Research Article |
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Corresponding author: Bernhard Seifert ( bernhard.seifert@senckenberg.de ) Academic editor: Thomas Schmitt
© 2025 Bernhard Seifert.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Seifert B (2025) Treachery pigmentation pattern leads to misidentification: Tapinoma melanocephalum (Fabricius), Tapinoma pygmaeum (Dufour) and Tapinoma jandai sp. nov. (Hymenoptera, Formicidae). Contributions to Entomology 75(2): 245-252. https://doi.org/10.3897/contrib.entomol.75.e154879
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Discrete color morphs of ants, in the case reported here morphs with whitish versus blackish gaster pigmentation, may be misinterpreted as different species. This became clear in a morphometric study including 91 worker individuals of Tapinoma jandai sp. nov., Tapinoma melanocephalum (Fabricius, 1793) and Tapinoma pygmaeum (Dufour, 1857). The three species are clearly separable by a principal component analysis of 17 morphological characters under exclusion of pigmentation characters. This result is confirmed by a clear separation in a linear discriminant analysis with the classification of any specimen being confirmed with posterior probabilities of p > 0.9994. The study showed that just the holotype series of Tapinoma pithecorum Seifert, 2022 is a rare color variant of Tapinoma pygmaeum and consequently the former taxon is a junior synonym of the latter. In contrast, all remaining samples that formed the basis of the description of Tapinoma pithecorum are recognized as an undescribed species which is introduced here as Tapinoma jandai sp. nov. A consideration of 14 taxa of minute species showing similarities to the new species revealed that none of these is suspected of representing a senior synonym of T. jandai sp. nov. and that Tapinoma indicum Forel, 1895 should be transferred to the genus Ravavy Fisher, 2009. Based on examples in Formica, Camponotus, Cardiocondyla and the two cases reported here, it is argued that ant classification by simple color patterns is under increased risk of misclassification.
Ant taxonomy, color dimorphism, ghost ant, numeric morphology-based alpha-taxonomy
The recent description of Tapinoma pithecorum by Seifert, 2022 that was supposed by
These alerting findings prompted a critical reinvestigation of the Tapinoma pithecorum material and an extension of sample size under inclusion of T. pygmaeum. The unpleasant result was that just the holotype series of Tapinoma pithecorum from the greenhouse (but not any other sample I had named T. pithecorum) belonged to T. pygmaeum when the evaluation excludes gaster pigmentation. The holotype series represents a probably rare case of Tapinoma pygmaeum nest population having a whitish gaster being distinctly lighter than the blackish brown head and mesosoma. On the other hand, in structurally and morphometrically clear Tapinoma melanocephalum which usually show in dried condition dark heads and mesosomas and whitish gasters, homogenously dark samples occur rarely. I suppose that alternate gaster pigmentation in these related species is directed by a simple mutation and cannot be used as a leading taxonomic character. Since we need a taxonomically available name for the outdoor populations named by
The backbone of the investigations presented here is Numeric Morphology-Based Alpha-Taxonomy (NUMOBAT,
Details of equipment, measuring, data analyses and classification procedures are given in
CL Maximum cephalic length measured between points A and B; A is the posteromedian margin point of head capsule; B is an imagined median point situated at the same transversal level as the most anterior points of clypeus left and right of clypeal excision. Bilateral asymmetries are averaged.
CS Cephalic size with CS=(CL +CW)/2.
CW Maximum cephalic width.
dAN Minimum distance of the inner (centripetal) margins of antennal socket rings which is best measurable in dorsofrontal view.
EL Eye length; maximum diameter of the compound eye over all structurally defined ommatidiae; bilateral mean.
ExCly Maximum depth of anteromedian clypeal excision as it appears in frontodorsal view and with median line of head positioned perpendicular in the visual field; bilateral asymmetries are averaged.
ExClyL/W Ratio ExCly /ExClyW.
ExClyW Width of clypeal excision at the level of the base centers of the two most apical and largest setae.
ExOcc Depth of excavation of posterior vertex with head in full face view. Procedure: Using high magnification with low depth of focus, adjust the head to the measuring position defined for CL. Focus both posterior corners of vertex until they form a sharp contour, adjust them to equal horizontal level within the visual field and superimpose the corners with the horizontal line of the cross-scale. Change the focal level until the median part of posterior vertex forms a sharp contour. Read the depth.
Full face view Dorsal aspect of head adjusted for measuring both maximum CL and CW. Use of high magnifications with low depth of focus improve the correct spatial adjustment.
Fu2L Median length of second funiculus segment in dorsal view. Dorsal view is given when the swiveling plane of 1st funiculus segment is positioned in the visual plane. Take care to really measure median length as the segment’s sides have unequal length and to recognize the real distal margin of the segment. The latter has a very thin cuticle, frequently producing a narrow, shining ribbon that seems to be, by optical impression, demarcated from the rest of the segment. The median line of the segment is visualized by centre of the patch reflecting the coaxial light.
