mesic forest; 29.V.1987; S. 



744 



L. MONOD 



Tillier. - WAM, 98/1892; 1 9; Col des Rousettes, rainforest; 11.11.1993; M. S. Harvey, N. I. 
Platnick, R. J. Raven. - WAM 98/1891; 1 juv.; same data as WAM 98/1892. - LM, without 
registration number; 2 6 , 3 9, 7 juv.; Col des Rousettes, track, 427 m, small patch of rainforest 
in between niaoulis savannah, in between stones in a scree covered by litter and tree roots, dryer 
habitat than previous spécimens; 21. IX. 2004; D. Gaillard, L. Monod. - AMNH, LP 5544; 1 juv.; 
Col des Rousettes, track, 427 m, small patch of rainforest in between niaoulis savannah, in 
between stones in a scree covered by litter and tree roots, dryer habitat than previous spécimens; 
2 1 .IX.2004; D. Gaillard, L. Monod. - MNHN, RS 8537; 2 9 , 1 3 juv.; Dothio, Col de Pétchékara, 
Station 287, 340 m, mesic forest; 27.X.1986; A. & S. Tillier. - LM, without registration number; 
3 6,4 9; Farino, Louis Barbou forest exploitation, 383 m, degraded humid forest, near dry 
creek, under stones; 7. X. 2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. Pocqueureux), L. 
Monod. - AMNH, LP 5547; 1 juv.; Farino, Louis Barbou forest exploitation, 383 m, degraded 
humid forest, near dry creek, under stones; 7. X. 2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. 
Pocqueureux), L. Monod. - LM, without registration number; 2 6 , 3 9; Farino, Louis Barbou 
forest exploitation, 477 m, degraded humid forest, under stones; 7.X.2004; C. Mille, S. Cazeres 
& T. Nuques (I.A.C. Pocqueureux), L. Monod. - AMNH, LP 5548; 1 juv.; Farino, Louis Barbou 
forest exploitation, 477 m, degraded humid forest, under stones; 7. X. 2004; C. Mille, S. Cazeres 
& T. Nuques (I.A.C. Pocqueureux), L. Monod. - QM, S 23332; 1 juv.; Forêt nord, sud de 
Noumea, Kwa Neie transmitter station, mixed-dry forest; 25.X.1988; T. C. (?) & R. Raven. - 
MNHN, RS 8349; 1 juv.; Goro, mesic forest; 3.IX.1975; P. Bouchet. - MNHN, RS 8537; 1 6, 

1 juv.; S of Grand Lac, Station 235, 280 m, mesic forest on peridotites; 26. XI. 84; A. & S. Tillier, 
P. Bouchet (MNHNP, Malacologie). - BPBM, Acc. No. 1979.380; 2 juv.; Hienghene, 10-50m; 
14-17.VIII.1979; G. M. Nishida. - AMNH, AH 4622-4643; 3 6, 17 9, 2 juv.; Koyaboa Forest 
(Poindimié), NE central coast, 16/22.X.1983; Saint Louis (Noumea), on the banks of a creek; 

