A revision of the genus Zyras (Zyras) Stephens, 1835 (Coleoptera, Staphylinidae, Aleocharinae). I. Current classification status and the redefinition of the genus

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DOI: 10.3897/zookeys.29.218
Abstract
The genus Zyras is redefined and redescribed based on the study of the type species Zyras haworthi (Stephens). Illustrations of all important morphological characters are provided. The status of the genus Zyras (s.l.) is discussed. A list of all species attributed to the subgenus Z. (Zyras), including the species described as Zyras (s.str.), is also given.
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 49
A revision of the genus Zyras (Zyras) Stephens, 1835
(Coleoptera, Staphylinidae, Aleocharinae).
I. Current classification status and
the redefinition of the genus
Peter Hlaváč1, Tomáš Jászay2
1 Na doline 14, SK-040 14 Košice, Slovakia 2 Museum of Šariš, Natural History Department, Radničné nám.
13, SK-085 01 Bardejov, Slovakia
Corresponding author: Peter Hlaváč (phlavac@stonline.sk)
Academic editor: Jan Klimaszewski|Received 25 October 2009|Accepted 10 November 2009|Published 11 December2009
Citation: Hlaváč P, Jászay T (2009) A revision of the genus Zyras (Zyras) Stephens, 1835 (Coleoptera, Staphylinidae,
Aleocharinae). I. Current classi cation status and the rede nition of the genus. Zookeys 29: 49–71. doi: 10.3897/
zookeys.29.218
Abstract
e genus Zyras is rede ned and redescribed based on the study of the type species Zyras haworthi
(Stephens). Illustrations of all important morphological characters are provided.  e status of the genus
Zyras (s.l.) is discussed. A list of all species attributed to the subgenus Z. (Zyras), including the species
described as Zyras (s.str.), is also given.
Keywords
Coleoptera, Staphylinidae, Aleocharinae, Lomechusini, Zyras, taxonomy, myrmecophily
Introduction
e Lomechusine genus Zyras Stephens, with currently 802 species and 54 subgenera
(Hlaváč, Newton and Maruyama, in prep.), is certainly one of the largest and most
problematic genera of the subfamily Aleocharinae. All species of the genus are believed
to be free predators (synechtrans) of various species of ants.  e taxonomic chaos in
this genus is manifested by the fact that 122 species were described only as Zyras with-
out any a lation to a subgenus. Another 68 species originaly described as Zyras were
subsequently synonymyzed or transferred to di erent genera.
ZooKeys 29: 49–71 (2009)
doi: 10.3897/zookeys.29.218
www.pensoftonline.net/zookeys
Copyright P. Hlavácˇ, T. Jászay. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
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Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
50
Until 2006, seven subgenera already have been raised to generic level, Myrmoe-
cia (Mulsant & Rey) by Seevers (1978), Aulacocephalonia (Bernhauer) and Homalo-
donia (Bernhauer) by Kistner and Jacobson (1981), Paramyrmoecia (Scheerpeltz) by
Kistner and Elliot (1985), Macrogerodonia (Bernhauer) and Stichodonia (Bernhauer)
by Pace (1986a, 1999b) and Trachydonia (Bernhauer) by Kistner (1997). Maruyama
(2006) revised the complex of subgenera Pella Stephens, Lepla Tottenham, Pellochro-
monia Reit ter and Myrmelia Mulsant & Rey, in which he synonymyzed all these taxa
with Pella and Pella was treated as avalid genus. Finally, Assing (2009) raised Peltodo-
nia (Bernhauer) to the genus level.  e genus Razia Blackwelder, 1952 (new name for
Allocota Bernhauer, 1916; not to be confused with Aloconota omson, 1958) is now
placed in the tribe Homalotini and it is a valid subgenus of Gyrophaena Mannerheim,
1830 (Newton and  ayer 2003).
As a consequence of these changes, the following 52 subgenera of the genus Zyras
have remained valid (Hlaváč, Newton and Maruyama, in prep.): Acanthocnemidonia
Bernhauer, Acrothoraconia Bernhauer, Androdonia Bernhauer, Anophthalmodonia Bern-
hauer, Antronia Bernhauer, Aplastonia Bernhauer, Apostenonia Bernhauer, Apterygodo-
nia Scheerpeltz, Aulacodonia Bernhauer, Botsa Blackwelder, Callodonia Bernhauer, Ca-
meronodonia Dvořák, Camonia Bernhauer, Cephalodonia Bernhauer, Colpodonia Bern-
hauer, Craspa Blackwelder, Crateodonia Bernhauer, Ctenodonia Wasmann, Dentothal-
monia Last, Diaulaconia Bernhauer, Euryalonia Bernhauer, Eurydonia Bernhauer, Eu-
ryncephalodonia Scheerpeltz, Euryndonia Bernhauer, Fealina Bernhauer, Glossacantha
Gemminger & Harold, Grammodonia Bernhauer, Hylozyras Iabloko -Khnzorian, Is-
othoracodonia Bernhauer, Lastia Dvořák, Leptodonia Bernhauer, Neotropopella Sche-
erpeltz, Pachydonia Bernhauer, Paragrammodonia Bernhauer, Parophthalmonia Bern-
hauer, Platydonia Bernhauer, Polydonia Bernhauer, Pycnodonia Bernhauer, Remionia
Blackwelder, Rhopalodonia Cameron, Rocnema Blackwelder, Sinozyras Pace, Subversoris
Last, Synthoracodonia Scheerpeltz, Taprodonia Cameron, Termidonia Motschulsky, Ter-
mitelia Cameron, Termitodonia Cameron, Trigonodonia Bernhauer, Trigonozyras Ca-
meron, Tropidonia Bernhauer, Visendor Last.
e logical question must be “Is it really possible that all these very numerous taxa,
distributed worldwide, form a monophyletic group?”  is question is not new and it
has already been raised twice,  rst by Seevers (1965: 238) [“ e taxonomic state of the
vast Zyras complex is one of great confusion and the proposal of more than 50 subgenera has
not helped clarify matters very much”] and later by Kistner (1972: 148) [“Also each species
alleged to Zyras (s. str.) in the literature will need to be checked before its inclusion in the
rede ned genus is settled. For example, I have not seen a species from sub-Saharan Africa
which really belongs to this genus. It is highly likely that Zyras in the future will be a very
small genus”]. We fully agree with these two opinions, except for the number of species
of true Zyras, which is anticipated to grow to at least 200 species.
e distribution of the genus seems to be mainly in the holarctic region with only
three species in the USA and Canada (Klimaszewski et al. 2005),  ve species in the
western Palaearctic region and a large number of species in eastern Palaearctic, Orien-
tal and perhaps also in the Australian regions. We have not seen, like Kistner (1972),
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 51
any true Zyras from sub-Saharan Africa or from America south of USA. Of six species
of Zyras reported by Ashe (Navarrete-Heredia et al. 2002) from Mexico, two species
already have been transferred to another genus, Pella fauveli (Sharp) and Myrmoecia
tapinomatis Mann (Klimaszewski et al. 2005).  e remaining four species should be
checked, but they, highly likely, will not be members of true Zyras either.
