Hypoponera punctatissima (Roger) and H. schauinslandi (Emery)
Two morphologically and biologically distinct species
Staatliches Museum fur Naturkunde Gorlitz
The ant known so far under the name ^Hypoponera punctatissima^ consists of two sibling species:
H. punctatissima (Roger, 1 859) and H. schauinslandi (Emery, 1 899), which can be reliably distinguished
by high-precision morphometry and discriminant functions. Phenotypic separation is clearly supported
by a number of biological differences. H. punctatissima is capable of long-range flight-dispersal and of
nesting in several types of outdoor habitats. For Europe north of 48 N, nesting in natural or semi-
natural habitats without endogenous heat production was unknown or extremely rare before 1950
(0 - 6 % of 1 6 overall observations) but documented in 25 % of 24 overall observations after 1950,
which is interpreted as a consequence of global climate change. H. punctatissima does not depend on
passive anthropogenic transport to invade temperate Europe north of 48 N. Most certainly it has
been present in England 1600 years b.p. or much earlier. H. schauinslandi, in contrast, is not capable
of outdoor nesting and long-range flight-dispersal in Europe north of 48 N and occurs there exclusively
in greenhouses, where soil arthropods are available as a main food source. This tropical-cosmopolitan
species depends on passive human transport when introduced to the temperate zone. Both species show
gynomorphic and ergatoid females, major and minor ergatoid males, destructive fighting between
major males but not between minor males, and a similar size of nest populations. Investigation of type
material by high-resolution discriminant analysis established the following synonymies - in any case
with error probabilities of p < 0.006: H. androgyna (Roger, 1859), H. tarda (Charsley, 1877),
H. punctatissima r.jugata (Forel, 1892) and H. punctatissima var. exacta (Santschi, 1923) are junior
synonyms of H. punctatissima, while H. dulcis var. aemula (Santschi, 1911) and H. ergatandria
subsp. bondroiti (Forel, 1911) are junior synonyms of H. schauinslandi. Four names H. abeillei
(Andre, 1881), H. indifferent (Forel, 1895), H. gibbinota (Forel, 1912) and//, lesnei (Bondroit, 1916)
- refer to species clearly different from both H. punctatissima and H. schauinslandi.
Hypoponera punctatissima (Roger) and H. schauinslandi (Emery) - zwei
morphologisch und biologisch trennbare Arten (Hymenoptera: Formicidae) Die bisher
unter dem Namen ^Hypoponera punctatissima bekannte Arneise besteht aus den
Zwillingsarten H. punctatissima (Roger, 1859) and H. schauinslandi (Emery, 1899),
die mittels hochauflosender Diskrimianzanalyse sicher unterscheidbar sind. Die phano-
typische Unterscheidung wird durch eine Reihe biologischer Unterschiede bestatigt.
62 Bernhard Seifert
H. punctatissima ist zu einem weitreichenden Ausbreitungsflug und zur ganzjahrigen
Etablierung von Nestern in verschiedenen Typen von Freilandlebensraumen befahigt.
Nestgriindungen in Freilandhabitaten, die keine endogene Warmeproduktion aufweisen,
waren vor 1950 in Europa nordlich 48 N unbekannt oder extrem selten (hochstens einer
von 16 Nachweisen), warden hier aber fur sechs von 24 Nachweisen nach 1950 dokumen-
tiert. Diese Zunahme wird als Folge der globalen Erwarmung, insbesondere der milderen
Winter interpretiert. H. punctatissima war bei der Besiedlung der temperaten Zone nordlich
48 N nicht von passivem anthropogenen Transport abhangig und konnte schon im
Atlantikum hier eingewandert sein. Zumindest England wurde sehr wahrscheinlich
spatestens im 4. Jahrhundert besiedelt. H. schauinslandi ist dagegen nicht zu einem
weitreichendem Ausbreitungsflug oder zur Etablierung von Freilandnestern in Europa
nordlich 48 N befahigt und tritt hier ausschliesslich in Warmhausern auf, wo kleine Boden-
arthropoden als Hauptnahrungsquelle verfugbar sind. Sie ist ein tropischer Kosmopolit,
dessen Einfilhrung in die temperate Zone unter gegenwartigen klimatischen Bedingungen
von passivem anthropogenen Transport zwingend abhangig ist. Beide Arten zeigen
gynomorphe und ergatoide Weibchen, einen Grossenbimorphismus der ergatoiden Mann-
chen, Beschadigungskampf zwischen den Major-Mannchen, Fehlen eines solchen
zwischen Minor-Mannchen und eine vergleichbare Demographic der Nester. Die
Untersuchung von Typenmaterial mittels hochauflosender Diskriminanzanalyse konnte
folgende Synonymien mit einer Irrtumswahrscheinlichkeit p < 0.006 zeigen: H. androgyna
(Roger, 1859), H. tarda (Charsley, 1877), H. punctatissima r. jugata (Forel, 1892) und
H. punctatissima var. exacta (Santschi, 1 923) sind jiingere Synonyme von H. punctatissima,
wahrend H. dulcis var. aemula (Santschi, 1911) and H. ergatandria subsp. bondroiti
(Forel, 1911) jiingere Synonyme von H. schauinslandi sind. Vier Namen - H. abeillei
(Andre, 1881), H. indifferens (Forel, 1895), H. gibbinota (Forel, 1912) und H. lesnei
(Bondroit, 1916) - beziehen sich auf vier deutlich von H. punctatissima und H. schauins-
landi verschiedene Arten.
The ant genus Hypoponera (Santschi, 1938) is of mainly tropical distribution and differs
from Ponera Latreille, 1804 in the female castes by the absence of a fenestra or thin
translucent spot on anteroventral subpetiolar lobe. According to the conceptions followed
so far (KUTTER 1977, COLLINGWOOD 1979, SEIFERT 1996), only two species of Hypoponera
have invaded Central Europe: H. eduardi (Forel, 1894) and H. punctatissima (Roger,
1859). H. eduardi is widely distributed in the Mediterranean but only exceptionally
found in S Central Europe. //. punctatissima^, in contrast, is known from all countries in
Central Europe and has extended its range to Fennoscandia. Within the myrmecofauna of
these regions, it is outstanding for its peculiarity of nesting in heaps or mounds of various
decomposing organic materials (DONISTHORPE 1915, STRADLING 1965, SEIFERT 1982, DBSSART
& CAMMAERTS 1995, DEKONINCK & VANKERK.HOVEN 2001) or for being recorded deep in a
coal mine in S England (YARROW 1 967).