16. XI.1979; A. Renévier. - WAM, 98/1887-8; 1 1 juv.; Ile des Pins, near Grotte de la 3ème, 
rainforest; 19.111.1993; M. S. Harvey, N. I. Platnick, R. J. Raven. - MNHN, RS 7333; 2 9; La 
Foa; IX. 1976; M. Droin. - MNHN, without registration number; 19; Grotte Le Cresson, 
18.IX.65.-QM, S 39694; 2 9, 1 juv.; Chute de la Madeleine, 230 m; 12.XI.2000; P. Bouchard, 
C. Burwell & G. Monteith (9918). - NHML, 1 927.11. 1 .2-5; 4 9; Mt Arago; P. D. Montague. - 
USNM, 00753; 1 9, 1 juv.; Mt Dor, 15 mi from Noumea; 22.X.1944; W. D. Crabb. - LM, 
without registration number; 3 6 , 2 9 ; Mt Humbold, near mine, 505 m, degraded primary forest, 
vine thicket, in between stones in a scree covered by leaf litter and humus and under larger 
stones; 4.X.2004; L. Monod. - AMNH, LP 5546; 1 juv.; Mt Humbold, near mine, 505 m, 
degraded primary forest, vine thicket, in between stones in a scree covered by leaf litter and 
humus and under larger stones; 4.X.2004; L. Monod. - QM, without registration number; 1 9 ; 
Mt Koghis, 500 m, rainforest, night collecting; 2-3.XI.2002; Burwell, G. Monteith & S. Wright 
(1 1088). - LM, without registration number; 3 (5,3 9 ; Mt Koghis, start of tracks, 440 m, rain- 
forest, under rocks; 26.IX.2004; D. Gaillard, L. Monod. - AMNH, LP 5545; 1 juv.; Mt Koghis, 
start of tracks, 440 m, rainforest, under rocks; 26.IX.2004; D. Gaillard, L. Monod. - AMNH, LP 
6224; 2 juv.; Mt Koghis, ca. 20 km N Noumea; 4.XII.2004; S. Huber. - CAS, without regis- 
tration number; 1 9, 2 juv.; S. E. slope of Mt Koyaboa (390 m), Poindimié, élévation less than 
500 ft, 3.VI.1985, L. Wishmeyer. - MNHN, RS 8537; 1 9 , 20 juv.; Mt Mé Ori, SE slope, Station 
29 a, 530 m, forêt humide; 7.V.1987; A. Mordan & S. Tillier. - MNHN, without registration 
number; 1 6,2 9,1 juv.; Mt Pouédihi forest; 18.IV. 1965. - MNHN, RS 8537; 1 juv.; Barrage 
de la Néaoua (Ouen Sieu), Station 210, 500 m, mesic forest; 20.XI.1984; A. & S. Tillier, P. 
Bouchet, M.-P. Triclot (MNHNP, Malacologie). - AMNH, LP 5542; 1 9 ; Road Nouméa-Rivière 
Bleue Forest Reserve, 177 m, vine thicket, in rock crevices; 17. IX. 2004; D. Gaillard & L. 
Monod. - MNHN, RS 8350; 5 juv.; Oua Tom, 130 m, mesic forest; 16.IX.1978. - MNHN, RS 
8537; 1 9, 2 juv.; Vallée de la Ouen Nondoué, Station 205, 70 m, Nothofagus forest; 20.X.1984; 
A. & S. Tillier, P. Bouchet, M.-P. Triclot (MNHNP, Malacologie). - NHM, 1924.11. 1.6-7; \ 6,1 
9 ; Plaine des Lacs, forest, under bark; 23.11.1914; D. Montague. -MNHN, RS 8537; 1 6, 1 9, 

2 juv.; Rivière Blanche, Parc Cagous, Station 256, 160/170 m, mesic forest; 1. IX. 1986; A. & S. 
Tillier. - MNHN, without registration number; 1 6,1 9, 6 juv.; Rivière Bleue Forest Reserve; 

17. VIII.1965. - MNHN, without registration number; 3 juv.; Rivière Bleue Forest Reserve; 

20. VIII.1965. - MNHN, without registration number; 2 juv.; Rivière Bleue Forest Reserve; 

21. XI. 1995; J. P. Hugot. - MNFTN, without registration number; 1 6; Rivière Bleue Forest 
Reserve, under bark; 17. VII. 1965; Stanniloner. - AMNH, without registration number; 1 juv.; 



LIOC HELES SCORPIONES, LIOCHELIDAE 



745 




FIG. 1 1 

Liocheles neocaledonicus (Simon, 1877). Maie, dorsal aspect. Reconstruction based on scien - 
tific illustrations and photographs of live spécimens. Scale line, 5 mm. 

Rivière Bleue Forest Reserve, 280 m, wet forest; 21.V.1987; N. I. Platnick, R. J. Raven. - QM, 
S 23328; 3 $, 3 juv.; Rivière Bleue Forest Reserve, N rainforest; 21.V.1987; R. Raven. - WAM, 
98/1885-6; 1 6, 1 9 ; Rivière Bleue Forest Reserve, 240 m, rainforest; 9.III. 1993; M. S. Harvey, 
N. I. Platnick, R. J. Raven. - LM, without registration number; 3 â , 3 9; Rivière Bleue Forest 
Reserve, trail between Pont Péri gnon and Kaori géant, 116-169 m, rainforest, under stones and 
in rock crevices; 18.IX.2004; D. Gaillard, L. Monod. - AMNH, LP 5543; 1 juv.; Rivière Bleue 