We are sure that many, if not all, species currently attributed to di erent subgenera
of Zyras, have nothing or very little in common with the true Zyras. For the subgenera
Diaulaconia (Hlaváč 2005: 153) and Glossacantha (Hlaváč 2005; Maruyama 2006,
respectivelly) that has been already discussed, although the new status of both genera
has not been o cially changed. In our opinion, Zyras is a very characteristic and well
de ned genus, and any further fragmentation should be carefully re-considered.
e majority of subgenera, still attributed to Zyras, were very poorly de ned main-
ly by Bernhauer [34 subgenera described in the period of 1928–1936, four were later
renamed by Blackwelder (1952) and one is synonym], Cameron [six subgenera, one
renamed later by Dvořák (1981)], Scheerpeltz (four subgenera in 1963–1976), Last
[4 subgenera, one later renamed by Dvořák (1984)] and Wasmann (3 subgenera, one
synonym) (Hlaváč, Newton and Maruyama, in prep.). Unfortunately even the  nal
and most recent papers by Pace (1999a, 1999c) did not bring any light to the Zyras
taxonomic problem.
Material and methods
e main goal of this article is to rede ne the genus Zyras based on its type-species,
Zyras haworthi (Stephens).  e material used for this study was taken from some insti-
tutions listed below.
Specimens prepared for the morphological study were examined with Leica S8A-
PO stereo-microscope and Nikon SMZ – 1B stereo-microscope with di use lighting
at magni cations up to 105×. Male genitalia and other dissected parts were studied
using a Meopta transmitted-light microscope at magni cation up to 450×. Genital
segments were dissected and treated with lactic acid. All drawings were made using
a drawing tube.  e dissected and mounted parts were mounted and pinned with
the specimen.
Depository abbrevations
e material is deposited in following collections:
CMS Collection of Michael Schülke, Berlin, Germany;
CPH Collection of Peter Hlaváč, Košice, Slovakia;
CVA Collection of Volker Assing, Hannover, Germany;
NMW Naturhistorisches Museum (Harald Schilhammer), Vienna, Austria;
SDEI Senckenberg Deutsches Entomologisches Institut (Lothar Zerche), Germany;
SMB Šarišské Múzeum Bardejov (Tomáš Jászay), Slovakia.
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
52
Tribe Lomechusini Fleming, 1821
Genus Zyras Stephens, 1835
Zyras Stephens, 1835: 430. Type species: Zyras haworthi (Stephens), established by
decision of the International Commission on Zoological Nomenclature (ICZN
1961, Opinion 599)
Zyras Stephens: Cameron, 1939: 497 (key to subgenera and species of India, Myanmar
and Sri Lanka); Kistner 1972: 143 (redescription); Scheerpeltz 1974: 39 (key to
African subgenera); Seevers 1978: 153 (key to Nearctic species); Dvořák 1980 (key
to species of Czech and Slovak Republic, including Pella, Lepla, Pellochromonia,
Zyras, Myrmoecia); Dvořák 1984: 200 (key to Palaearctic species including China);
Hastir and Gaspar 2001 (biology); Klimaszewski et al. 2005: 714 (restricted rede-
nition, key to species of America north of Mexico).
Diagnosis. Zyras (Zyras) can be distinguished from the other genera of Lomechusini by a
combination of the following characters: 1) whole body shiny, unicoloured or often with
di erent colouration for head, pronotum, elytra and abdomen, sometimes covered with
long setae; 2) all antennomeres petiolate, antennomere III about as long as pedicel; 3)
neck absent; 4) pronotum usually with well-de ned median antebasal fovea; 5) abdomen
simple, not physogastric, parallel-sided; 6) simpli ed aedeagus, median lobe with enlarged
basal capsule and narrowly conical apical lobe; 7) spermatheca sclerotized, very small with
narrowly elongate capsule and with extremely long and highly coiled spermathecal duct.
Description. Body (Fig. 1) slender, subparallel-sided. Body length highly variable,
ranging from 3–9 mm, whole body always shiny, unicoloured, or often with di erent
colouration for head, pronotum, elytra and abdomen.
Head (Figs 2, 3) slightly wider than long, posterior margin slightly covered by an-
terior edge of pronotum, neck absent; temples long, round, about as long as diameter
of eyes or longer; occipital suture present, not visible dorsally, ventrally reaching hy-
postoma, hypostoma narrow; surface with erect or appressed setae. Gula long, evenly
divergent from anterior to posterior; submentum fused to gula, broadly expanded an-
teriorly. Eyes large, oval in lateral view, prominent.
Antennae (Fig. 4) with all antennomeres petiolate, antennomere III about as long
as II, when bent backwards slightly exceeding base of pronotum.
Labrum (Fig. 5) much wider than long, with shallow median excavation; surface
covered with numerous pseudopores and about ten real pores, except on posterior and
lateral areas; antero-lateral areas each with 4–6 macrosetae of di erent length.
Mandibles (Figs 6–9) almost symmetrical, lacking teeth, with 3–4 small setae
present dorsally, mesal areas of dorsal and ventral surfaces covered with numerous
pseudopores, prostheca with inner margin pubescent.
Maxilla (Fig. 10) is generalized in shape like for other Lomechusini, with elongate galea
and shortened lacinia, palpomere I minuscule, palpomere II slightly curved and gently short-
er that palpomere III, terminal palpomere about three times as short as III, pointed apically.
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 53
Mentum (Fig. 11) trapeziform, posterior and anterior margins truncate, with setae,
surface covered with pseudopores bearing minute setae, and about 6–8 pores with
longer setae present on antero-lateral areas.
Labium (Fig. 12) with prementum with two real pores and one setal pore meso-
laterally, with about 30–40 pseudopores behind medial setae, apodeme with large,
truncate process, lateral lobes of apodeme gently curved, assimetrical, bifurcate apicaly.
Ligula bilobed, each lobe long, pointed apically, with  ne apical seta. Labial palpus
with  rst segment long, more than twice as long as second and clearly thicker, third
segment slightly longer than second and about twice as slender as second.
Pronotum slightly wider than long (Fig. 1), anterior margin straight, posterior mar-
gin evenly rounded, posterior corners small but well-de ned, smooth or entirely or
Figure 1. Zyras (Zyras) haworthi (Stephens), dorsal view (approximative length of photograped speci-
men is 7.1 mm)
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
54
Figures 4–5. Zyras (Zyras) haworthi (Stephens). 4 antennae (scale = 0.5mm) 5 labrum, dorsal (scale = 0.1mm)
Figures 2–3. Zyras (Zyras) haworthi (Stephens). 2 head, dorsal aspect 3 head, ventral aspect (scale = 0.5mm)
2
45
3
partially coarsely punctured and sparsely covered with erect long setae, macrosetae on
lateral margins present, well-de ned, median antebasal fovea usually present (may be
missing in some species, for instance Z. bakerianus).