Hypoponera punctatissima and H. schauinslandi - two distinct species 63
A recent morphological investigation had the surprising result that //. punctatissima
in the conception of all contemporary authors consists of two clearly different species
and that the natural history of these ants must be written anew. An improved morphometric
methodology very near to that described by Seifert (2002) enabled a certain discrimination
of both workers and gynes of the two sister species on the individual level. Despite this
clear separation, a diagnostic character perceptible by simple visual inspection (such as
characteristic setae, carinae, curvatures, dents etc.) was not discovered. Hence, the
distinction of H. punctatissima from its sister species H. schauinslandi (Emery, 1899) is
purely morphometric at the present stage. The results of these studies, of type
investigations in 12 related taxa, and a differential biology of the two sister species are
presented in this paper.
2. Materials and methods
2.1 . Equipment, measuring, and data evaluation
Workers were evaluated for a minimum of 10 and gynes for a minimum of 9 numeric
characters, at magnifications of 200 -320 x using a Wild M10 high-performance stereo-
microscope equipped with a 1.6x planapochromatic objective. All measurements were
made on mounted and dried specimens using a pin-holding stage, permitting endless
rotations around X, Y, and Z axes. A LEICA cross-scaled ocular micrometer with
120 graduation marks ranging over 65 % of the visual field was used. A cross-scale is
inevitable for exact measurements of characters such as PrOc or FoDG. A mean measuring
error of 0.6 um was calculated for small and well-defined structures such as petiole
width, but one of 2 um for larger structures with difficult positioning and some definition
error such as mesosoma length. To avoid rounding errors, all measurements were recorded
in urn even for characters for which a precision of 1 um is impossible. In order to reduce
irritating reflections of the cuticular surfaces, a plastic diffuser was positioned as close as
possible to the specimen. This method considerably improved the resolution of micro-
sculpture and the measuring accuracy for tiny structures such as pubescence hairs. Any
measurement refers to real cuticular surface and not to the diffuse pubescence surface
(important in CW, PEW, FR, MW).
The process to discriminate the most similar sister species included (a) the reduction of
variance in primary data (SEIFERT 2002), (b) the removal of allometric variance by species-
specific functions valid for species pairs (SEIFERT 2002), and (c) a canonical discriminant
analysis with an SPSS program.
2.2. The morphometric characters
CL - maximum cephalic length in median line; the head must be carefully tilted to
the position with the true maximum. Excavations of occiput reduce CL. Anterior
reference point in Ponera and Hypoponera is the upper clypeal protrusion
(do not use the clypeal protrusion below mandibular level!).
CS - cephalic size; the arithmetic mean of CL and CW, used as a less variable
indicator of body size.
CW - maximum cephalic width.
EyePos - a distance between a posterior and anterior reference point measured when
head adjusted as given for CL. Posterior reference point: intersection point
of the median line of head with an orthogonal line directed to eye centre.
Because of bilateral head asymmetries use an average of both eyes. Anterior
reference point: anteriormost point of clypeus as given for CL.
FoDG - mean distance of foveolae on dorsum of 1st gaster tergite. Count the number
of foveolae n in an area A. FoDG is then sqrt A. /sqrt n. To enable most effective
counting, select a mirroring part and align the counting areas longitudinally.
Explicitly, counting was performed within squares of 1 x 10 graduation marks
(GRM) at a magnification of 320 x. In this case, 10 GRM corresponded to
42.5 |im. For example, if 40 foveolae in seven 10 x 10 GRM squares were
counted, then FoDG is calculated as 42.5 * sqrt (7/40) = sqrt (7 * 42.5 * 42.5)7
sqrt (40). In the LE1CA cross-scaled ocular micrometer the space between the
GRM and the cross line is exactly 10 GRM wide. Hence, various numbers of
10*10 GRM squares, connected or not, can be easily defined. In Hypoponera,
pubescence may be patchily removed to reduce the counting error.
FL - maximum divergence of frontal lobes.
FR - minimum distance between frontal carinae.
ML mesosoma length measured in lateral view. Anterior reference point in the
worker: the point where anterior pronotal slope curves in anterior pronotal
extension(shield). Anterior reference point in the gyne: the anteriormost
point of the face of pronotal slope. Posterior reference point in both castes:
caudalmost point of median propodeum. (Note: after the first suture at
propodeal end there is still a sclerite sometimes partially hidden by petiole. If
this sclerite is not fully visible, measure to the suture and multiply with 1 .03).
MW - maximum overall mesosoma width (workers) or maximum width before the
NOH petiole node height; measured in a right angle from a reference line beginning
at the transition point between caudal node profile and caudal petiolar neck
and ending at the most frontodorsal point of node corner (Fig. 1).
PEL petiole length; horizontal distance from the tip of the frontolateral node corner
to the caudalmost point of petiole (Fig. 1).
PEW maximum width of petiole.
PrOc - a preocular distance between a posterior and anterior reference point measured
when head adjusted as given for CL. Posterior reference point: intersection
point of the median line of head with an orthogonal line directed to anterior
eye margin. Because of bilateral head asymmetries use an average of both
eyes. Anterior reference point: anteriormost point of clypeus as given for CL.