746 



L. MONOD 



Forest Reserve, trail between Pont Pérignon and Kaori géant, 116-169 m, rainforest, under 
stones and in rock crevices; 18.IX.2004; D. Gaillard, L. Monod. - AMNH, LP 4323; 2 juv.; 
Rivière Bleue Forest Reserve, Pic du Grand Kaori, 400 m, under dead wood branches; 
1 1.V.2005; J. Murienne. - MNHN, RS 8537; 1 9, 1 juv.; Rivière Bleue Forest Reserve, mesic 
forest/alluvia, station 250, parcelle VI M, 160 m; 1. VIII. 1986; A. & S. Tillier. - MNHN, RS 
8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 a, parcelle VI H, 160 
m; 15.IX.1986; Y. Letocart, A. & S. Tillier. - MNHN, RS 8537; 2 juv.; Rivière Bleue Forest 
Reserve, mesic forest/alluvia, station 250 b, parcelle VI I, 160 m; 2.X.1987; A. & S. Tillier. - 
MNHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 d, par- 
celle VI I, 160 m; 4.XII.1987; A. & S. Tillier. - MNHN, RS 8537; 1 juv.; Rivière Bleue Forest 
Reserve, mesic forest/alluvia, station 250 e, parcelle VI J, 160 m; 5.1.1987; A. & S. Tillier. - 
MNHN, RS 8537; 1 â, 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 g, 
parcelle VI O, 160 m; 6.III.87; A. & S. Tillier. - MNHN, RS 8537; 1 9; Rivière Bleue Forest 
Reserve, mesic forest/alluvia, station 250 h, parcelle VI O, 160 m; 6.IV.1987; A. & S. Tillier. - 
MNHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 i, 
parcelle VI W, 160 m; 30.V.1987; A. Mordan, A. & S. Tillier. - MNHN, RS 8537; 1 S ; Rivière 
Bleue Forest Reserve, mesic forest/alluvia, station 250 j, parcelle VI G, 160 m; 12. VI. 87; A. & 
S. Tillier. - MNHN, RS 8537; 2 cT, 2 juv.; Rivière Bleue Forest Reserve, mesic forest, station 
251, parcelle VII V, 170 m; 15.VIII.1986; A. & S. Tillier. - MNHN, RS 8537; 2 juv.; Rivière 
Bleue Forest Reserve, mesic forest, station 251 a, parcelle VII U, 165 m; 22.IX.1987; A. & S. 
Tillier. - MNHN, RS 8537; 1 (5,1 9,1 juv.; Rivière Bleue Forest Reserve, mesic forest, station 
251 b, parcelle VII O, 170 m; 13.X.1986; A. & S. Tillier. - MNHN, RS 8537; 1 juv.; Rivière 
Bleue Forest Reserve, mesic forest, station 251 d, parcelle VII U, 170 m; 11. XII. 1986; A. & S. 
Tillier. - MNHN, RS 8537; 4 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 e, 
parcelle VII U, 170 m; 13.1.1987; A. & S. Tillier. - MNHN, RS 8537; 1 6,2 9 ; Rivière Bleue 
Forest Reserve, mesic forest, station 251 f, parcelle VII L, 170 m; 12.11.1987; A. & S. Tillier. - 
MNHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 g, parcelle VII 
K, 170 m; 16.111.1987; A. & S. Tillier. - MNHN, RS 8537; 1 â; Rivière Bleue Forest Reserve, 
mesic forest, station 251 h, parcelle VII R, 170 m; 14.IV.1987; A. & S. Tillier. - MNHN, RS 
8537; 1 d,l 9, 2 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 i, parcelle VII G, 
170 m; 14.V.1987; A. Mordan, A. & S. Tillier. - MNHN, RS 8537; 1 S, 4 juv.; Rivière Bleue 
Forest Reserve, mesic forest, station 251 k, parcelle VII, 170 m; 16.VII.1986; A. & S. Tillier. - 
MNHN, RS 8537; 2 juv.; Rivière Bleue Forest Reserve, Station 255, Mt Kaala, S slope, dry - 
forest; 27.VIII.1986; A. Chazeau & S. Tillier. - QM, S 39696; 1 d, 2 juv.; Rivière Bleue Forest, 
Kaori géant; 11-12.XI.2000; P. Bouchard, C. Burwell & G. Monteith (9957). - QM, S 23331; 
2 juv.; Rivière Bleue Reserve, near Kaori géant, 120 m; 25.V.1984; G. Monteith & D. Cook. - 
QM, S 39711; 1 9, 1 juv.; Rivière Bleue Forest Reserve, Panoramic track; 12.XI.2000; P. 
Bouchard, C. Burwell & G. Monteith (9951). - QM, without registration number; 1 6; Rivière 
Bleue Forest Reserve, Pic du Grand Kaori, 250 m; 16-18.XI.2002; S. Wright (11198). - QM, 
S 59098; 1 juv.; Rivière Bleue Forest Reserve, Pic du Grand Kaori, 250 m; 29.1.2002; G. B. 
Monteith (8922). - QM, S 23339; 19,3 juv.; Rivière des Pirogues (headwaters), 350-400 m; 
22.V.1984; G. Monteith & D. Cook. - MNHN, without registration number; 1 S, 1 juv.; Thi 
Forest Reserve; 5.VIII.65. - QM, S 23330; 1 9; Thi Forest Reserve, 150 m; 21.V.1984; 
G. Monteith & D. Cook. - AMNH, without registration number; 2 juv.; Touaourou, under stones; 
20.X.1959; B. Malkin. - USNM, 00753; 1 9 ; Touaourou; 25.XII.1960; J. P. E. Morrison (3927, 
gto VI). - WAM, 98/1889-90; 19,1 juv.; 2 km N of Troulala; 14.111.1993; M. S. Harvey, N. I. 
Platnick, R. J. Raven. 