Elytra subparallel-sided (Fig. 1) with well-de ned sutural groove throughout, com-
bined width of elytra about twice as wide as long on suture, sutural striae well-de ned
through whole length. Scutellum triangular.
Mesoventrite (Figs 13, 14) much shorter than metaventrite, coarsely and densely punc-
tured, setation only on sides, mesoventral process large, round. Mesocoxal cavities sepa-
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 55
rated, isthmus wide but short. Metaventrite rectangular, slightly wider than long, shining,
irregularly and coarsely punctured, with long erect setae, metaventral process truncate.
Legs (Kistner, 1972: 145, Fig. 3, B, C, D) with procoxae very long, only slightly
shorter than profemur, mesocoxae suboval, metacoxae subtriangular; pro- and mes-
otrochanter small, metatrochanter slightly larger, rounded at apex; femora  attened,
narrowest at base, with punctures and setation; all tibiae straight, very slightly dilated
apicad, with dense setation, each apex with two stout setae; tarsal formula 4-5-5, tarsi
generalised, metatarsus twice as long as protarsus.
Abdomen simple, not physogastric, parallel-sided, at base with punctures and with
long setae on side; paratergites well-developed on tergites III–VII; tergites with pos-
terior margin straight; sternite with surface moderately to densely covered with setae;
eighth abdominal segment (Figs 15–18) species characteristic and also bearing sexual
characters; tergite VIII (Figs 15, 17) narrowed posteriad, truncate apically, apex crenate
or dentate, sternite VIII (Figs 16, 18) narrowed posteriad, at apex rounded in male
and truncate in female, apical setae in female longer and thicker than in male. Ninth
tergite as in (Figs 19, 20), lateral lobes in male asymmetric, absent in female as in most
Aleocharines (Maruyama 2006). Ninth sternite of male as in (Figs 21, 22) elongate,
somewhat pointed at base, expanded apicad and truncate at apex, with well-de ned
sharp corners, oblong in middle, bearing setae which are longer and dense on sides.
Aedeagus (Figs 23–25) simple, median lobe composed of enlarged basal capsule and
narrowly conical apical lobe, copulatory piece weakly-de ned or absent. Parameres (Fig. 26)
long, clearly longer than apical lobe, composed of condylite, paramerite and velum, apical
lobe of paramerite bears four macrosetae, apical lobe elongate, exceeding apex of velum.
Spermatheca (Fig. 26) sclerotized, very small with narrowly elongate capsule, cap-
sule curved and slender apicad, with extremely long spermathecal duct, coiled many
times forming a bundle.
Biology. In general, it is believed that all species of Zyras (Zyras) are free living
predators or synechtrans of di erent species of ants according to Wasmann’s categori-
Figures 6–9. Zyras (Zyras) haworthi (Stephens). 6 left mandible, dorsal aspect (scale = 0.1mm) 7 rigth
mandible, dorsal aspect (scale = 0.1mm) 8 rigth mandible, ventral aspect (scale = 0.1mm) 9 left mandible,
ventral aspect (scale = 0.1mm)
6879
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
56
Figures 10–12. Zyras (Zyras) haworthi (Stephens). 12 maxilla (scale = 0.1mm) 11 mentum (scale =
0.1mm) 12 labium (scale = 0.1mm)
Figures 13–14. Zyras (Zyras) haworthi (Stephens). 13 metaventrite and mesoventrite lateral (scale =
1mm) 14 metaventrite and mesoventrite ventral (scale = 1mm)
13
10
11
12
14
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 57
Figures 15–18. Zyras (Zyras) haworthi (Stephens). 15 tergite VIII male 16 sternite VIII male 17 tergite
VIII female 18 sternite VIII female (scale = 0.5 mm)
15
16
17
18
Figures 19–20. Zyras (Zyras) haworthi (Stephens). 19 tergites IX and X male (scale = 0.5 mm) 20
tergites IX and X female (scale = 0.5 mm)
19 20
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
58
Figures 21–22. Zyras (Zyras) haworthi (Stephens). 21 sternite IX male (scale = 0.5 mm) 22 apex of
sternite IX male (scale = 0.1mm)
Figures 23–24. Zyras (Zyras) haworthi (Stephens). 23 aedeagus, ventral aspect 24 aedeagus, lateral aspect (scale = 0.5mm)
21 22
23 24
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 59
zation (Hölldobler and Wilson, 1990). However there is almost no clear evidence for
this. Hastir and Gaspar (2001) showed that it is di cult to classify Zyras haworthi in
any of Wasmann’s categories.  ey have also shown that Zyras haworthi is more numer-
ous in localities where Formica sanguinea Latreille is also present.
Distribution. Actually worldwide distributed, but most probably restricted to the Pale-
arctic, Nearctic, Oriental and Australian regions (see discussion above, Introduction). How-
ever, due to current taxonomic problems of the genus, it is not possible to be more precise.
Zyras (Zyras) haworthi (Stephens)
Figures 1–26
Aleochara haworthi Stephens, 1832: 126. Type locality: Norfolk
Bolitochara elegans Heer, 1839: 350. Type locality: Helvetia
Zyras nigricollis Motschulsky, 1845: 41. Type locality: Tschougoue , Russie méridionale
Material examined. Austria: 1 : Austria, Mistelbach env., Eichenwald, 11.V.2002/M.