Hypoponera punctatissima and H. schauinslandi - two distinct species
Fig. 1 Measuring of petiole length (PEL) and of petiolar node height (NOH) in Ponera and
2.3. Sources of material
Institutions or private collections from which material was studied have the following
coll. Burger collection of F. Burger/Weimar
coll. Dauber collection of J. Dauber/Giessen
coll.Yamauchi collection of K. Yamauchi/Gifu
DEI Deutsches Entomologisches Institut Eberswalde
MCSN Genova Museo Civico di Storia Naturale Genova
MHN Geneve Museum d'Histoire Naturelle Geneve
MNHN Paris Museum National d'Histoire Naturelle Paris
NM Leipzig Naturkundemuseum Leipzig
NHM Basel Naturhistorisches Museum Basel
NHM Wien Naturhistorisches Museum Wien
SM Frankfurt Senckenberg-Museum Frankfurt
SMN Gorlitz Staatliches Museum fur Naturkunde Gorlitz
UM Oxford University Museum Oxford: Hope Entomological Collections
ZMHU Berlin Zoologisches Museum der Humboldt-Universitat Berlin
66 Bernhard Seifert
2.4. Investigated material of H. punctatissima and H. schauinslandi
45 samples of H. punctatissima with 1 1 ergatoid males, 38 gynomorphic and 48 workers
ergatoid female specimens were morphometrically investigated:
AUSTRIA: Dornbach, 1862; Linz-27 km ESE; Gutau, 1974.07.30; Purgstall (15.07.53 E,
48.03.05 N), 1978.07.28. BELGIUM: Bruxelles, Quartier Leopold, railway station, 1994.08.16;
Leuven, centre ofthe city, 1998.07.10; Limburg, Overbroek, Gelinden, 1998.08.16. CANADA:
Montreal, Insectarium, 2003.05.28. ENGLAND: Oxford, St.Giles Road West, greenhouse,
1877, type//, tarda; Westerham 1921.10.29; Westerham, 1921.06.05. FINLAND: Jyvaskyla,
pre 1925 (leg. Sahlberg). FRANCE: Reims, 1934.06.24; Reims, 1934.07.18. GERMANY: Berlin,
pre 1860, type H. punctatissima (Roger); Leipzig-City, 1912.07.15; Miltitz near Leipzig,
1912.06.01; Fahrnau, Hiihnerberg, pre 1925; Berlin, pre 1945; Marburg/Lahn, Niederweimar,
1949.07.04; Halle, Passendorfer Wiesen, 1953.07.22; Dresden, Dresdener Heide,
Saugartenmoor, vii. 1975; Untergrochlitz near Greiz, 1 977.01 .14; Untergrochlitz near Greiz,
1977.07.05; Leipzig-City, 1978.08.; Doberschiltz near Eilenburg, 1981.09.27; Doberschiltz
near Eilenburg, 1981 . 10.04; Moschwig near Bad Schmiedeberg, 1 982.08. 15; Leipzig, Auwald,
1983. 08. 08.; Leipzig, Auwald, 1983.08.12; Ruhla, 1987.06.28; Rudingshain-E, Niddahange,
1990.07.23; Mainz-Finthen-W, Hollenberg, chalky sand dune, 1990.09./10.; Mainz-Finthen-
W, Hollenberg, young orchard, 1990.09./10.; Gotha, Seeberge, Heilige Lehne, 1993.08.14;
Konigshain near Gorlitz, Hochstein-0.5 km SE, 1994.08.09; Rottewitz near Meissen, 2001 ;
Hamburg-Eimsbuttel, 2002.06.26. LUXEMBURG: Luxemburg-City, City Bath 1918.08.
MADAGASCAR: Imerina, pre 1893, type H. jugata. MOROCCO: Tanger, 1904; Volubilis,
pre 1921; Rabat, pre 1921. THE NETHERLANDS: Mijdrecht, 1926.10.08 (van der Wiel).
NORWAY: Roa near Oslo, pre 1945 (Eidmann). POLAND: Rauden, pre 1860, type
H. punctatissima; TUNISIA: Al Hammamat (36.24 N, 10.37 E), 1917.08., type H. exacta.
29 samples of H. schauinslandi with 2 ergatoid males, 24 gynomorphic and 45 ergatoid
female specimens were morphometrically investigated:
BELGIUM: Bruxelles, Botanical Garden, greenhouse, 1909.11., type H. bondroiti;
Maarkedal (50.47N, 3.37E), home for the aged, 1997.01. ENGLAND: London, butcher's
shop, from electrocutor trap, 2000; Berks, Wilden, butterfly house, 2000.02.25. GERMANY:
Potsdam-1.5 km WNW, Sanssouci, greenhouses, 1939.07.19; Potsdam: Babelsberg,
greenhouse Orchidflora, 1939.07.21; Potsdam-1.5 km WNW, Sanssouci, orchid houses,
1940.10.08; Potsdam-1.5 km WNW, Sanssouci, orchid houses, 1940.06.19; Berlin,
Zoological Garden (aquarium), pre 1945; Berlin, Zoological Garden, pre 1945 (Heinroth);
Leipzig, Botanical Garden, greenhouse, 1955.03.11 ; Wuppertal, Zoological garden, 1960.04;
Frankfurt/M., Senckenberg-Anlage, at window, 1973.07.13; Frankfurt/M., Senckenberg-
Anlage, at window, 1974.07.10; Leipzig, Botanical Garden, greenhouse, 1978.08; Berlin-
Neukolln, Britzer Garten, butterfly house, 1995.01.12; Erfurt, butterfly house, 1999.02.27;
Diisseldorf-City, Aquazoo, greenhouse, 2002; Gorlitz-City, Naturkundemuseum,
greenhouse, 2002.03.21; Berlin-Friedrichsfelde, Tierpark, greenhouse, 2002.10.02; Konstanz,
botanical garden of university, greenhouse, 2003.04.24. HAWAII: Laysan Island (25.46
N,171.44 W): Schauinsland, pre 1900, type//, schauinslandi. INDONESIA: Java: Cibodas,
2000.01.05. JAPAN: Okinawa: Yona, 1999.10.03; Okinawa: Miyakelsl., 2001.07.26. POLAND:
Breslau, Botanical Garden, greenhouse pre 1945. SWITZERLAND: Zurich, Zoological
Hypoponera punctatissima and H. schauinslandi - two distinct species 67
Garden, greenhouse, 2001.01.28. TANZANIA: Kilimanjaro: Kiboscho, 1400 m, 1904, type
H. aemula. Victoria: Nyanza, Acip. di Sesse, Bugala, 1908.
2.5. Investigated type material of Hypoponera
For any of the 1 2 names treated here type material was available and any of the names
could be referred to Hypoponera. The text on the type labels is cited with each writing
error, abbreviation or sign and illegible syllables are mentioned. A backslash between the
citations means a change to the next label on the same pin. Differences between text of
labels and statements in original descriptions were in no case as strong that genuine type
identity might be doubted. Material is listed here under the original names in chronological
Ponera punctatissima Roger, 1859: 1 paralectotype worker (des. by Seifert 2003) in
MNHN Paris, apparently labelled by Roger himself Roger \ Ponera punctatissima. n. sp. ?
Rauden Berlin \ MUSEUM PARIS COLL. O. SICHEL I867, and a copy of a handwritten
comment of Roger in German. 1 paralectotype worker in DEI, labelled not in Rogers
handwriting Rauden. 1 paralectotype worker in ZMHU Berlin, labelled not in Rogers
handwriting Schlesien Rauden Roger, S. \ Ponera punctatissima Rog. \ Type.
Ponera androgyna Roger, 1859: 2 ergatoid syntype males in ZMHU Berlin, labelled
not in Rogers handwriting Schlesien Rauden Roger, S. \ Ponera punctat. Rog. v. andro-
gyna Rog \ Type.
Ponera tarda Charsley, 1877: 2 syntype workers in UM Oxford with the following
labels: Ex Coll. J. O. Westwood. Pres. 1857 by Rev. F. W. Hope. Named 1896 by Edw.