Fu2W Maximum width of second funiculus segment in dorsal view. Procedure: use the same spatial adjustment of the ant as in measuring Fu2L but rotate it in the visual plane by 90°. The use of transmitted-light is important to visualize the real cuticular surface which is often obscured by pubescence.
Fu3L Median length of third funiculus segment in dorsal view measured under the same conditions as given for Fu2L.
IFu2 Index Fu2L / Fu2W.
ML Mesosoma length from the caudalmost point of lateral metapleuron to rear margin of anterior pronotal fringe (in workers). In workers, if anterior measuring point is concealed, keep the orientation of measuring line, choose a higher magnification, measure from the caudalmost point of lateral metapleuron to the level of promesonotal border and multiply by 1.415. This factor is an average estimated from several species.
MGr Depth of metanotal groove / depression in lateral view; the upper reference line extends between the highest points of mesonotum and propodeum perpendicular to which depth measuring is performed.
MW Maximum pronotal width.
nExCly Bilateral sum of pubescence hairs and smaller setae protruding just visibly at magnifications > 200× across margin of clypeal excision. The two large anteriormost setae are not counted.
PoOc Postocular distance taken in the same spatial adjustment as in measuring CL: distance from the transversal level of posterior eye margin to median point of hind margin of head; bilateral asymmetries are averaged.
SL Scape length excluding articulatory condyle. Transmitted light is useful to visualize the real distal end of scape that may be obscured by pubescence.
The separation of Tapinoma melanocephalum, T. pygmaeum and T. jandai sp. nov. was successful on worker individual level without any character selection or the necessity to form nest sample means. A principal component analysis considering all 17 numeric characters given in Table
Morphometric data of three Tapinoma species in the sequence arithmetic mean ± standard deviation [minimum, maximum]. The columns with ANOVA data are placed between the columns of the compared species. F values of the strongest differences are given in heavy type.
| Tapinoma melanocephalum (i = 51) | ANOVA F1,66, p | Tapinoma jandai (i = 17) | ANOVA F1,45, p | Tapinoma pygmaeum (i = 30) | |
|---|---|---|---|---|---|
| CS [µm] | 438 ± 15 [409, 493] | 11.74, 0.001 | 454 ± 21 [422, 489] | 31.80, 0.000 | 489 ± 20 [443, 517] |
| CL/CW | 1.161 ± 0.027 [1.103, 1.213] | 8.17, 0.006 | 1.141 ± 0.017 [1.107, 1.169] | 0.05, n.s. | 1.143 ± 0.028 [1.101, 1.218] |
| SL/CS | 0.981 ± 0.023 [0.907, 1.025] | 456.8, 0.000 | 0.841 ± 0.023 [0.810, 0.907] | 1.02, n.s. | 0.847 ± 0.017 [0.815, 0.889] |
| ExOcc/CS [%] | 0.21 ± 0.30 [0.00, 1.39] | 8.76, 0.004 | 0.47 ± 0.37 [0.00, 1.27] | 15.72, 0.000 | 0.98 ± 0.46 [0.08, 2.19] |
| ExCly/CS [%] | 1.51 ± 0.49 [0.64, 2.94] | 42.3, 0.000 | 0.63 ± 0.46 [0.00, 1.66] | 0.08, n.s. | 0.60 ± 0.29 [0.00, 1.10] |
| ExClyW/CS [%] | 12.66 ± 1.45 [9.80, 15.49] | 4.21, 0.044 | 11.79 ± 1.70 [7.07, 14.60] | 0.73, n.s. | 11.34 ± 1.71 [8.98, 16.52] |
| ExClyL/W [%] | 0.119 ± 0.036 [0.05, 0.23] | 43.44, 0.000 | 0.053 ± 0.035 [0.00, 0.11] | 0.01, n.s. | 0.054 ± 0.028 [0.000, 0.114] |
| nExCly | 1.88 ± 1.11 [0.0, 5.0] | 11.64, 0.001 | 3.24 ± 2.11 [0.0, 9.0] | 28.00, 0.000 | 0.93 ± 0.86 [0.0, 3.0] |
| dAN/CS | 0.339 ± 0.008 [0.318, 0.358] | 33.48, 0.000 | 0.326 ± 0.009 [0.313, 0.342] | 2.33, n.s. | 0.323 ± 0.007 [0.304, 0.337] |
| PoOc/CL | 0.469 ± 0.011 [0.434, 0.488] | 57.28, 0.000 | 0.495 ± 0.014 [0.461, 0.518] | 1.96, n.s. | 0.499 ± 0.009 [0.476, 0.511] |
| EL/CS | 0.274 ± 0.010 [0.255, 0.307] | 113.5, 0.000 | 0.246 ± 0.008 [0.234, 0.260] | 0.58, n.s. | 0.245 ± 0.007 [0.230, 0.259] |
| MGr/CS [%] | 2.23 ± 0.80 [1.19, 4.52] | 19.21, 0.000 | 1.31 ± 0.56 [0.20, 2.57] | 2.91, n.s. | 1.56 ± 0.44 [0.72, 2.47] |
| MW/CS | 0.635 ± 0.018 [0.594, 0.676] | 57.3, 0.000 | 0.675 ± 0.022 [0.650, 0.737] | 48.53, 0.000 | 0.621 ± 0.027 [0.561, 0.683] |
| ML/CS | 1.240 ± 0.040 [1.119, 1.320] | 69.91, 0.000 | 1.152 ± 0.027 [1.094, 1.196] | 1.10, n.s. | 1.161 ± 0.027 [1.119, 1.228] |
| Fu2/CS [%] | 8.60 ± 0.38 [7.62, 9.35] | 179.3, 0.000 | 7.26 ± 0.28 [6.58, 7.65] | 67.32, 0.000 | 8.22 ± 0.43 [7.45, 9.10] |