Schülke (CMS); 1 : Bisamberg near Vienna, 25.VII.10, coll. Künnemann (SDEI); 1
: Steiermark, coll. Stierlin (SDEI); 1 : Styria, Reitter, coll. Künnemann (SDEI). Azer-
baijan: : Kasp.Meer-Geb., Talysch, 1897, von Heyden (SDEI). Bosnia and Hercego-
vina: 3♂♂: Bjelašnica Plan., Gipfelregion, Bosn., 1902, coll. Leonhard (SDEI); 2♂♂:
Figures 25–26. Zyras (Zyras) haworthi (Stephens). 25 paramera (scale = 0.5mm) 26 spermateca (scale
= 0.5mm)
25 26
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
60
Bosnia, 1902, Maklen-Pass, O. Leonhard (SDEI); 1 : Herzegovina, Jablanica, 1901,
O. Leonhard (SDEI). Bulgaria: 4♂♂, 3♀♀: NO Bulg., Senokos, 20km OTolbuchin,
4.5.-19.6.1987, leg. Penev, Querc. roboris et pedunculi orae, Barberfallen (SDEI); 1 :
Bulg., Or. Eminska Planina, Vlas, 10.V.1987, leg. Behne, Heinig, aus Laubschicht ge-
siebt unter Quercus und Ulmus carpinifolia, (CVA); 2♀♀: Bulg., Samokov, M. Hilf,
1911, Coll. O. Leonhard (SDEI); 1 : NO Bulg., Srebarna, 20km Wsilistra, 17.6.-
11.7.1987, leg. Penev (SDEI); 1 : Bulg., Rila-Geb., 23.VII.1985, Musala, 2500 m,
leg. Zerche (SDEI). Czech Republic: 3♂♂: Prag, Coll. Kraatz (SDEI); 1: Prag, Ska-
litz[ky], Coll. Weise (SDEI); 1: Prag, coll. Stierlin (SDEI); 1: Bohemia (SDEI); 1,
2♀♀: Prag, (Dieck), coll. von Heyden (SDEI); 2♀♀: Prag, Lokay, coll. von Heyden
(SDEI); 1: Moravia, Coll. Koltze (SDEI); 3♂♂, 2♀♀: Prag, Coll. Koltze (SDEI); :
Slatz, Prag, Stussiner, Coll. Weise (SDEI); 1ex.: Paskau [= Paskov], V-95, coll. Koltze
(SDEI). Croatia: 3ex.: Oestr [=Oesterreiches] Küstenland, Fužine, 1906, leg. M. Hilf,
Coll. O. Leonhard (SDEI). France: 1: France, Alpes Maritimes, Col de Castillon,
Südseite 350 m nördl. Monti, I. Wolf leg., 5.5.1996 (CMS); 1ex.(lacking last segments
of abdomen): France, Midi Pyréneés, S/Ax les  ermes, Umg. L´Hospitalet, Ariege,
1300m, 21.VI.1999, leg. I. Wolf (CVA); 1: Francia, A. Marittime, Sospel, 1.VI.1999,
500m, leg. F. Angelini (CVA); 1: Corsica, 1905, Cervione, Coll. O. Leonhard, 21.7.
(SDEI); 1, 1: Rhone, Cl. Rey, Coll. Leonhard (SDEI). Germany: 1: D-Nieder-
sachsen, E Schladen, Hedeper, Westerberg, VI.2001, L. Schmidt (CVA); 1: D. Nie-
dersachsen, Hildesheim-Ochter-Trillkegut, 10.VII.2001, Sprick (CVA); 1, 1: D.
Sachsen-Anhalt, Stendal, Badingen, 24.VI.1993, Sprick (CVA); 1: Erzgebirge, Stoll-
berg, Uhmann, 4.6.24 (SDEI); 1: Mark, Umgb., Oderberg, 29.V.27 (SDEI); 1: Er-
lang [=Erlangen], Coll. Kraatz (SDEI). Italy: 1: Italia, Lazio, Colli Albani, Commune
Rocca di Papa, Monte Cava, 600–800m, 9.05.1998, leg. I. Wolf, (CMS); 1: Calabria-
Orsomarso, Grisolia (CS), 11.IX.1993, 700m, leg. F. Angelini; 1, 1: Toscana, I.
Elba, Cavo (LI), 15.5.1998, 250m, F. Angelini (CVA); 1, 1: Calabria, Sta. Eufemia,
lg. Paganetti, coll. Franklin Műller (SDEI). Macedonia: 1: Skoplje, Mac., 25.VI.54,
leg. F. Schubert (NMW). Portugal: 1: P-Algarve, 52, W Monchique, 554m, Ru-
bus, 37°18'53N, 8°33'55W, 15.IV.2002, Meybohm (CVA); 1: P-Algarve, 46, Mon-
chique, above road to Alferce, 591m, 37°19'20N, 8°31'52W, 10.IV.2002, Meybohm
(CVA). Poland: 1: Liegnitz [=Legnica], Pfeil, coll. von Heyden (SDEI); 1: Schwd-
nitz [Schweidnitz = Šwidnica], Letzner (SDEI); 1: Gerhardt, Liegnitz [= Legnica],
Coll. Rottenberg, (SDEI); 1, 1: Mühlgast [near Wohlów.], Rottenberg (SDEI). Ro-
mania: 1: NE Romania, Negulesti, 25km SE Piatra Neamt, 500m, 18.vi.1996, P.
Hlaváč lgt. (CPH); 1: Banat 1909, Herkulesbad, leg. M. Hilf, Coll. O. Leonhard
(SDEI); 1: Banat 1909, Orsova, leg. M. Hilf, Coll. O. Leonhard, 7.VI. (SDEI). Ser-
bia: 1: Serbia, Pirot env. Zvonce, banja Prskalo, 520m presev [sifting], above river,
28.IV.2002, lgt. T. Jászay (SMB). Slovakia: 1: Slovakia (Košice), Herľany env., sifting,
1.v.2004, P. Hlaváč lgt. (CPH); 1: Slovakia centr., Bacúch, 6.vii.1990, T. Lackner lgt.
(CPH); 1: Pieniny-Poľana, zemné pasce [traps], 30.6.1989, leg. Svatoň (SMB); 1:
Nová Bošáca, St. hora, 6.5.93, Majzlan lgt. (SMB); 1: Slovakia centr., Trenčín env.,
24.6.99, O. Majzlan lgt. (SMB); 1: Parkan [= Štúrovo], Coll. Weise (SDEI). Sloveni-
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 61
ja: 1: Slo Zalog, 3.6.1996, Zg. Kašelj, B. Drovenik leg (CVA). Spain: 2♀♀: Spanien,
Lugo, Sierra Ancares, 2.8.1984, leg. J. C. Otero (CVA). Turkey: 1: TR Mersin (33),
road to Arslanköy, 5km SE Aladağ, 700m, 36°54'45N 34° 31'44E, leg. 10.5.2004 Bra-
chat & Meybohm (CVA); 1: TR.-Mersin [51], road to Arslanköy, 5km SE Aladağ,
700m, 36°54'45N, 34° 31'44E, 10.V.2004, C. Besuchet (CVA); 1: TR Antakya 10,
400m, E Yeşilkent, 30.IV.2002, 36°57N, 36°15E, Meybohm & Brachat (CVA); 2♂♂:
Ovacik, Tunceli, 1400m, Asm, F. Schubert, 6/76 (NMW).
Description. Body (Fig. 1) 5.5–7.0 mm long. Head and pronotum dark reddish-
black, elytra reddish-brown, lateral part of apices of elytra with black spot, legs yellow-
ish-brown, antennae yellowish-brown with antennomeres I–III lighter.
Head 1.15 times as wide as long, on sides with long erect golden setae. Antennae
(Fig. 4) with scape almost twice as long as pedicel, pedicel slightly longer than antenno-
mere III, antennomeres IV–VII about the same length, IV–VI slightly transverse, an-
tennomeres VII-X rhombic, clearly extended apicad, X shortest, terminal antennomere
about as long as antennomere III, almost twice as long as wide at base, round at apex.
Pronotum (Fig. 1) 1.25 times as wide as long and 1.28 times as long as head, ir-
regularly punctured, setose and with about  ve long black macrosetae on each side.
Mesoventrite (Fig. 14) regularly, coarsely punctured on the whole surface. Metaven-
trite 1.5 times as wide as long and 3.5 times as long as mesoventrite (measured in mid
line), smooth in the middle, coarsely punctured on sides, here also with erect golden setae.