Sounders as Ponera punctatissima^ (printed and handwritten), ^burrowing in loose
earth in greenhouse R. S. Charsley OXFORD^ (in Charsley's handwriting), Ponera tarda
Charsley. In greenhouse near the Woodstock R Oxford - An P. punctatissima? sec.
W. J. White + F. Smith in litt. Sept. 1879 (in Westwoods handwriting), and TYPE HYME
2500 172; 2/2 Ponera tarda Charsley, 1877 SYNTYPES ?? HOPE ENT COLL., OUMNH
Ponera abeillei Andre, 1881: 2 syntypes in MNHN Paris: 1 worker type labelled Ajaccio
Abeille \ MUSEUM PARIS COLLECTION ERNEST ANDRE 1914 \ TYPE \ H. abeillei
(Andre); 1 type worker with same labelling but bottom label abeillei (Andre).
Ponera punctatissima r. jugata Forel, 1892: 1 type gyne without scapes, labelled
Typus \ P. punctatissima Roger R. jugata Forel Imerina (Sikora) \ r. P. jugata Forel,
Ponera punctatissima r. indifferens Forel, 1895: 4 syntypes in MHN Geneve: 1 gyne
labelled Typus \ P. punctatissima Rog ? r. indifferens Forel Moramanga (Sikora Juli 93)
75 type\ 2 workers Typus \ Pan. punctatissima ? r. indifferens type Moramanga (Sikora
Juli 93) 75 \ 7. P. indifferens Forel; 1 gyne, labelled 50 Mg \ Ponera punctatissima
Roger ? r. indifferens Forel Mockay (possible reading also Mokay or Mackay).
68 Bernhard Seifert
Ponera punctatissima var. schauinslandi Emery, 1899: 2 gyne syntypes on two different
pins in MCSN Genova: the first syntype labelled Ins. Laysan Schauinslandi and ^Ponera
punctatissima var Schauinslandi Emery in Emery's handwriting and a red label
SYNTYPUS Ponera punctatissima var. schauinslandi Emery, 1899', the second syntype
labelled Ins. Laysan Schauinslandi in Emery's handwriting and a with a red label
SYNTYPUS Ponera punctatissima var. schauinslandi Emery, 1899.
Ponera dulcis var. aemula Santschi, 1911: 1 worker lectotype (by present designation)
and 1 worker paralectotype on separate pins, both labelled Type (handwritten),
MUSEUM PARIS AFRIQUE ORIENT. ALL. KILIMANDJARO (ZONE DES CULTURES,
KIBOSCHO (1400m) CH. ALLUAUD 1904 (printed) and Ponera dulcis var. aemula
Sant (handwritten), stored in NHM Basel. Lectotype with CW 532 |im, paralectotype
CW 545 urn.
Ponera ergatandria subsp. bondroiti Forel, 1911: following syntypes labelled with a
printed Typus label, stored in MEN Geneve: 1 worker labelled Jardin Botan. Bruxelles
15-11-09 \ 6 ergatoide minor in Bondroit's handwriting and /*. ergatandria Forel r.
Bondroiti Ford type Bruxelles Serre chaud in Forels scribbling. This specimen shows
a well-developed stinger and no trace of male genitalia. 3 workers labelled Bruxelles
11-09 in Bondroit's handwriting and P. ergatandria Forel r. Bondroiti For 9 type Serre
chaud Bruxell in Forels scribbling. 1 alate gyne labelled ^Bruxelles Jardin Botan. 5-11-
09 in Bondroit's handwriting and P. ergatandria Forel r. Bondroiti For 9 type
Bruxelles Serre chaud \ r. P. Bondroiti Forel in Forels scribbling. 1 ergatoid male labelled
Jardin Botan. Bruxelles 15-11-09 \ 6 ergatoide in Bondroit's handwriting.
Ponera opaciceps var. gibbinota Forel, 1912: 1 type worker, labelled Typus \ Ponera
opaciceps Mayr 9 var. gibbinota Jery & Kew \ v. gibbinota Forel, (other possible
readings of Forel's scribbling Jery: Kery, Hery, Guy), MHN Geneve.
Ponera lesnei Bondroit, 1916: 1 worker type, labelled 772.2 \ Bois.de. Colombes 7
Juillet 87. Sous les pierres \ MUSEUM PARIS FRANCE P.LESNE 19 \ TYPE \ Ponera
lesnei Bondr. Type and in Forel's handwriting Ponera ragusai Em. ssp. santschii Em
(= lesnei Bond., MNHN Paris.
Ponera punctatissima var. exacta Santschi, 1923: 1 lectotype (by present designation)
and 2 paralectotype workers on the same pin, labelled Tunisie Hammamat (Santschi
VIII) \ P. punctatissima v. exacta Sat, NHM Basel. The Lectotype (top specimen) with
CS 617, SL/CS 0.777 and MW/CS 0.647.
3. Results and discussion
3.1. Synonymic lists of H. punctatissima and H. schauinslandi
Based upon investigation of type material the following synonymic lists are presented
(an argumentation why these synonymies are established is given in section 3.2.).
Hypoponera punctatissima (Roger, 1859)
Ponera punctatissima Roger, 1859; POLAND and GERMANY: Rauden and Berlin
Hypoponera punctatissima and H. schauinslandi - two distinct species 69
Ponera androgyna Roger, 1859; POLAND and GERMANY: Rauden and Berlin
Ponera tarda Charsley, 1877; ENGLAND: Oxford
Ponera punctatissima r. jugata Forel, 1892; MADAGASCAR
Ponera punctatissima var. exacta Santschi, 1923; TUNISIA: Hammamet
Hypoponera schauinslandi (Emery, 1899), stat. nov.
Ponera punctatissima var. schauinslandi Emery, 1899; HAWAII: Laysan Island
Ponera dulcis var. aemula Santschi, 1911; TANZANIA: Kilimandjaro; syn. nov.
Ponera ergatandria subs, bondroiti Forel, 1911; BELGIUM: Bruxelles; syn. nov.