DlAGNOSlS: L. neocaledonicus can be distinguished form L. waigiensis by the 
following characters: (1) fémur as long as or shorter than carapace length; (2) maie 
pedipalp chela: movable finger with basai lobe présent as a low hump with 2-3 larger 
conical teeth; fixed finger with suprabasal lobe low/reduced; (3) prolateral process of 
pedipalp patella low, forming two distinct spiniform processes (bifid), thèse sometimes 
fused but not forming a large médian spine; (4) prosoma with coarse spiniform 
granules at least on anteromedian surface, rest of prosoma usually with sparse smaller 
spiniform granules; (5) metasoma: dorsosubmedian carinae of segments III-IV with 



I /()( 7//.7 /..S S( OKPIOMS. [ KK HI I.IDAi: 



747 





B 





FIG. 12 

Liocheles neocaledonicus (Simon, 1877). (A) Maie (MNHN-RS 8537, Rivière Bleue Forest 
Reserve), dorsal aspect. (B) Same, ventral aspect. (C) Female (MNHN-RS 8537, Rivière Bleue 
Forest Reserve), dorsal aspect. (D) Same, ventral aspect. Scale lines, 5 mm. 



moderate to strong posterior spiniform granules (distinctly larger than preceding 
granules); ventrosubmedian carinae of segment I with 1-2 pairs of strong subposterior 
spiniform granules and 1-3 pairs of well developed medial granules; ventrosubmedian 
carinae of segment II with one pair of subposterior and 1-3 pairs of medial well deve- 



748 



L. MONOD 



loped spiniform granules; ventrosubmedian carinae of segment III with one pair of 
well developed subposterior spiniform granules. 

Description of adult male (habitus see Figs 11, 12A, B): Coloration'. As in 
figure 1 1 . Pedipalps black to reddish brown. Chelicerae: dorsal surface light brown to 
orange, fingers slightly infuscated. Carapace black to dark brown. Tergites black to 
dark brown, slightly lighter than carapace. Coxapophyses, sternum, génital operculum, 
pectines and sternites light brown to orange. Metasoma black to dark brown. Telson 
light brown to orange (paler than metasoma). Legs light brown to orange (paler than 
tergites). 

Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma, pro- 
soma, legs and pedipalp finely punctated. 

Chelicerae: Medial and basai teeth of fixed finger fused into a bicusp. Movable 
finger: dorsal margin with four teeth (one subdistal and one basai); distal dorsal tooth 
smaller than distal ventral tooth; ventral margin smooth. 

Pedipalp: Fémur as long as or shorter than carapace. Chela virtually asetose. 

Chela fingers morphosculpture: Denticulate edge even distally, with double row 
of primary denticles often fused at base, with inner accessory denticles (IAD), without 
enlarged granules. Chela movable finger: suprabasal lobe well developed, gently 
rounded dorsally and lacking a sharp conical tooth, wider than high, not overlapping 
fixed finger; suprabasal lobe and corresponding notch contiguous or at most with a 
reduced gap; basai lobe reduced to low hump, with 2-3 larger conical teeth. Chela fixed 
finger: suprabasal notch distinct and deep; suprabasal lobe présent and distinct, low 
and reduced; basai lobe absent. 

Trichobothriotaxy (Fig. 13): Chela manus without accessory trichobothria; Dt 
trichobothrium midpalm or slightly less than midpalm; four V trichobothria; V 3 and V 4 
separate; Esb trichobothrium more distal than Eb group, midway between Eb group 
and Est; Eb 3 trichobothrium close to Ebj_ 2 ; Est trichobothrium midpalm or nearly so. 
Chela fixed finger, db trichobothrium on dorsal surface; esb, eb, est, et equidistant 
(distance est-esb similar to distance esb-eb) or est closer to esb; eb trichobothrium at 
base of finger, behind point of articulation between fixed and movable fingers, above 
esb-et axis; esb trichobothrium below est-et axis, at base of finger, behind point of 
articulation between fixed and movable fingers; two i trichobothria. Patella: d 2 tricho- 
bothria distal to patellar spur; et séries with three trichobothria; est séries with one 
trichobothrium; em séries with two trichobothria; esb séries with two trichobothria; 
em-esb séries with four trichobothria m two groups esb j_ 2 and em j_ 2 ; eb séries with 
five trichobothria in two groups eb 1 and eb 2 _ 5 or eb 1 leb 4 _ 5 and eb 2 _ 3 ; three v tricho - 
bothria. 