Abdomen two-coloured, densely punctured apex of visible tergites III–V dark red-
dish-brown, base much lighter and bearing setae, tergites VI–VII dark reddish-brown
almost on the whole surface, densely punctured at base and with few but long apical se-
tae, paratergite III–VII well de ned covered with setae, paratergites III–V punctured.
Sexual dimorphism: females have di erent structure of abdominal tergites and
sternites VIII, IX and X (Figs 15–21). Tergite VIII with small median tubercle in male,
tubercle lacking in female, truncate apex is much narrover in male than in female, me-
dian excavation deeper in male than in female; sternite VIII round in male, truncate in
female; tergite IX with asymmetric lateral lobes in male, lateral lobes lacking in female.
Host ants. e following ant genera have been reported as hosts of Zyras (Zyras):
Formica sp., Lasius (Dendrolasius) sp., Liometopum sp., Myrmica sp.
Distribution. Almost whole Europe, Armenia, Azerbaijan, Georgia, Russia, Alge-
ria, Tunisia, Japan
Checklist of species of the subgenus Zyras (Zyras) (Stephens)
A detailed world catalogue of the tribe Lomechusini is near completion (Hlaváč, New-
ton and Maruyama, in. prep.).  e purpose of this checklist is to provide a listing of all
species which were described as, or placed subsequently in the subgenus Zyras (Zyras),
or Zyras (s.str.). All species described as Zyras without subgeneric a lation are excluded
from this list.  e list is arranged by zoogeographical regions.  is division is a practi-
cal solution for the better orientation within the genus.
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
62
Nearctic region:
is area includes America north of Mexico, i.e. USA and Canada. All three species
listed here are true Zyras and were revised recently (Klimaszewski et al. 2005).
1. obliquus (Casey, 1894: 325) (Myrmedonia). Distribution: USA, Canada
= pseudohaworthi Klimaszewski, 2002: 53
2. planifer (Casey, 1894: 326) (Myrmedonia). Distribution: USA
= schwarzi Wasmann, 1894: 207 (Myrmedonia)
3. rudis (LeConte, 1866: 372) (Myrmedonia). Distribution: USA
West Palaearctic region:
is area, as de ned here, includes the whole of Europe, northern Africa, Arabian
peninsula, Caspian region and Siberia to Lake Baikal. All  ve species recorded here are
true Zyras.
4. collaris (Paykull, 1789: 50) (Staphylinus). Distribution: Europe, Russia (Euro-
pean), Azerbaijan, Georgia, Algeria
5. cultus Last, 1960: 14. Distribution: Algeria, Eritrea, Angola
6. fulgidus (Gravenhorst, 1806: 163) (Aleochara). Distribution: Europe, Russia
(European), Georgia, Iran, Near East
7. haworthi (Stephens, 1832: 126) (Aleochara). Distribution: Europe, Armenia,
Azerbaijan, Georgia, Russia, Algeria, Tunisia, Japan
= elegans (Heer, 1839: 350) (Bolitochara)
= nigricollis Motschulsky, 1845: 41
8. maculipennis Gridelli, 1921: 87 (Zyras fulgidus var. maculipennis). Distribution:
Caucasus, Kazachstan, Uzbekistan
East Palaearctic region:
is area includes Siberia east from Lake Baikal, Nepal, northern India, China, Tai-
wan, Koreas, Japan and Russia Far East. Currently there are 36 species described from
this region and one nomen nudum.
9. abacus Dvořák, 1984: 194. Distribution: Kazakhstan, Kyrgyzstan
10. beijingensis Pace, 1993: 114. Distribution: China (Beijing)
11. britannorum Pace, 1992: 140. Distribution: Nepal
12. chinkiangensis Bernhauer, 1939: 148. Distribution: China (Jiangsu)
13. coloratus Cameron, 1939: 545 [Zyras (Pella)]. Distribution: India (Uttar Pradesh)
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 63
14. condignus Last, 1969: 279. Distribution: India (Uttar Pradesh), Nepal, Vietnam
= distinctus Cameron, 1939: 540, preoccupied, not Zyras distinctus Biering, 1937
15. cylindricornis Dvořák, 1981: 56. Distribution: Japan, Korea, China (Liaoning)
16. exasperatus Schubert, 1908: 610. Distribution: India (Himachal Pradesh)
17. fratrumkadooriorum Pace, 1998: 968. Distribution: China (Hong Kong)
18. fugax (Sharp, 1888): 289 (Myrmedonia). Distribution: Japan, Korea
19. gardneri Cameron, 1939: 538. Distribution: India (West Bengal)
20. hirsutiventris (Champion, 1927: 245) (Myrmedonia). Distribution: India (Ut-
tar Pradesh), Nepal
21. hauserianus Bernhauer, 1933b: 46. Distribution: China (Xinjiang)
22. hongkongensis Pace, 1999a: 684. Distribution: China (Hong Kong)
23. championi Cameron, 1939: 537. Distribution: India (Himachal Pradesh)
24. illecebrosus Last, 1982: 81. Distribution: Mongolia
25. iridescens (Sawada, 1970: 49). (Bolitochara) Distribution: Japan
26. kraatzi Schubert, 1908: 609. Distribution: India (Himachal Pradesh, Uttar Pra-
desh), Nepal
= ignicauda (Champion, 1927: 245) (Myrmedonia)
27. manjushri Pace, 1992b: 142. Distribution: Nepal
28. morvani Pace, 1986b: 182. Distribution: Nepal
29. nigroaeneus Cameron, 1939: 543. Distribution: India (Himachal Pradesh)
30. notaticornis Pace, 1998: 971. Distribution: China (Hong Kong, Zheijang)
31. optatus (Sharp, 1888: 289) (Myrmedonia). Distribution: Japan
32. pallipes Pace, 1992: 140. Distribution: Nepal
33. particornis (Sharp, 1888: 290) (Myrmedonia). Distribution: Japan, Russia (Far
East), Korea, China (Liaoning)
34. perforatus (Champion, 1921: 178) (Myrmedonia). Distribution: India (Sikkim/
West Bengal, Uttar Pradesh), Nepal
35. pictus (Sharp, 1874: 11) (Ilyobates). Distribution: Japan, Korea
36. pindarae (Champion, 1921: 179 (Myrmedonia). Distribution: India (Uttar
Pradesh), Nepal
37. restitutus Pace, 1993: 114. Distribution: China (Sichuan)
38. ru cauda Cameron, 1939: 543. Distribution: India (West Bengal), Nepal
39. seminigerrimus Bernhauer, 1933a: 54. Distribution: China (Sichuan)
40. shaanxiensis Pace, 1998: 971. Distribution: China (Shaanxi)
41. sibiricus Bernhauer, 1914: 68. Distribution: Russia: Siberia, Korea
42. song Pace, 1993: 112. Distribution: China (Yunnan)
43. songanus Pace, 1993: 114. Distribution: China (Beijing)
44. wei Pace, 1993: 114. Distribution: China (Guizhou)
e occurence of all species of Zyras (Zyras) from Asia is usually restricted to a rath-
er small area, with the exception of Zyras geminus Kraatz, which is widely distrbiuted
as reported in the literature.