Hypoponera bondroiti, sensu YAMAUCHI et. al 1 996 [morphometric data and investigation
of syntopic material]
3.2. Differential diagnosis of H. punctatissima and schauinslandi
The types of all available names possibly representing synonyms of H. punctatissima
and H. schauinslandi were investigated in this study to ensure that no senior synonym of
H. schauinslandi remained undetected. The identity of 12 taxa was checked in this way.
Four names - H. abeillei (Andre, 1881), H. indifferens (Forel, 1895), H. gibbinota (Forel,
1912) and H. lesnei (Bondroit, 1916) - refer to species clearly different from both
H. punctatissima and H. schauinslandi (see section 3.3.).
Primary morphometric data of workers and ergatoid females specimens of 5 European
species of Hypoponera and of the gynes of H. punctatissima and H. schauinslandi are
given in Tabs 1 and 2. No conspicuous external characters such as carinae, dents,
curvatures, sculptural elements, setae etc. were discovered that could allow a discrimination
of the sister species H. punctatissima and H. schauinslandi by subjective visual
inspection. Reliably possible was the separation of workers by discriminant functions,
the details of which are given in the Appendix (section 4). The resulting discriminant
scores D(10) given as arithmetic mean standard deviation [lower extreme, upper extreme]
H. punctatissima D(10) 3.068 1.004 [+ 0.61, + 4.84] n = 48
H. schauinslandi D(10) -2.8150.995 [-4.91, - 0.61] n = 45
1 00 % of the 93 initial classifications were confirmed by D(l 0) with error probabilities of
p < 0.05 in the worst case and any of the involved type individuals was allocated to either
cluster with an error probability p < 0.006 (Fig. 2). Hence the synonymies stated above are
D(10) was 3.02, 3.21, 2.83 in the lectotype and two paratypes of H. punctatissima from
Rauden, 2.37 and 0.77 in the syntypes of//, tarda from Oxford, 3.80, 4.17 and 4.29 in the
lectotype and two paratypes of//, exacta from Al Hammamat, -0.99 and - 1.54 in the
syntypes of H. aemula from Kilimandjaro and - 3.21, - 4.28 and - 2.93 in the syntypes of
H. bondroiti from Bruxelles.
A reliable separation of gynes was possible by discriminant functions, the details of
which are given in the Appendix (section 4). The resulting D(9) were:
H. punctatissima D(9)
H. schaitinslandi D(9)
3.294 0.816 [+1.77, + 5.15]n = 37
-4.316 1.234 [-6.38,-1.77]n = 24
Any of the 61 initial classifications was confirmed by D(9) and any specimen including
the types was allocated to either cluster with an error probability p<0.001 (Fig. 3). Hence,
the synonymies stated above are fully confirmed also in the gynes: D(9) was 1.77 the
paratype of H. punctatissima from Rauden, minus 5.48 and minus 4.14 in the syntypes of
H. schauinslandi from the Laysan Island, and minus 5.35 in the syntype of H. bondroiti
(n = 38)
(n = 48)
-6,0 -5,0 -4,0 -3,0 -2,0
1,0 2,0 3,0 4,0 5,0 6,0
Fig. 2 Canonical discriminant values D(10) of workers of H. schauinslandi (Emery, 1899) and
H. punctatissima (Roger, 1859). Position of type specimens is indicated by letters:
B - H. bondroiti (Forel, 1911), A - H. aemula (Santschi, 1911), T - H. tarda (Charsley,
1877), P - H. punctatissima, and E - H. exacta (Santschi, 1923)
The type gyne ofH.jugata from Imerina, in which both scapes are missing, could not be
included in the analysis above. The primary data of this type gyne were CS 677, CL/CW
1.145, MW/CS 0.770, PEW/CS 0.493, PEL/CS 0.385, ML/CS 1 .654, NOH/CS 0.376, FODG
13.5. To objectively assess the position of this taxon, a separate analysis calculating a
discriminant D(8) was run, the details of which are given in the Appendix (section 4).
Hypoponera punctatissima and H. schauinslandi two distinct species
The exclusion of the powerful character SL from the analysis did not decrease the
discriminative performance dramatically: 100% of the initial classifications still were
confirmed, 98.4 % of individuals were classified with error probabilities of p < 0.01, and
any of the involved type individuals was allocated to either cluster with an error probability
p < 0.001. The type of H.jugata showed a D(8) of 1.70 with a probability to belong to
H. punctatissima of p > 0.999. The overall D(8) for 62 gyne specimens were:
H. punctatissima D(8) 2.481 0.816 [+ 0.65, + 3.78] n = 38
H. schauinslandi D(8) -3. 124 1.240 [- 5.1 1, - 0.65] n = 24
Comparative statements on the character combination of ergatoid males are not possible
here because only one specimen plus one head of one H. schauinslandi male was available.
However, with CS given in mm, the size-corrected scape length index
SL/CS cor = SL/CS /(- 0.3589 CS + 0.8741)
of these two individuals was clearly lower than in all 1 1 H. punctatissima males: 0.857
and 0.885 against 1.000 0.01 8 [0.968, 1.028]. These data promise a clear separation of
both sister species when more specimens with complete character sets would be available.
H. punctatissima j
(n = 35)
(n = 23)
-8,0 -7,0 -6,0 -5,0 -4,0 -3,0 -2,0 -1,0 0,0
Fig. 3 Canonical discriminant values D (9) of gynes of H. schauinslandi (Emery, 1899) and
H. punctatissima (Roger, 1859). Position of type specimens is indicated by letters:
B - H. bondroiti (Forel, 1911), S - H. schauinslandi, and P - H. punctatissima
Tab. 1 Morphometry of worker and ergatoid female individuals of 6 species of Hypoponera occurring
in Europe given as arithmetic mean standard deviation [minimum, maximum]. Characters in
heavy type are most diagnostic
(n = 48)
(n = 45)
(n = 2)
0.840 0.0 14
0.1 48 0.008
O.I 22 0.008
0.1 19 0.007
11. 69 0.67
11. 70 0.70
1.469 0.0 18
0.683 0.01 6
0.402 0.0 15
3.3. Taxa belonging neither to H. punctatissima nor to H. schauinslandi
Hypoponera abeillei (Andre, 1881)
The two worker syntypes from Ajaccio strikingly differ from both H. punctatissima and
H. schauinslandi in many characters - e.g. much larger CL/CW, SL/CS, and PEL/CS or
much smaller FR/CS (Tab. 1). The elongated head and petiole is outstanding among
European Hypoponera. The specimens show no visible ommatidiae and a very small
subpetiolar corner but no suggestion of a fenestra or translucent spot. This is clearly a
Hypoponera indifferens (Forel, 1895)
This taxon from Madagascar originally had been described as a race of H. punctatissima
Hypoponera punctatissima and H. schauinslandi two distinct species
Tab. 2 Morphometry of Hypoponera gynes given as arithmetic mean standard deviation [minimum,
maximum]. Characters in heavy type are diagnostic. Strings of asterisks are a visual indication
of characters useful in separating H. punctatissima and H. schauinslandi
(n = 38)
(n = 24)
(n = 2)
0.758 0.0 15
0.727 0.0 10
0.753 0.01 8
2. 158 0.073
2. 197 0.064
2. 120 0.036
but BOLTON (1995) formally raised it to species rank. Morphometric results given here
clearly confirm this view.
