Ponera testacea EMERY, 1895 stat. n.

Csősz, S. & Seifert, B., 2003, Ponera Testacea Emery, 1895 Stat. N. - A Sister Species Of P. Coarctata (Latreille, 1802) (Hymenoptera, Formicidae), Acta Zoologica Academiae Scientiarum Hungaricae 49 (3), pp. 201-214 : 207-213

Ponera testacea EMERY, 1895 stat. n.

( Figs 4, 6, 8, 10 )

P. coarctata var. testacea EMERY, 1895 Memoria letta alla R. Accademia delle Science dell’ Instituto di Bologna (5) v. 5 p. 294.

Lectotype (worker) by present designation: Bonifacio, leg. REVEL 1872 , deposited inthe “ EMERY collection”, MCSN Genova ( Italy). Its measurements: CL 655; CW 495; FR 65; FL 140; SL 460; ML 840 ; PEH 360; PEW 300. Para-lectotypes (3 workers) deposited in“Forel collection”, Muséum d’Histoire Naturelle (Genéve, Switzerland). One specimen Rapallo / Liguria / Mai 1891 / Flach and two other specimens (at one pin) labelled by EMERY “Gallia merid.” and with a blue label “Cotypus”.

Material studied – Altogether 59 nest samples with 147 workers and 31 gynes were investigated. The localities are givenbelow inalphabetical order.

Austria – Gainfahrn, leg. G. Mayr 19th century; Kirchberg a. Wechsel, leg. Ganglbauer 1901; Klosterneuburg, leg. Hummler appr. 1860; Loich, Steinfeldberg (28 km SW St. Pölten), leg. Faber 1965.06.05; Oberndorf, leg. Rausch 1970.07.31; Pernitz (37 km SW of Vienna), leg. Franz; Bulgaria – Melnik, Valley of the Melnik River, leg. Seifert 1982.08.30 and 1982.09.01; Melnik 2 km N, leg. Seifert 1982.08.31; Melnik: Rozen 8 km N, 1400 m, leg. Seifert 1982.09.01. Corsica – Bonifacio, 1872, leg: Revel 1 lectotype worker. Croatia – Buccari, 1927.04.21.; Gospic, 1918.08. leg. Bíró; Krivorije, Dalmatie, 1927. leg. Fodor; Sucurac, Dalmazia, (No. 55/2) leg. Novák; Zelenika, leg. Gerger; Zengg; Dalmatien, Castelnuovo, leg. Reiter; France – Gallia merid. with a blue label “Cotypus” 2 paralectotype workers. Germany – Bad Kreuznach 4 km SSW: Münster am Stein, leg. Rüschkamp 1929.09.18; Eisleben vic.: Wormsleben, leg. Seifert 1983.05.28; Friedrichsaue near Aschersleben, leg. Seifert 2002.05.13; Herborn, Schloss 2 km SSE, Galgenkopf, 232 m, leg. Wolf 1949.06.09; Jena vic.: NSG Leutratal, Mesobrometum, coll. Dunger 1974; Jena vic.: NSG Leutratal, Arrhenateretum, coll. Dunger 1974; Kaiserstuhl: Oberbergen 2 km N, leg. Lamparsky 1990.08; Kyffhäuser: Steinthaleben 1 km SSE, leg. Seifert 1987.05.25; Kyffhäuser: Bad Frankenhausen 1 km NW leg. Seifert 1987.05.26; Mainz-Gonsenheim, leg. Dauber 1994.07.; Marburg /Lahn, Weimar’scher Kopf, leg. Wolf 1949.04.29; Marburg /Lahn, leg. Wolf 1951.09.08; Meissen vic.: Niedermuschütz 2 km NE, leg. Seifert 1982.07.29; Münster a. S. (hill top under stone), 1929.09.18. leg. F. Rüschkamp; Sangerhausen vic.: Ichstedt–NW, leg. Weipert 1998.07.08. Dillenburg, leg. Wolf 1946.08.10. Hungary – Bp. Sashegy, (Cribnope), 1886.03.28.; Budapest, 1914. leg. Horváth; Farkasd, 1901.10.08. (No. 718/280); Füle, 1933.05.16. Úhelyi; Pálmatér, Com. Békés; Hu. Com. Bors. Nagysalló, 1923.08.24. leg. Dudich; Keő, 1881.09.09. (No. 1.) leg. André 1885; Kun. szt. Miklós, 1928.08. Újhelyi; Pálmakert, 1912.09.19. leg. Horváth; Pusztapoó, 1929.02.11. leg. Vásárhelyi; Pusztapoó, 1929.02.11. leg. Vásárhelyi; Rimaszombat 1909.07.10. leg. Szabó; Sátoraljaújlak, 1882.10.03. (No. 718/275); Szigetszentmiklós, 1909.05.15. leg. Bíró; Szigetszentmiklós, 1912.10.12. leg. Bíró; Szigetszentmiklós, 1912.10.16. leg. Bíró; Szigetszentmiklós, 1912.10.19. leg. Bíró; Szöd, 1920.05.15. leg. Bíró; Újhely (No. 718/276); Valkó, 1903.09.13. leg. Bíró; Veszprém, 1881.09.20. (No.718/281); Villány, leg. Sieber 1982.05; Italy – Rapallo (Liguria) V. 1891, leg Flach; Portomzurizio, 1914.11. leg. A. Andreini. Romania – Déva, leg. Mallász; Hermannstadt vicinity, 1948.05.20. leg. Dr. Worell; Kolozsvár, 1882.10.18. (No. 718/277). Serbia – Bacaka. Spain – Montnegre, Barcelona, 1998.09.17. leg. Espadaler. Switzerland – Pfynwald, Sion 1 km S leg. Seifert 1990.07.14.; Veyrier GE, 11.12.