Carinae: Chela manus: dorsal secondary carina obsolète; digital carina distinct, 
costate or granular, stronger than external secondary carina; external secondary carinae 
absent or obsolète; ventroexternal carina granular or crenulate; ventromedian and 
ventrointernal carinae absent or obsolète; internomedian carina présent, granular (at 
least sparsely or faintly). Patella: prolateral process présent (distinct), low, forming two 
distinct spiniform processes (bifid), thèse sometimes fused but not forming a large 
médian spine; internodorsal and dorsomedian carinae distinct; dorsoexternal carina 



LIOCHELES SCORPIONES, LIOCHELIDAE 



749 




FIG. 13 

Liocheles neocaledonicus (Simon, 1877), female paratype (ZMH), pedipalp with trichobothrial 
pattern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) 
Trochanter, fémur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral 
aspect. Scale line, 2.5 mm. 



750 



L. MONOD 



absent or obsolète; externomedian carinae granular or costate; ventroexternal carina 
présent and distinct, costate or granular. Fémur: dorsoexternal carina présent and 
distinct, at least as a ridge, usually more distinct in proximal half, equally developed 
than dorsointernal carina; dorsointernal carina présent and distinct; internomedian 
dorsal carina absent, without spines or granules (or with a single basai spine); interno- 
median ventral carina absent, without spines, or at most vestigial, with a single spine 
at proximal extremity; ventromedian carina absent or obsolète; ventrointernal carina 
présent, granular. 

Macrosculpture: Chela fingers granular (at least sparsely in proximal half). 
Chela fixed finger: area around dbldsbldst trichobothria granular (at least in the most 
proximal part), dbldsbldst trichobothria in three distinct smooth dépressions (one 
around each trichobothrium). Chela: dorsal intercarinal surface entirely and densely 
granular, with medium-sized spiniform granules; retrolateral intercarinal surface 
granular; ventral intercarinal surface with granulation présent along prolateral and 
retrolateral edges only, smooth medially; prolateral intercarinal surface at least sparsely 
granular, granulation less distinct along prolateral ventral edge. Patella: dorsal and 
ventral intercarinal surfaces entirely granular, at least a reticulated network of granules 
présent; retrolateral intercarinal surface at least sparsely granular; prolateral inter- 
carinal surface: proximal half at least sparsely granular, distal half usually less granu- 
lar. Fémur: dorsal intercarinal surface densely granular except for distal end; dorsal and 
prolateral intercarinal surfaces with small to medium-sized spiniform granules; retro- 
lateral and prolateral intercarinal surfaces at least sparsely granular; ventral intercarinal 
surface granular proximally, distal part without granulation. 

Carapace: Anterior margin with shallow médian notch. Anterior furcated 
suture/sulcus présent and distinct. Médian ocular tubercle situated anteromedially, 
medium-sized, occupying about 1/7 of carapace width, at least slightly raised; super- 
ciliary carinae présent, with at least few granules. Médian ocelli separated by at least 
half diameter of médian ocellus, at least twice the size of latéral ocelli. Three pairs of 
latéral ocelli; ocelli of equal size, equidistant, close together, almost touching each 
other. Margin behind latéral ocellus without spines. Carapace entirely granular, at least 
sparsely; large spiniform granules on anteromedian surface, rest of prosoma usually 
with smaller spiniform granules, evenly and sparsely distributed. 

Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without 
distinct prominence; posterior margin of pretergites smooth, without spines of 
granules. Posttergites I-VII, with minute spiniform granules, uniformly and sparsely 
distributed; I-VI entirely granular, at least sparsely in posterior half, granulation absent 
or vestigial on ridges in some spécimens; VII entirely granular, at least sparsely. 
Posttergites III-VII with distinct reticulated network of ridges and dimples, surface 
uneven; I-VI with latéral transversal sulcus. 

Ventral morphology: Anterodistal tip of coxa III without swelling or bulge. 
Sternum (Fig. 14A) subpentagonal (anterior width approximately equal to or slightly 
less than posterior width); length less than or equal to posterior width. Pectines 
(Fig. 14A) moderately long (distal edge reaching, but not surpassing distal edge of 
coxa of leg IV), with fulcrae and three marginal lamellae; 7-12 pectinal teeth, long, 
straight, entirely covered by sensory papillae. Stigmata (spiracles) half-moon shaped 



uochei.es scorpiones, liochelidae 



751 




FIG. 14 

Liocheles neocaledonicus (Simon, 1877), pectines and génital opercula, ventral aspect. (A) Maie 
(H. sarasini paralectotype, ZMH). (B) Female {H. sarasini paralectotype, ZMH). Scale line, 
1 mm. 



752 



L. MONOD 



(with a distinct curve), short (less than 1/3 of sternite width). Sternite VII without 
longitudinal carinae. 