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
64
45. geminus (Kraatz, 1859: 27) (Myrmedonia). Distribution: Sri Lanka, India, Indo-
nesia (Java, Sumatra), Taiwan , Philippines, Japan (Iriomote Island)
sinorum Rougemont, 2001: 111. Distribution: China (Hong Kong), NOMEN
NUDUM
Indochina:
is region includes southern Myanmar,  ailand, Cambodia, Vietnam and continen-
tal Malaysia. ere are currently 21 species included.
46. alboantennatus Pace, 1986a: 460. Distribution: Myanmar
47. angkoricola Pace, 2004: 292. Distribution: Cambodia,  ailand
48. bartolozzii Pace, 2003: 68. Distribution: Malaysia (Malay peninsula)
49. benenensis Pace, 2001: 196. Distribution: Vietnam
50. birmanus Scheerpeltz, 1965: 350. Distribution: Myanmar
51. chumphonensis Pace, 2004: 292. Distribution:  ailand
52. drugmandi Pace, 2004: 293. Distribution:  ailand
53. ferrugineiventris Scheerpeltz, 1965: 349. Distribution: Myanmar
54. ferrugineus Cameron, 1939: 541. Distribution: Myanmar
55. fustigans Pace, 2000a: 77. Distribution:  ailand
56. glabricollis Scheerpeltz, 1965: 358. Distribution: Myanmar
57. kambaitiensis Scheerpeltz, 1965: 347. Distribution: Myanmar
58. malaisei Scheerpeltz, 1965: 345. Distribution: Myanmar
59. nitens Cameron, 1944: 108. Distribution: Malaysia (Malay peninsula)
60. pseudobirmanus Scheerpeltz, 1965: 351. Distribution: Myanmar
61. quasar Dvořák, 1996: 6. Distribution: Vietnam
62. semiasperatus Scheerpeltz, 1965: 353. Distribution: Myanmar
63. setosipennis Scheerpeltz, 1965: 356. Distribution: Myanmar
64. setosivestis Scheerpeltz, 1965: 354. Distribution: Myanmar
65. thaiorum Pace, 1986a: 460. Distribution:  ailand
66. variolatus Pace, 2003: 68. Distribution: Malaysia (Malay peninsula)
Asian tropical islands:
is region includes all Indonesia excluding Papua, Malaysia (Borneo: Sabah and
Sarawak) and Philippines.  ere are 23 species described.
67. adulescens (Pace, 1987: 212) (Drusilla). Distribution: Malaysia (Borneo: Sabah)
68. alboterminalis Pace, 2008: 150. Distribution: Malaysia (Borneo: Sabah)
69. alternans (Cameron, 1925: 45) (Myrmedonia). Distribution: Indonesia (Sumatra)
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 65
70. bryanti Cameron, 1943b: 141. Distribution: Malaysia (Borneo: Sarawak)
71. daiaccorum Pace, 2008: 152. Distribution: Malaysia (Borneo: Sabah)
72. drescheri Cameron, 1939: 17. Distribution: Indonesia (Java)
73. elegantulus Cameron, 1939: 20. Distribution: Indonesia (Java)
74. facundus Last, 1969: 279. Distribution: Indonesia (Java)
= semirufus (Cameron, 1939: 18), preoccupied, not Myrmedonia semirufa Bern-
hauer, 1902
75.  avorufus Cameron, 1939: 18. Distribution: Indonesia (Java)
76. gratellus Cameron, 1939: 20. Distribution: Indonesia (Java)
77. kinabaluensis Pace, 2008: 148. Distribution: Malaysia (Borneo: Sabah)
78. louwerensi Cameron, 1939: 19. Distribution: Indonesia (Java)
79. matangensis Cameron, 1943b: 141. Distribution: Malaysia (Borneo: Sarawak)
80. montanus (Bernhauer, 1915: 152) (Astilbus). Distribution: Malaysia (Borneo:
Sarawak)
81. mortuorum Pace, 1990: 99. Distribution: Philippines (Luzon)
82. nigerrimus Cameron, 1943b: 142. Distribution: Malaysia (Borneo: Sarawak)
83. paederinus Pace, 2008: 153. Distribution: Malaysia (Borneo: Sabah)
84. pallipyga Pace, 2008: 150. Distribution: Malaysia (Borneo: Sabah)
85. preangeranus Cameron, 1939: 17. Distribution: Indonesia (Java)
86. pervariolosus Pace, 2008: 150. Distribution: Malaysia (Borneo: Sabah)
87. punctipennis Cameron, 1939: 18. Distribution: Indonesia (Java)
88. quadriterminalis Pace, 2008: 149. Distribution: Malaysiad (Borneo: Sabah)
89. shiva Pace, 1987: 216. Distribution: Indonesia (Lombok, Bali)
Southern India and Sri Lanka:
is region includes India south from Himalayan region and Sri Lanka.  ere are 9
species included.
90. castaneus (Motschulsky, 1861: 150) (Myrmedonia). Distribution: Sri Lanka
91. hastatus Fauvel, 1904: 64. Distribution: India (Karnataka)
92. hirtus (Kraatz, 1859: 25) (Myrmedonia). Distribution: Sri Lanka, Taiwan
93. indicus Cameron, 1944: 108. Distribution: India (Karnataka)
94. parageminus Pace, 1988: 335. Distribution: Sri Lanka
95. rufescens Cameron, 1939: 534. Distribution: Sri Lanka
96. nilgiriensis Cameron, 1939: 537. Distribution: India (Tamil Nadu)
97. optimus Cameron, 1939: 534. Distribution: India (Tamil Nadu)
98. proximus Cameron, 1939: 538. (Zyras (s.str.), TL: Nilgiri Hills) Distribution:
India (Tamil Nadu)
Peter Hlaváč & Tomáš Jászay / ZooKeys 29: 49–71 (2009)
66
Afrotropical region and temperate South Africa:
is region includes sub-Saharan Africa and the Republic of South Africa. All species
attributed today to the true Zyras were described by Horace Last (1956, 1960, 1962,
1977, 1980) and Roberto Pace (1996). We suspect that these African species are not
true Zyras and belong to other subgenera.