Worker ( Figs 4, 6, 8 , Table 1 ) – Average body length 2.5–3.0 mm. Colour light ochraceous yellow to brownish yellow. Whole body covered with thick decumbent hairs. Mandibles triangular, with 3 apical teeth at the tip of the masticatory border, followed by a regular series of 9–14 minute denticles. Clypeus produced anteriorly, with a rather distinctly raised longitudinal carina. Eyes very small, or invisible, with 0–3 minute facetes. Scape with subdecumbent and suberect hairs. Tibiae frequently with short, decumbent hairs only. Alitrunk relatively lower than in coarctata . Furrow between the mesonotum and anepisternum often missing, or hardly visible. Petiole low and stubby in profile with the anterior and posterior surface running almost parallele. Dorsal surface of petiole large and flat viewed from above, forming more approaching to a half circle than in coarctata . The subpetiolar tooth-like process of petiole forms a well-visible caudoventral triangular projection. Subpetiolar fenestra slightly larger than in coarctata . First gaster tergite with very dense hairs. Average distance of the fine cuticular foveolae on first gaster tergite 16.4 (14.7–18.7) µm. For morphometric measurements of altogether 106 workers see Table 1 .

Gyne ( Fig. 10 ) – Similar to the worker. Average body length 3.0 mm. Colour light ochraceous yellow to brownish yellow. Whole body with thick decumbent hairs. The frontal furrow clearly reaches the anteriormost ocellus. Scape with subdecumbent and suberect hairs. Tibiae with exuberant short, decumbent hairs only. Alitrunk relatively lower than in coarctata . Metanotal furrow between the katepisternum and anepisternum not shallow, hardly visible, or sometimes missing. Petiole low and stubby in profile, the anterior and posterior surface running almost parallel in profile. The subpetiolar tooth-like process of petiole often forms a well-visible triangular caudoventral projection. First gaster tergite with very dense hairs, similar to the worker.

Morphometric measurements of 9 gynes (SEIFERT): CS 671±12 [650,690], FoDG 16.36±1.03 [14.9,17.8], PEL/NOH 1.238±0.063 [1.130,1.342], PiMe 8.26±1.10 [7.3,10.0], CL/CW 1.234±0.023 [1.190,1.255], SL/ CS 0.811±0.011 [0.797,0.828], ML / CS 1.663±0.023 [1.635,1.703], MW / CS 0.761±0.025 [0.735,0.811], PEW/ CS 0.532±0.020 [0.513,0.571], PEL/ CS 0.441±0.014 [0.411, 0.457], NOH/ CS 0.357±0.017 [0.330,0.392].