The two syntype gynes from Moramanga and Mackay show the following standard
measurements: CS 634, 646; CL/CW 1.198, 1.224; SL/CS 0.788, 0.824; MW/CS 0.752, 0.763;
PEW/CS 0.563, 0.640; PEL/CS 0.420, 0.394; ML/CS 1.728, 1.746; NOH/CS 0.451, 0.516;
FODG 12.3, 10.6; PrOc/CL 0.278, 0.280; FL/CS 0.229 (width of rupture subtracted), 0.239;
FR/CS 0.097, 0.116. There is a median rupture of clypeus and anterior vertex in the
Moramamga specimen. A comparison with the data presented in Tab. 2 shows that CL/CW
is completely above the known range in H. punctatissima and H. schauinslandi and that
4 other characters are significantly different. The most striking difference, however, is the
much larger PrOc/CL of H. indifferens, which ranges between 0.147 and 0.168 in
H. punctatissima and H. schauinslandi.
74 Bernhard Seifert
Characters of the two syntype workers from Moramanga correlate with those of the
gynes: CS 570, 606; CL/CW 1 .242, 1 .232; SL/CS 0.772, 0.760; MW/CS 0.691 , 0.703; PEW/
CS 0.521, 0.533; PEL/CS 0.403, 0.401 ;ML/CS 1.522, 1.544; NOH/CS 0.416, 0.424; FODGn.m,
15.8; EyePos/CL 0.379, 0.379; FR/CS 0.093, 0.083. A comparison with the data presented in
Tab. 1 shows that CL/CW, PEL/CS and NOH/CS are clearly above and FR/CS below the
known range in H. punctatissima and H. schauinslandi. Again the most striking difference
is the eye position: EyePos/CL ranges between 0.251 and 0.284 in H. punctatissima and
H. schauinslandi. Vertex foveolae are wider and deeper in H. indifferens and its mandibles
10-dentate and rather homodont.
Hypoponera gibbinota (Forel, 1912)
It remains unclear from which region of the world this ant had been introduced to a
greenhouse in S England. The type worker of H. gibbinota is very distant from members
of the H. punctatissima group and differs in having the anteromedian clypeal margin
clearly notched, a much stronger microsculpture, a well-developed microreticulum on the
gaster tergites (obscuring foveolae!), a much longer scape, a much lower PEL/NOH and a
clearly larger EyePos/CL. By morphometry, the most similar European species is H. eduardi
but H. gibbinota differs by notched clypeus, longer scape and stronger microreticulum
on gaster tergites. Data of the H. gibbinota type worker: CS 678, CL/CW 1 .200, SL/CS
0.884, MW/CS 0.664, PEW/CS 0.500, PEL/CS 0.337, ML/CS 1.427, NOH/CS 0.405, FODG
immeasurable, EyePos/CL 0.246, FR/CS 0.157, FL/CS 0.259, PEL/NOH 0.832.
Hypoponera lesnei (Bondroit, 1916)
H. lesnei, described from Seine-et-Oise: Bois de Colombes near Paris, has been listed by
BERNARD (1967) as a synonym of H. ragusai (Emery 1894) that was described from Sicily.
I cannot comment on Bernard's unexplained speculation because types of the latter taxon
were not seen. For the problems considered here, it is important to state that the H. lesnei
type worker definitely belongs to Hypoponera and that it is clearly heterospecific from
both H. punctatissima and H. schauinslandi or any other species described in this paper.
There is no species within this study combining a large CL/CW, FR/CS, and PEL/NOH
with small SL/CS. The character combination of the type worker is: CS 550, CL/CW 1.283,
SL/CS 0.728, MW/CS 0.642, PEW/CS 0.45 1, PEL/CS 0.387, ML/CS 1.518, NOH/CS 0.3 19,
FODG 13.0, FR/CS 0.139, PEL/NOH 1 .213, 4 ommatidiae per eye.
3.4. Habitat selection of H. punctatissima and H. schauinslandi
Significant differences in the nesting sites of Hypoponera schauinslandi and Hypoponera
punctatissima are exposed when the consideration is restricted to localities in Europe
north of 48 northern latitude.
H. schauinslandi was exclusively found in greenhouses or other heated buildings with
air temperatures constantly above 19 C (usually between 22 and 27 C). No nest sites
outside houses are known (Tab. 3). 94 % of the H. schauinslandi sites were greenhouses
with tropical or subtropical plants situated in zoological and botanical gardens, in butterfly
parks, plant stores, museums etc. The only exception, a home for old aged, most probably
had a room with a lot of potted plants. This type of habitat selection indicates H. schauins-
Hypoponera punctatissima and H. schauimlandi - two distinct species 75
landi to be a true tropical species, the anthropogenic introduction of which into the north
temperate zone began no earlier than in the 1 860s, when tropical plants and animals were
imported in higher numbers and could be kept in greenhouses with stable heating conditions
throughout the year.
In H. punctatissima, only 22 % of nests were found in heated houses but 78 % under
open-air conditions. Among these outdoor nests, 62 % were found in heaps of decom-
posing organic material such as saw dust, hay, flotsam deposited on the sea shore, or
manure from cow or horse stables. Many of these heaps, especially when consisting of
wet and fine-grained material, may show an enormous heat production, which may force
the ants in summer to construct their brood chambers 1 - 2 cm below the surface (Klaus
Lippold pers. comm.) but prevents deeper layers from freezing even in the coldest winters.
SEIFERT (1982) has doubted permanent establishment of nests and successful over-
wintering of H. punctatissima in outdoor habitats without endogenous heat production
(ONHP sites), but this idea is no longer acceptable in the context of rising global tempe-
ratures. In continental Europe north of 48 N, there is an apparent trend to increase nesting
in ONHP habitats during the 20th century. The chronological sequence of six recordings
from such habitats is: a xerothermous meadow in the Passendorfer Wiesen near Halle in
1953, a park lane in the city of Leipzig in 1978, a xerothermous grassland near Mainz-
Finthen in 1 990, a xerothermous orchard near Mainz-Finthen in 1990, a Molinia stand in a
peat bog (!) near Hutschenhausen in 1993, and a xerothermous granite rock near Gorlitz in
1 994. It is noteworthy that there no clear record exists from ONHP habitats, but 1 5 records
from other habitats prior to 1 950, while there are 6 records from ONHP habitats and 1 8 from
other habitats after 1950. This weakly significant trend (X 2 2.72, p < 0.10) is also confirmed
by literature data of H. punctatissima in which the species identity is unclear: only one
record from a garden in England (DONISTHORPE 1927) could possibly be attributed to an
ONHP nest site before 1950.