99. basilewskyi Last, 1956: 219. Distribution: Rwanda, Cameroon, Burundi, Gha-
na, Dem. Rep. Congo
100. bonus Last, 1977a: 947. Distribution: Dem. Rep. Congo
101. bramtonus Last, 1962: 222. Distribution: Senegal, Ghana, Dem. Rep. Congo
102. conjectus Last, 1960: 12. Distribution: Gabon, Spanish Guinea, Angola
103.  avipennis Last, 1956: 218. Distribution: Burundi, Ghana, Nigeria, Rwanda,
Dem. Rep. Congo
104. mutarensis Pace, 1996: 231. Distribution: Zimbabwe
105. nakuruensis Pace, 1996: 231. Distribution: Kenya
106. nigritus Last, 1980: 176. Distribution: Rep. South Africa (Cape, Natal)
107. planctos Last, 1977b: 81. Distribution: Ghana
108. punctus Last, 1967: 109. Distribution: Angola, Burkina Faso, Nigeria, Dem.
Rep. Congo
= punctus var. croceus Last, 1967: 110
109. tambachensis Pace, 1996: 228. Distribution: Kenya
Australian region:
Australia plus New Zealand, New Guinea and New Caledonia.  ere are only three
species known from this region.
110. biroi Last, 1980: 161. Distribution: New Guinea
= biroi var. sub avus Last, 1980: 161
111. hirsutus Cameron, 1943a: 353. Distribution: Australia (Victoria)
112. papuanus Pace, 2000b: 159. Distribution: Papua New Guinea
Neotropical region:
Mexico, Central and South America (including Caribbean islands).  ere are two spe-
cies from this vast area known today. We believe that true Zyras do not occur in Neo-
tropics and both species probably belong to other genera of Lomechusini.
113. distinctus Bierig, 1937: 281. Distribution: Cuba
114. paecesanus Pace, 1997: 36. Distribution: Colombia
A revision of the genus Zyras (Zyras) Stephens, 1835. I. Current classi cation status and... 67
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A revision of the genus Zyras (Zyras) Stephens, 1835 (Coleoptera, Staphylinidae, Aleocharinae). I. Current classification status and the redefinition of the genus.pdf
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Supplementary resources

  • ... Vávru (Ostrava). Literárne údaje o Staphylinidae po roku 1993 nachádzame roztrúsené v množstve faunistických príspevkov , alebo katalógov: Ádám (2008), Assing (1998, 1999a, 1999b, 1999c, 2003a, 2003b, 2006b, 2007b, 2008a, 2008b, 2009a, 2009b, 2009c, 2009d, 2009f, 2009g, 2010b, 2010c, 2010d, 2013 ), Assing & Wunderle (2008), Baranová & Jászay (2010), Benedikt (2014), Benedikt et al. (2015), Boháč (2001), Boháč & Matějíček (2002), Blažek & Jászay (2014), Cunev (1994Cunev ( , 1997Cunev ( , 1997aCunev ( , 1998Cunev ( , 1999aCunev ( , 2000Cunev ( , 2001Cunev ( , 2002), Cunev & Jászay (1998), Cunev et al. (1995), Franc (1993a, 1993b, 1995, 2010a, 2010b), Franc & Mlejnek (2000), Hlaváč (2000), Hlaváč & Jászay (2009), Holecová & Zach (1996) , Janák (1993) Jászay (1994, 1996, 1998a, 1998b, 1998c, 1999a, 1999b, 1999c, 2001, 2004), Jászay & Boháč (1994), Jászay & Hlaváč (1997Jászay & Hlaváč ( , 2006Jászay & Hlaváč ( , 2013), Jászay et al. (2000), Jászay & Kodada (1997aJászay & Kodada ( , 1997b), Jászay & Majzlan (2004), Kočárek (2000), Kocian (1996Kocian ( , 1997aKocian ( , 1997b), Kolimár (2000a, 2000b), Korbel (1995), Korbel et al. (1997), Kováč et al. (2007), Krištofík et al. (1993, 1995), Langraf & Schlarmannová (2015), Majzlan (1993Majzlan ( , 1994Majzlan ( , 1996Majzlan ( , 1997Majzlan ( , 1999aMajzlan ( , 1999bMajzlan ( , 1999cMajzlan ( , 2002Majzlan ( , 2003Majzlan ( , 2006Majzlan ( , 2007aMajzlan ( , 2007bMajzlan ( , 2008Majzlan ( , 2010aMajzlan ( , 2010bMajzlan ( , 2010cMajzlan ( , 2010dMajzlan ( , 2011aMajzlan ( , 2011bMajzlan ( , 2012aMajzlan ( , 2012bMajzlan ( , 2012cMajzlan ( , 2013aMajzlan ( , 2013bMajzlan ( , 2013cMajzlan ( , 2013dMajzlan ( , 2014aMajzlan ( , 2014bMajzlan ( , 2015aMajzlan ( , 2015b), Majzlan & Boháč (2012), Majzlan & Cunev (2011), Majzlan & Fedor (2009), Majzlan & Gajdoš (2007), Majzlan & Jászay (1994Majzlan & Jászay ( , 1997), Majzlan O. & Majzlan J. (2011), Majzlan & Kicková (2001) , Majzlan & Rychlík (1992Rychlík ( , 1993Rychlík ( , 1998Rychlík ( , 2002Rychlík ( , 2012 ), Majzlan & Vidlička (2009), Majzlan et al. (2004Majzlan et al. ( , 2010), Majzlan & Zápražný (2005), Mantič (2008, 2012), Mantič & Benedikt (2011), Maruyama (2006), Mock et al. (2009), Olšovský (2007), Papáč (2008), Papáč et al. (2007Papáč et al. ( , 2009 ), Paśnik (2006a, 2006b), Potocký (2015), Rendoš et al. (2012Rendoš et al. ( ), Šustek (1994Rendoš et al. ( , 2000Rendoš et al. ( , 2001Rendoš et al. ( , 2002), Schülke & Makranczy (2012), Šustek & Krištofík (2003Šustek & Krištofík ( , 2009), Šustek & Stanko (2012), Schülke (2003, 2010), Vogel (2006Vogel ( , 2007b). Údaje k podčeľadi Scaphidiinae sú uvedené v prácach Löbl (1997, 1998b) a podčeľadi Dasycerinae v prácach Löbl (1998a) a Löbl & Calame (1996). ...
  • ... A specimen of this genus was reported from Grand Cayman by Blackwelder (1947). True members of the genus Zyras are not known from the West Indies (Hlaváč and Jászay 2009 ...
    Article
    Full-text available
    In 1947, 20 species of Staphylinidae were reported from the Cayman Islands as a result of an Oxford University expedition there in 1938 which made extensive use of a light trap. The list is here expanded to 62 spe­cies based on collections by R. R. Askew, G. E. Ball, E. A. Dilbert, B. K. Dozier, E. J. Gerberg, P. J. Fitzgerald, M. C. Thomas, and R. H. Turnbow since 1970, all of whom also used light traps except for a collection or two by flight intercept trap.
  • ... However, some species are known to be positively associated with Formicidae (i.e. 'myrmecophilous'), such as species in the genus Zyras (Staphylinidae; Akino, 2002; Hlaváč and Jászay, 2009), the most abundant Coleoptera genus in our study. Consequently , this may be the reason why we found significant association between Formicidae and Staphylinidae and total Coleoptera and why the biomass of Staphylinidae and total Coleoptera was lower at regulated rivers. ...