Metasoma (Fig. 15C): Segments I-V as long as or slightly longer than in female 
(Fig. 15D), not flattened laterally. Segments I-IV with dorsomedian furrow shallow, 
weak to absent or only visible on segments I-III; dorsosubmedian and dorsolateral 
carinae absent or obsolète; ventrolateral and paired ventrosubmedian carinae présent 
and distinct on at least some segments. Segment I: width less than or equal to height; 
médian latéral carina présent and distinct; dorsosubmedian carinae with posterior 
spiniform granules weak to absent (not noticeably larger than preceding granules); 
dorsomedian posterior spiniform granules weak to absent; ventral surface without 
posterior spiniform granules; ventrosubmedian carinae with 1-2 pairs of moderate to 
strong subposterior spiniform granules and 1-3 pairs of well developed medial 
granules. Segment II: dorsosubmedian carinae with posterior spiniform granules weak 
to absent (not noticeably larger than preceding granules); dorsomedian posterior spini- 
form granules weak to absent; ventrolateral carinae without posterior spiniform 
granules; ventrosubmedian carinae without posterior spiniform granules, with one pair 
of subposterior and 1-3 pairs of medial well developed spiniform granules. Segment 
III: dorsosubmedian carinae with posterior spiniform granules moderate to strong 
(distinctly larger than preceding granules); ventrolateral carinae with ridges indistinct 
or weak and smooth; ventrosubmedian carinae with ridges indistinct or weak and 
smooth (sometimes with a few reduced spiniform granules), with one pair of well 
developed subposterior spiniform granules. Segment IV: dorsosubmedian carinae with 
posterior spiniform granules moderate to strong (distinctly larger than preceding 
granules); ventrosubmedian carinae with ridges indistinct or weak and smooth (some- 
times with few reduced spiniform granules in posterior half), without subposterior 
spiniform granules; ventrolateral carinae with ridges indistinct or weak and smooth. 
Segment V: dorsal surface smooth, without smooth shiny dépression in posterior half 
between ventrolateral carinae; dorsolateral carinae absent or obsolète; ventromedian 
carina absent or obsolète, ridges absent or weak and smooth; ventrolateral carinae 
présent and distinct, smooth or nearly so (rarely one pair of vestigial granules 
posteriorly); anal arch crenulate, at least with few reduced teeth. Telson as long as or 
slightly longer than metasomal segment V; vesicle smooth, without granules, un - 
modified. 

Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae 
often weakly developed). Fémur IV with ventromedian carinae vestigial (only 
expressed distally) or indistinct (only .scattered granules). Tibiae I-II: retrolateral 
margins without spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spini- 
form macrosetae; retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV 
(Fig. 15B): two ventrosubmedian rows of secondarily setiform macrosetae; ventro- 
median row of spinules absent or vestigial; basai spinules présent (at least one, usually 
in a short row); terminal ventromedian spinules absent; ungues shorter than telotarsus. 
Telotarsi I-IV: prolateral/retrolateral rows with 4/4, 4/4, 5/5 and 5/5 spiniform macro- 
setae respectively. 

Hemispermatophore (Fig. 16): Distal lamina at least slightly curved, approxi- 
mately of same size as basai part or slightly longer; distal crest absent; single lamellar 



LIOCHELES SCORPIONES, LIOCHELIDAE 



753 




FIG. 15 



Liocheles neocaledonicus (Simon, 1877). (A) Carapace of female paratype (ZMH), dorsal 
aspect. (B) Left tarsus IV of maie {H. sarasini paralectotype, ZMH), ventral aspect. (C) 
Metasoma of maie {H. sarasini paralectotype, ZMH), latéral aspect. (D) Metasoma of female 
paratype (ZMH), latéral aspect. Scale lines, 2.5 mm (A), 2 mm (C, D), 0.5 mm (B). 



754 



L. MONOD 




FIG. 16 

Liocheles neocaledonicus (Simon, 1877), maie (LM, Mount Koghis), left hemispermatophore. 
(A) In toto, dorsal aspect. (B) Détail of capsular région, entai aspect. (C) Idem, ventral aspect. 
Scale lines, 1 mm. 

hook in basai 1/3 of distal lamella or below; basai extrusion absent; transverse ridge 
présent and distinct, merging with anterior edge above base of lamellar hook, at 
approximately same level as base of lamellar hook. Lamella thin, folded only 
proximally and unfolded towards flattened distal extremity (tip and base approximately 
of same width); longitudinal 'spine', accessory hook and accessory lobe absent; 
lamellar tip below base of lamellar hook and above tip of distal lobe. Distal lobe well 
developed as a distinct hump, without distinctive hook-like shape, without accessory 
hook, carinae or crest. Basai lobe well developed (spoon shaped), merging with 
accessory anterior basai lobe; distal edge without accessory fold toward ectal part, 
forming a 90° angle with lamella; basai edge without accessory fold (no groove), 
forming a 135-150° angle with lamella. 

Book lungs (after Kamenz & Prendini, 2008): Lamellar surface with simple 
trabeculae. Lamellar distal edges with arcuate bow-like structures. Posterior spiracle 
edge with chisel-like structures. 