Morphometric measurements of 16 gynes ( CSŐSZ): CL 725±14 [710,750], CW 588±13 [570,610], CS 656±13 [640,680], FR 76±5 [68,80], FL 160±7 [150,168], SL 530±12 [515,550], ML 1066±25 [1040,1110], MH 499±25 [480,550], PEH 442±15 [420,460], PH 240±11 [225,260], PL 240±10 [230,260], PEW 0.525±0.017 [0.50,0.566], CL/CW 1,233±0.0154 [1.210,1.256], FL/FR 2.113±0.928 [1.938,2.206], PH / PL 1.004±0.515 [0.923,1.061], ML / MH 1.972±0.051 [1.902,2.018], CL/SL 1.368±0.332 [1.324,1.413]

Male – Average body length smaller than 3.00 mm. Colour always black. Whole body with thick decumbent hairs. Head with sparse shorter hairs. Eyes with long hairs among the facetes. Alitrunk less robust inprofile view thanin coarctata .

Differential diagnosis and comments – In the workers, P. testacea differs from coarctata by a number of metric characters ( Table 1 ). These are smaller absolute size, lower distance of small foveolae on dorsum of first gaster tergite, a lower height vs length ratio of petiole node, a lower average pigmentation score, a lower height vs. length ratio of petiole, and a wider frons ratio (FR/ CS). These differences are repeated in the gynes. Both a multivariate discriminant analysis with 9 characters and 107 individuals ( CSŐSZ) and a multiple discriminant function with 7 characters and 126 individuals (SEIFERT) resulted inperfect separationof the two species ( Figs 13 , 14 ). A simplified discrimination for the practical use is provided by a multiplicative index CS × PH / PL ( Fig. 12 ) that is 546±27 [477–598] in testacea and 679±33 [618–776] in coarctata . Preliminary attempts to find reliable morphological discriminators of males, including genitalia characters, were not successful.

Designation ofthe lectotype – EMERY’ s original material of Ponera was found to contain no specimens with any type labelling as it had been stated by TAYLOR. Altogether 15 specimens were received from the Museo Civico di Storia Naturale , Genova and one specimen was fixed by present designation as lectotype of Ponera coarctata var. testacea EMERY, 1895 . The lectotype had beencollected inBonifacio by R EVEL in1876. Bonifacio is a small village at the S coast of Corsica. The specimen is in agreement with the type localities and descriptory statements given by EMERY who stated “… Francia meridionale, la Liguria, Corsica, e la Spagna … colore testaceo uniforme, o talvolta piu o meno bruno sul dorso…”. Other specimens that could belong to the original material of EMERY were not available from the Museo Civico. EMERY repeatedly sent ant material to FOREL and three specimens within the Forel collection in Muséum d’Histoire Naturelle, Genéve are considered by us as belonging to the original series. One specimen, collected by FLACH at Rapallo (Liguria) inMay 1891, and two other specimens (at one pin) labelled by EMERY “Gallia merid.” and with a blue label “Cotypus” made by FOREL were designated as paralectotypes during the course of this study.

Distribution and biology – EMERY (1895, 1909) roughly stated the occurrence of testacea in the Mediterranean region (S France, entire Italy, Corsica, and Spain). Our examinations revealed that it is widely distributed in C Europe and spreads north to 51°30’N at least (Eisleben, Germany). According to the data base of SEIFERT, 122 sites with Ponera are known from Germany, stretching north to 52°05’N (Niedersachsen). Since the possible occurrence of the sibling species has not been recognised earlier, voucher specimens were available for only 34 sites. P. testacea was found in 16 and coarctata in 24 localities with syntopic occurrence of both species inas many as 6 sites. P. testacea is in Germany typical for open and very xerothermous grasslands on limestone, silicate rock, or sand. It has so far not been found in xerothermous woodland with higher humidity and seems to be rare inurbanhabitats. P. coarctata is found in any of these habitat types, on the average it is less xerothermophilic and is expected to spread farther north.

InHungary, coarctata is widely distributed and frequently found in loessy grasslands as well as deep in forests, and occurs mainly in less xerothermous habitats. P. testacea oftenhad beencollected inwarm and dry habitats like sandy grasslands and dry rocky calcareous grasslands, altogether 20 localities are known with testacea and with only one syntopic occurrence of both species.

The situation in other countries is almost unknown. The few data of testacea from Switzerland, and Romania do not indicate rarity, since voucher specimens for coarctata are similarly sparse.

As result of the species splitting and the most difficult discrimination of coarctata and testacea , a post hoc interpretation of literature data is almost impossible and the natural history of both species must be written anew. However, the most detailed studies onthe biology of Central European Ponera ( LIEBIG et al. 1995, 1997) canbe exclusively referred to Ponera coarctata as our recent investigation of LIEBIG’ s voucher specimens showed.