The clear interspecific nest site differences allow a most probable allocation of the
following literature records on outdoor nesting to H. punctatissima without having seen
voucher specimens: a bone heap in Queenborough/E Kent (DONISTHORPE 1915), a coffee
waste heap in a quarry near Ashbourne/Derbyshire (STRADLING 1965), four horse dung
heaps at Exeter/Devon, Trescott/Staffordshire and Hopwood/Warwickshire (TIMMINGS &
STRADLING 1993), a compost heap in Comberton/Worcesterhire and a pile of wood chippings
in a large coppice wood near Pershore/Worcestershire (WHITEHEAD 1994) and mounds of
mowed grass in Belgium (several cases, DESSART & CAMMAERTS 1 995). Summer temperatures
in the coffee waste heap were 35 C at 25 cm depth and there was a high abundance of
Collembola near to surface. The preferred temperature range of Hypoponera within the
horse dung heaps was 22 32 C.
Confirmation by skilled determiners is urgently needed for some spectacular statements
of RESSL (1995) on nesting in several natural sites in Austria, mainly xerothermous
grasslands, in the years 1956- 1966. Repeated occurrence of workers in winter nests of
small mammals (in particular Talpa), as stated by Ressl, would add a really new trait to the
biology of H. punctatissima. However, the few specimens of Ressl found in the collection
of NHM Vienna turned out as Ponera (Schlick-Steiner und Steiner in litt.) and it remains to
be studied if the many missing specimens possibly stored in the collection of Walter
Faber were also misidentified.
Tab. 3 Nesting sites verified by examined voucher specimens. GH = greenhouses, HHD = heated
human dwellings, ODHP = outdoor habitats with endogenous (decompository) heat production,
ONHP = outdoor habitats with no endogenous heat production
Head capsules from a sewer beneath a heated Roman bath house in Yorkshire, older
than 1500 years b.p. were once determined by Chris O'Toole as Hypoponera punctatissima
(O'Toole, pers. comm. 2003.02.26). The whereabouts of voucher specimens of this
archaeological sample investigated in the 1970s are unknown, but a correct naming is
supported by the circumstances of finding that speak against Ponera, by the most certain
absence of H. schauinslandi from England before 1850, and by a significant probability of
H. punctatissima to occur in Roman England. The large potency for long-range flight-
dispersal (see section 3.5) and the good chance to survive temperate winters indicate that
H. punctatissima should have reached north temperate Europe independent of anthro-
pogenic transport and might have invaded these areas already during warmer climatic
periods beginning with the Atlantic. Only the British Isles with their mild winters should
have provided conditions for early invasion.
3.5. Swarming and dispersal of H. punctatissima and H. schauinslandi
Observations throughout the year of alate gynes of the two sibling species in Europe
north of 48 N significantly differ (Tab. 4). In H. punctatissima, 19 observations were
made between 28 May and 16 August (= 95 %) but only one outside this period
(5 November, referring to an intranidal observation). This pronounced summer peak is
obviously both a consequence of a real production maximum in the many outdoor nests
and of favourable flight temperatures from late June to mid August. In fact 14 out of 18
summer observations refer to catches of alate gynes in dispersal and only 5 to intranidal
collecting. Take-off of alate gynes from a nest in a xerothermous rock area was once
directly observed near the town of Gorlitz: the gynes flew off on 9 August 1994,
18.00 - 18.15 h local time when air temperatures were 22 C (after a daily maximum of
25 C). Very high flight temperatures and much sunshine are obviously not necessary.
The mean and maximum of daily air temperatures were 18. 7 2. 3 [15.0, 21.5] C and 24.1 3.2
[19,28] C during eight flight days with weather recordings available - these means are
only 1.5 2.5 K higher than average seasonal values.
In H. schauinslandi, 8 observations (= 80 %) fall into the period from 9 November to 25
February but only two were made on 10 and 13 July (in both cases referring to an alate
gyne caught at window panes in a building). These data suggest H. schauinslandi alates
mainly to develop in winter and rarely in summer which appears unintelligible in view of
rather constant temperature, and humidity conditions provided in greenhouses throughout
the year. However the data are significant - when weighted against the overall frequency
of greenhouse collecting: within 23 greenhouse collecting days of different investigators
Hypoponera punctatissima and H. schauinslandi - two distinct species
from October to March recorded in the author's files, 8 yielded alate gynes and 15 not
while all 20 collections from April to September yielded no alates (X 2 6.40, p < 0.02).
A possible explanation might be that development of alates is depressed by the long-day
light regime of the summer period.
The mode of dispersal of both sibling species in temperate Europe N of 48 N is most
different. There is no suggestion that H. schauinslandi gynes are capable of long-range
flight-dispersal in climates with July isotherms below 20 C. No alate gynes in dispersal
were caught outside buildings. In two cases they were caught on a window pane inside a
building in Frankfurt City and in another one in an electrocution trap in a butcher's shop
in London City. If outdoor flight-dispersal should ever occur in H. schauinslandi, it should
not end in finding a suitable nesting site. Hence, dispersal of H. schauinslandi undoubtedly
depends upon passive anthropogenic transport to heated houses with living soil.
A zoogeographic characteristics of this species is: tropical, cosmopolitan, tramp species.
In H. punctatissima, as much as 86 % of dispersing alates were caught outside of
buildings and as much as 50 % far away from the next human settlement (Tab. 5). In one
case an alate gyne was caught in a pitfall trap in a small, isolated peat bog in the centre of
a compact 36 km 2 forest. This clearly indicates a high potency for long-range flight-dispersal
in temperate climates. This adaptation and the well-documented ability for permanent
nesting in natural habitats makes it most probable that H. punctatissima was not depending
upon passive anthropogenic introduction into areas north of 48 N - though human
civilisation undoubtedly provided the basis for a considerable increase of its abundance.
A zoogeographic characteristic of this species is: tropical to south temperate, cosmopolitan.
Tab. 4 Occurrence throughout the year of alate gynes verified by examined voucher specimens
Tab. 5 Spatial occurrence of flying gynes verified by examined voucher specimens. inBu = within
human buildings, ouBu = outside human buildings but within human settlements, open = far
away from human settlements
3.6. Other traits of the biology of//, punctatissima and H. schauinslandi
Very little is known about other traits of the biology in H. punctatissima. Absence of
voucher specimens also makes an interpretation of published statements difficult - e.g. it
is not clear to which species the observations of HAMILTON (1979) could refer.