LIOCHELES SCORPIONES, LIOCHELIDAE 



755 



Table 3 

Meristic data for adult maies and females of Liocheles neocaledonicus (Simon, 1877). 







maies 






females 






Carapace 


Length 


7.2 


6.2 


6.9 


7.8 


6.4 


8.6 


Anterior width 


4.2 


3.8 


4.8 


4.7 


3.9 


5.4 




Posterior width 


8.6 


7.4 


8.0 


9.4 


7.7 


9.3 


Metasomal 


Length 


2.7 


2.3 


2.4 


2.4 


2.1 


2.5 


segment I 
















Width 


2.1 


1.9 


1.9 


2.1 


1.9 


2.3 


Metasomal 
















segment V 


Length 


4.5 


4.2 


4.4 


4.4 


3.6 


4.8 




Width 


1.5 


1.2 


1.4 


1.7 


1.3 


1.8 




Height 


1.6 


1.4 


1.5 


1.8 


1.4 


1.8 


Telson vesicle 


Width 


1.6 


1.3 


1.5 


1.8 


1.3 


1.7 




Height 


1.7 


1.4 


1.6 


1.8 


1.4 


1.8 


Pedipalps 


Fémur length 


6.5 


5.6 


6.5 


6.6 


5.2 


7.4 




Fpmur width 

1 C 1 J 1 Ul VV IVitll 


3.0 


2.6 


2.9 


3.2 


2.5 


3.4 




Patella length 


6.8 


5.9 


6.8 


7.4 


5.6 


8.1 




Patella width 


3.2 


2.7 


3.4 


3.4 


2.6 


4.2 




Chela length 


14.9 


12.8 


14.3 


15.4 


12.0 


16.9 




Chela width 


5.8 


4.8 


5.3 


6.2 


4.7 


6.4 




Chela height 


3.5 


2.6 


3.3 


3.5 


2.5 


4.2 




Chela movable 
















finger length 


7.7 


6.7 


7.5 


7.9 


6.2 


9.1 


Total length 




49 


40 


44 


51 


42 


58 



Description of adult female: Same characters as in maie except as follows. 

Pedipalp: Chela fingers linear or nearly so, without lobes and notches. 

Ventral morphology (Fig. 14B): Génital operculum oval to semi-oval, as wide 
as long (or wider than long), approximately same width as sternum; sclerites partly 
fused, médian suture distinct; posterior notch weakly developed. Pectines short (distal 
edge not reaching distal edge of coxa of leg IV). Six to eleven pectinal teeth covered 
with sensory papillae only in distal portion. 

INTRA-SPECIFIC VARIABILITY: In maies the lobe on the cutting edge of the 
movable finger and the corresponding notch on the fixed finger of the pedipalp chela 
can be more or less developed. Some spécimens display very low lobes, while others 
have more pronounced lobes. Moreover, the fitting between the lobe and the notch is 
also variable; in most spécimens there is no gap between them, or at most a very 
narrow posterior gap. However, some maies have slighty larger posterior gaps. Bigger 
spécimens tend to have a more pronounced lobe and notch, and usually also display 
larger posterior gaps. The posterior spiniform granules on the dorsosubmedian carinae 
of metasomal segments III and IV are usually distinctly larger than the preceding 
granules, however there is no clear size différence in some spécimens. Pectinal teeth 
count variation is as follow: 7-12 in maies and 6-11 in females. 



756 



L. MONOD 



ACKNOWLEDGEMENTS 

An extramural training at the MHNG in 1998 was partially funded by the 
Department of Cultural Affairs of the City of Geneva. I am deeply indebted to the 
following persons: Wilson Lourenço and Volker Mahnert who supervised the présent 
study; the late Jacqueline Heurtault and Yves Coineau who provided work space and 
infrastructure at the MNHN; Peter Schwendinger, Bernd Hauser and Charles Lienhard 
who did the same at the MHNG. I also wish to thank the following persons: (1) for 
granting access to the collections of their respective institutions, or providing and pre- 
paring loans: Léon Baert (MRHN); Johnathan Coddington and Dana de Roche 
(USNM); Hieronymus Dastych (ZMH); Paul D. Hillyard and Janet Beccaloni 
(NHML); Charles Griswold (CAS); Mark S. Harvey and Julianne Waldock (WAM); 
Franz Krapp (ZMAK); Erik Van Nieukerken, B. Van Bekkum-Ansari and Leonne 
Vermond (NNHM); Lorenzo Prendini, Randy Mercurio and Jeremy Huff (AMNH); 
Robert Raven, Phil Lawless and Owen Seeman (QM); Al Samuelson (BPBM); (2) for 
providing helful scientific advice, information, comments and/or support: Alain de 
Chambrier (MHNG), Danièle Decrouez (MHNG), Yvan Lôbl (MHNG), Dmitri 
Logunov (MM), Lorenzo Prendini (AMNH), Christine Rollard (MNHN) and Peter 
Schwendinger (MHNG); (3) for providing bibliographie références: the late N. A. 
Locket, Victor Fet and Matt E. Braunwalder (Arachnodata); (4) for making the habitus 
color illustrations: Vladimir Timokhanov; (5) for taking the habitus photographs: 
Claude Ratton (MHNG); (6) for issuing collecting permits in New Caledonia and/or 
assisting with the permit application process: Jean-Jérôme Cassan (DDEE), Richard 
Farman (DRN), Cendrine Meresse (DRN); (7) for assistance and hospitality during 
field work: Delphine Gaillard, Sylvie Cazeres (IAC), Christian Mille (IAC) and 
Thierry Nuques (IAC); (7) for their precious advice concerning the réalisation of 
various illustrations: Michèle Bertoncini (MNHN) and Jacques Rebière (MNHN); (8) 
for kindly reviewing the manuscript and providing useful constructive comments: 
Peter Schwendinger and Erich Volschenk. 