    Article
    River regulation can alter the structural complexity and natural dynamics of river ecosystems substantially with negative consequences for aquatic insects. However, there have been few studies of regulation effects on the export of emergent insects into terrestrial ecosystems. In northern Scandinavia, we compared emerged aquatic insect and terrestrial invertebrate biomass between four strongly regulated and four free-flowing rivers using fortnightly measurements at three upland-forest blocks in each over one summer. The biomass of emerged aquatic insects was significantly lower along regulated rivers than free-flowing rivers. Biomass in Linyphiidae, Opiliones, Staphylinidae, total Coleoptera, Formicidae and total terrestrial invertebrates was also lower along regulated rivers. Aquatic insect biomass did not explain the entire regulation effect on terrestrial invertebrates but did explain significant variations among Linyphiidae, total Coleoptera, Formicidae and total terrestrial biomass. Variations in Formicidae also explained significant variance among several terrestrial taxa, suggesting some keystone role in this group. Overall, our results suggest that river regulation affects upland-forest invertebrate communities, with at least some of these effects arising from links between aquatic emergence and terrestrial predators. The data highlight the need to consider areas beyond the riparian zone when assessing the effects of river regulation. Copyright © 2012 John Wiley & Sons, Ltd.
  • Article
    The taxonomy of Lomechusini Fleming has a complex history. Recent studies have shown that this group is polyphyletic; however, little is known about the evolutionary interrelationships among its constituent genera. The goals of the present study are to infer the phylogenetic relationships of Falagonia Sharp and closely related genera; to define the boundaries of those genera based on synapomorphic characters; and to explore the evolution of myrmecophily within the lineage. The phylogenetic analyses are based exclusively on morphological characters of adults. A total of 36 operational taxonomic units were used for the analysis. The best trees were selected based on maximum parsimony and Bayesian inference. During the parsimony reconstruction, different weighting strategies were used to recover the most robust phylogenetic hypothesis. Although minor differences were observed in the results of the different analyses, the topologies were consistent throughout. Several groups of genera proposed by Seevers (1965), such as the 'Tetradonia' and 'Ecitopora' groups, were not recovered. Thus, these may represent nonmonophyletic groups that were based on nonsynapomorphic diagnostic characters. Our analyses consistently recovered the genera Asheidium Santiago-Jiménez, Delgadoidium Santiago-Jiménez, Falagonia, Newtonidium Santiago-Jiménez, Pseudofalagonia Santiago-Jiménez, Sharpidium Santiago-Jiménez, Tetradonia Wasmann and Thayeridium Santiago-Jiménez, forming a monophyletic group that we have called the 'Asheidium complex'. Falagonia mexicana Sharp shows seven autapomorphies, none of which were used to establish the genus. Based on the phylogenetic results, myrmecophily has evolved independently at least three times within the lineage. This study, based on morphological characters, is one of the first approaches towards gaining an understanding of the phylogenetic relationships within the polyphyletic tribe Lomechusini.
  • Article
    Macrozoque hoplandrioides n. gen. and n. Sp. (Coleoptera: Staphylinidae: Aleocharinae), is described based on speci-mens recently collected from the Chimalapas forest in Mexico. The systematic position of Macrozoque is discussed and is proviosionally placed in the false Lomechusini clade. A distribution map, keys and illustrations are provided.
  • Article
    Full-text available
    All names of species and higher taxa proposed in the tribe Lomechusini (Staphylinidae: Aleocharinae) up to the end of the year 2010 are cataloged, with a full bibliography of original descriptions. The original combination of the described species, original citation of their type localities and their world distributions are given. The following replacement names are proposed for newly discovered primary and secondary junior homonyms: Brachysipalia elgon nom. nov. (= B. elgonensis Pace), Drusilla sculpticollides nom. nov. (=Drusilla sculpticollis Pace), Macrogerodonia bolivianides nom. nov. (=M. boliviana Pace), Orphnebius curticornides nom. nov. (= O. curticornis Pace 2008), Pachorhopala ituri nom. nov. (= P. gerardi Bernhauer 1932), Zyras (Camonia) aethiopicides nom. nov. (=Zyras (Camonia) aethiopicus Pace) and Zyras (Zyras) neoparageminus nom. nov. (= Zyras (Zyras) parageminus Pace). The following secondary homonyms are reinstated as valid: Drusilla opacicollis Cameron (= D. vedda Pace), and Zyras (Grammodonia) erraticus Last (= Z. (G.) adjectus Last). The following new synonymies are proposed (junior synonym first): Aenictobiina Kistner (= Aenictoteratini Kistner), Athexeniina Pace (= Myrmedoniina Thomson), Thoracophagus Kistner (= Drusilla Leach), Urodonia Silvestri (= Pedinopleurus Cameron), Lomechusoides hosodai K. Sawada (= L. amurensis (Bernhauer)), Diplopleurus notabilis Pace (= Trachydonia incredibilis Bernhauer), and Acanthoglossa badia Motschulsky (= Zyras (Glossacantha) affinis Kraatz) (the last-mentioned being a reversal of precedence). The following new combinations are established: Aenictocleptis antennarius (Pace) ex Zyras (Glosacantha); Drusilla congoensis (Kistner) ex Thoracophagus; Pedinopleurus notabilis (Silvestri) ex Urodonia; Pedinopleurus reptans (Pace) and Pedinopleurus tumorfrontalis (Pace ) ex Zyras (Diaulaconia); Pseudastilbus fortispina (Pace) ex Zyras (Glosacantha) and Deroleptus draco (Assing) ex Orphnebius. A lectotype is designated for Lomechusoides ganglbaueri (Bernhauer) to fix this name as a synonym of L. amurensis (Wasmann). The following genera are newly included into Lomechusini, subtribe Myrmedoniina: Aenictophila Seevers, Athexenia Pace, Cephaplakoxena Pace, Malaybergius Kistner, Mimaenictus Kistner & Jacobson, Procantonnetia Kistner & Jacobson, Steysborgia Kistner & Jacobson and Weissflogia Kistner. The genus Myrmigaster Sharp is removed from Lomechusini to Oxypodini. Nomenclatural problems with the names Lomechusa Gravenhorst, Lomechusoides Tottenhamand Atemeles Dillwyn, and recent taxonomic changes to the genus Orphnebius Motschulsky, are also discussed. Lomechusini as currently constituted includes 207 genera and 2205 species or subspecies placed in the subtribes Lomechusina (3 genera, 40 species or subspecies), Termitozyrina (11 genera, 16 species) and Myrmedoniina (193 genera and 2149 species or subspecies). Their regional distribution patterns and extensive associations with ants and termites are briefly summarized.
  • Article
    The Catalogue of Palaearctic Coleoptera provides information about all beetles occurring in Europe, North Africa and Asia north of the tropics. © 2015 by Koninklijke Brill nv, Leiden, The Netherlands. All rights reserved.