Figures

Fig 12. Values of the index CS × PH / PL show no overlap between species based on altogether 139 workers (based on CSŐSZ’ dataset)

Figs 4–11. 4–5 = Lateral aspect of the petiole: 4 = Ponera testacea, 5 = P. coarctata; 6–7 = petiole feature of the worker from above: 6 = P.testacea, 7 = P.coarctata; 8–9 = Lateral aspect of the worker mesosoma and petiole: 8 = P. testacea, 9 = P. coarctata; 10–11 = Lateral aspect of the gyne

Fig. 13. Multivariate discriminant analysis of 103 worker individuals of Ponera coarctata and P. testacea as combined histogram for all groups. Distribution of the sample along the canonical root. Number of variables inthe model: 9, Wilks’ Lambda: 0.10839, F(9, 82) = 74.945, P <0.0001 (based on CSŐSZ’ dataset)

Fig. 14. Multiple linear discriciminant score D(7) = 0.068 FoDG +0.002 CS –0.43 PEL/NOH +0.02 PiMe –0.13 CL/CW –0.2 FR/CS –0.10 PEW/CS.for 126 individual workers of Ponera coarctata and testacea (based onSEIFERT’s dataset)

Tables

Table 1. Morphometric measurements of workers of Ponera testacea and P. coarctata. The length of a string of asterisks is proportional to the discriminative power of a character

	CSŐSZ	SEIFERT		CSŐSZ	SEIFERT
	coarctata (n = 72)	coarctata (n = 74)		testacea (n = 58)	testacea (n = 48)
CS	635 ±14	638±17	***	576±13	587±14
	[595, 662]	[597, 678]		[533, 613]	[547, 615]
CL/CW	1.243± 0.028	1.255 ±0.017	**	1.279± 0.035	1.278 ±0.018
	[1.183, 1.336]	[1.219, 1.302]		[1.242,1.387]	[1.239, 1.317]
SL/ CS	0.790± 0.098	0.811 ± 0.016		0.788 ± 0.116	0.811 ±0.013
	[0.751, 0.897]	[0.773, 0.851]		[0.782,0.841]	[0.787,0.833]
FL/ CS	0.251± 0.032	0.246 ± 0.008		0.238 ± 0.035	0.246 ±0.009
	[0.226, 0.283]	[0.231, 0.264]		[0.217, 0.261]	[0.231,0.272]
FR/ CS	0.115± 0.016	0.116 ±0.010	*	0.113± 0.018	0.125 ±0.010
	[0.100, 0.137]	[0.095, 0.137]		[0.102,0.139]	[0.101,0.147]
ML /CS	1.412 ± 0.173	1.470 ± 0.027		1.393 ± 0.205	1.477 ±0.039
	[1.359, 1.627]	[1.404, 1.521]		[1.351, 1.496]	[1.422, 1.564]
PEW/ CS	0.472± 0.059	0.481 ± 0.016		0.492 ± 0.073	0.498 ±0.020
	[0.448,0.559]	[0.443, 0.521]		[0.483, 0.535]	[0.452, 0.549]
PEH/ CS	0.667± 0.082			0.634 ±0.094
	[0.641, 0.780]			[0.617, 0.711]
PH / CS	0.364± 0.045			0.339 ±0.050
	[0.335, 0.441]			[0.325, 0.369]
MW / CS		0.662± 0.014			0.658 ±0.013
		[0.634,0.711]			[0.628,0.686]
PL / CS	0.339± 0.042			0.357 ±0.053
	[0.322,0.398]			[0.335,0.388]
PEL/ CS		0.425± 0.013			0.440 ±0.012
		[0.384,0.459]			[0.401,0.460]
FL/FR	2.196± 0.022			2.117 ±0.020
	[1.880,2.540]			[1.810,2.420]
PH / PL	1.073±0.043		*	0.950±0.034
	[1.000,1.190]			[0.900,1.050]
CL/SL	1.384± 0.029			1.394 ±0.022
	[1.310,1.460]			[1.330,1.440]
CSxPH/ PL	680±33		***	546±27
	[618,776]			[477,598]
NOH/ CS		0.367±0.011	*	0.346±0.013
		[0.337,0.388]		[0.322,0.370]