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REVUE SUISSE DE ZOOLOGIE 



Tome 118 — Fascicule 4 



Pages 

Schwendinger, Peter J. & Ono, Hirotsugu. On two Heptathela species 
from southern Vietnam, with a discussion of copulatory organs and 
systematics of the Liphistiidae (Araneae: Mesothelae) 599-637 

Landry, Bernard, Roque-Albelo, Lazaro & Hayden, James E. A new 
genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the 
Galapagos Islands, Ecuador 639-649 

Kadej, Marcin & Hâvas, Jin. A new species of Anthrenus Geoffroy, 1762 

(Coleoptera: Dermestidae) from Oman, with a key in related species. 65 1 -657 

Mahnert, Volker & Schmidl, Jurgen. First record of the subfamily 
Pycnocheiridiinae from South America, with the description of 
Leptocheiridium pfeiferae gen. n., sp. n. (Arachnida: Pseudoscor- 
piones: Cheiridiidae) 659-666 

Shi, Li, Yang, Ding & Gaimari, Stephen D. Four new species from China 

and Southeast Asia (Diptera, Lauxaniidae, Homoneurinae) 667-693 

Lôbl, Ivan. On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphi- 

diinae) of the Philippines, II 695-721 

Monod, Lionel. Taxonomic emendations in the genus Liocheles 

Sundevall, 1833 (Scorpiones, Liochelidae) 723-758 



REVUE SUISSE DE ZOOLOGIE 



Volume 1 1 8 — Number 4 

Pages 

Schwendinger, Peter J. & Ono, Hirotsugu. On two Heptathela species 
from southern Vietnam, with a discussion of copulatory organs and 

systematics of the Liphistiidae (Araneae: Mesothelae) 599-637 

Landry, Bernard, Roque-Albelo, Lazaro & Hayden, James E. A new 
genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the 

Galapagos Islands, Ecuador 639-649 

Kadej, Marcin & Hâvas, Jirï. A new species of Anthrenus Geoffroy, 1762 

(Coleoptera: Dermestidae) from Oman, with a key in related species. 651-657 

Mahnert, Volker & Schmidl, Jùrgen. First record of the subfamily 
Pycnocheiridiinae from South America, with the description of 
Leptocheiridium pfeiferae gen. n., sp. n. (Arachnida: Pseudoscor- 

piones: Cheiridiidae) 659-666 

Shi, Li, Yang, Ding & Gaimari, Stephen D. Four new species from China 

and Southeast Asia (Diptera, Lauxaniidae, Homoneurinae) 667-693 

Lôbl, Ivan. On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphi- 

diinae) of the Philippines, II 695-721 

Monod, Lionel. Taxonomic emendations in the genus Liocheles 

Sundevall, 1833 (Scorpiones, Liochelidae) 723-758 



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clature must be strictly followed. Cite the authors of species on their first mention. 

Results. Thèse 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 ail new taxa, synonymies or combinations. Primary types must 
be deposited in a muséum or similar institution. In taxonomic papers the species heading should be followed by 
synonyms. material examined. description, distribution, and comments. Ail material e.xamined 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 "maie" and "female" (text file: S = 6 . £ = 9). 

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. 

Références. The author-date system (name-year System) must be used for the citation of références in the text. 
e.g. White & Green ( 1995) or (White & Green. 1995). For références with three and more authors the form Brown 
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below). The list of références must include ail publications cited in the text and only thèse. Références 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 référence. 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. Thèse, Genève. 43 pp. 
Pknard. E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61 . 
Mertens. R. & Wermith. 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 Muséum of Natural History. Chicago. XII + 861 pp. 

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