78 Bernhard Seifert
For H. schauinslandi and referring to the population on the Okinawa Islands, YAMAUCHI
et al (1996) provided the following basic data:
(1) nests are weakly polygynous and polydomous colonies occur.
(2) there is a distinct size dimorphism in ergatoid males: CS in minors is 508 - 615 jim
and in majors 695 - 808 Jim.
(3) a differential behaviour exists between males: majors fight with majors - no killings
but damage of appendages was observed in 7 out of 8 cases. Majors do not
attack minors because minors seem to mimic females. Minors do not fight among
themselves. Both minor and major males mate with both gynomorphic and ergatoid
(4) there is a distinct polymorphism of female reproductives: gynomorphic queens
have large eyes, 3 ocellae, a large mesosoma and 3 + 3 ovarioles and are able to
fly. Ergatoid queens are similar to workers but have a slightly larger size, larger
eyes, a spermatheka (that is always inseminated) and 3 + 3 ovarioles. Workers are
always without ovarioles and a spermatheka.
(5) the arithmetic mean, standard deviation and upper and lower extremes were in 26
nest populations: 29.2 34.8 [1,184] for workers, 3.7 4.5 [0,20] for ergatoid queens
(these found in 22 nests), 0.73 1.06 [0,4] for gynomorphic queens (found in 10
nests), 0.38 0.64 [0,2] for major males (found in 8 nests), 0.58 1.24 [0,6] for
minor males (found in 9 nests) and 5.30 9.47 [0,39] for alate gynes (found in 12
In its European greenhouse habitats, H. schauinslandi was repeatedly observed to
nest under very moist conditions in log or under bark. Collembola and other microarthropods
seem to be its most preferred prey, which was also observed for H. punctatissima in a
coffee waste heap. The obvious dependence of both H. schauinslandi and
H. punctatissima upon small epigeal and hypogeal arthropods and difficulties to use
other food sources prevents them from having a wider synanthropic distribution.
The following observations, where voucher specimens are available, indicate a high
similarity of H. punctatissima with H. schauinslandi in a number of basic traits:
(1) at least occasional polygyny/polydomy is suggested by observations of Klaus
Lippold (pers. comm.) who stated 1 found nest by nest closely together under
the whole surface of a large sawdust heap. I could not really decide if it were many
different societies or a single one.
(2) presence of size dimorphism in ergatoid males: CS was 615 - 722 jim in minors
and 762 - 879 Jim in majors. In size and general habitus, the latter resemble
(3) fighting between major males is indicated by traumatological findings: 3 out of 8
morphometrically investigated major males showed healed injuries as completed
peroxidase reactions indicated. In two cases the funiculus was cut off after the
pedicellus, in another case the tibio-femoral junction was damaged. No injuries
were observed in 19 minor males and these obviously do not fight among
themselves as the presence of 17 uninjured minor males within a nest sample from
Doberschutz 1981.09.21 suggests.
Hypoponera punctatissima and H. schauinslandi - two distinct species 79
(4) polymorphism in female reproductives: gynomorphic and ergatoid specimens are
observed. The ergatoids are most similar to workers but have larger CS, ML/CS
and MW/CS. However, the only conspicuous external character to distinguish
ergatoids from workers is the larger eye. The eye size index (arithmetic mean of
maximum and minimum eye diameter divided by CS) is 0.044 - 0.069 in workers
and 0.077 - 0.1 12 in ergatoid females.
(5) As data for worker nest populations are available: 29, 30, 40, 50, 171 and 175.
Production of alate gynes was 25 and 29 in two reported cases.
Singularity of ergatoid males within H. punctatissima nests seems to be rare. For the
nest samples seen by the author and supplemented by statements of the collectors, 2, 2,
2, 3, 3, and 17 ergatoid males were recorded. Major and minor males can be found within
the same nest.
4. Appendix - discriminant functions
Allometric variance was removed in H. punctatissima and H. schauinslandi by the
following functions with CS given in mm:
for workers and ergatoid females:
CL cor = CL/CW /(- 0.32782*CS + 1 .3665)
SL = SL/CS /(- 0.1 1300*CS + 0.8202)
FR cor =FR/CS/(+0.03960*CS +0.0991)
MW cor = MW/CS /(+ 0.29324*CS + 0.4853)
PEW cor = PEW/CS/(+0.51137*CS + 0.1615)
PEL OT = PEL/CS /(+ 0.09268*CS + 0.3 1 56)
NOH cor =NOH/CS/(+ 0.33847*CS + 0.1582)
FoDG cor =FoDG/(-3.835*CS+ 14.68)
ML cor = ML/CS /(+ 0.30388*CS + 1 .3024)
in gynomorphic females:
CL cor =CL/CW/(-0.35182*CS + 1.3615)
SL OT = SL/CS /(- 0.20526*CS + 0.8840)
MW cor = MW/CS /(+ 0.04802*CS + 0.7426)
PEW cor = PEW/CS /(+ 0.08920*CS + 0.4345)
PEL OT = PEL/CS /(- 0.05723*CS + 0.4254)
NOH cor =NOH/CS /(+ 0.04261*CS + 0.3532)
FoDG cor =FoDG/(-2.335*CS + 14.14)
ML cor = ML/CS /(+0. 141 06*CS+ 1.5694)
80 Bernhard Seifert
With these size-corrected values the following canonical discriminants were calculated
for workers and ergatoid females:
D(10) = - 79.04 + 20.740 CS + 4.429 CL cor + 78.305 SL cor + 7.143 PEW cor - 3.301 PEL cor
- 8.530 NOH cor + 1 .088 FR cor + 4.983 ML cor - 6.797 MW cor - 0.03 1 FoDG cor
for gynomorphic females:
D(9)= - 84.44 + 29.133 CS - 3.999 CL cor + 46.127 SL cor + 12.373 PEW cor + 18.1 18PEL cor
+ 8.518NOH cor - 20.687 ML cor + 1.147 MW cor + 0.595 FoDG cor
for gynomorphic females under exclusion of scape length:
D(g)= -51.924 + 37.438 CS + 8.647 CL CO] . + 2.088 PEW cl , + 29.229 PEL cor
+ 13.820NOH cor -33.337ML cor -3.291 MW cor + 1.034 FoDG cor
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Manuscript accepted: 1 7 November 2003
Dr. Bernhard Seifert
Staatliches Museum fur Naturkunde
02806 Gorlitz, Germany
E-Mail : firstname.lastname@example.org. sachsen.de