| Fu3/CS [%] | 12.42 ± 0.73 [10.83, 13.77] | 176.0, 0.000 | 9.69 ± 0.75 [8.66, 11.42] | 0.74, n.s. | 9.83 ± 0.37 [9.20, 10.79] |
| IFu2 | 0.873 ± 0.046 [0.775, 0.963] | 93.57, 0.000 | 0.751 ± 0.040 [0.671, 0.808] | 113.6, 0.000 | 0.922 ± 0.059 [0.842, 1.095] |
Principal component analysis of worker individuals of three Tapinoma species based on 17 morphological characters. T. jandai sp. nov. (crosses), T. pygmaeum of normal pigmentation (black dots), T. pygmaeum with whitish gaster (white dots = the holotype series of T. pithecorum), T. melanocephalum with whitish gaster (white squares), T. melanocephalum with dark gaster (black squares).
In order to assess the probability of the placement of the individuals in the PCA, a character-reduced linear discriminant analysis (LDA) was run using the first four principal components as characters (Fig.
Linear discriminant analysis of worker individuals of three Tapinoma species based on four morphological characters. T. jandai sp. nov. (crosses), T. pygmaeum of normal pigmentation (black dots), T. pygmaeum with whitish gaster (white dots = the holotype series of T. pithecorum), T. melanocephalum with whitish gaster (white squares), T. melanocephalum with dark gaster (black squares).
The morphometric data presented in Table
539.82*Fu2-41.107*MW-6.910
offers a perfect separation with all Tapinoma jandai sp. nov. having values < 0 and all T. pygmaeum values > 0. A safe separation of T. jandai sp. nov. from T. melanocephalum is possible by the discriminant
80.53*PoOc-74.20*SL+54.56*MW-4.648
with all Tapinoma jandai sp. nov. having values > 0 and all T. melanocephalum values < 0.
The name is given in honor of Milan Janda who has collected the holotype specimen and provided valuable other material for this study.
• Holotype worker labelled “PNG: 5.104°S, 145.775°E, 15 m, Madang: Baitabag, partially fragmented lowland rainforest, M. Janda 2004.01.01-MJ13337” and “Holotype Tapinoma jandai Seifert”; • one paratype with the same collecting data label on another pin; • 8 mounted paratype workers on two pins labelled “ INDIA: 31.38682°N, 76.47164°E, Himachal Pradesh: Bakhra, 650 m, H. Bharti 2016.05.12 -EX1750”; • 38 paratype workers with identic labelling stored in ethanol; depository SMN Görlitz. The paratype sample coded EX1750 was included in the genomic investigation of
Combining the knowledge from this study with two genetically identified samples (
Worker (Figs
It is a tropical species with basically unknown biology. The holotype sample was collected in a partially fragmented lowland rainforest, the paratype in a dense woodland and the genetically identified sample from China in a tropical botanical garden. The frequency of Tapinoma jandai sp. nov. in all collections and all antweb.org images studied is about 10% of that of T. melanocephalum and there seems to be no worldwide spreading. This indicates that Tapinoma jandai sp. nov. has a much lower tramp species potency than T. melanocephalum – if it has any.
The large divergence time of 13 My between Tapinoma jandai sp. nov. and T. melanocephalum estimated by
Tapinoma minutum Mayr, 1862
The imaged type from Sidney (CASENT0915549) differs by a much longer ratio SL/CL and by the eyes placed more towards the median.
Tapinoma indicum Forel, 1895
The imaged type from Poona (CASENT0909774) has a much shorter head, much broader clypeus and the eyes placed more towards the median. The propodeal profile is not visible in the image but according to head morphology and following the worker diagnosis given by
Tapinoma fragile Smith, 1876
The imaged type from Mauritius: Rodigues (CASENT0102974) has a much longer scape, much narrower mesosoma and is probably a synonym of Tapinoma melanocephalum.
Tapinoma funiculare Santschi, 1928
The imaged type from Tonkin, Yambey (CASENT0911570) differs in having a remarkably oval head in dorsal view and a much longer scape.
Tapinoma atriceps Emery, 1888
The imaged type from Brazil, Rio Grande do Sul (CASENT0904029) has a much longer scape and a narrower mesosoma. The taxon is suspicious to represent a junior synonym of Tapinoma melanocephalum.
Tapinoma breviscapum Forel, 1908
Two imaged type specimens from São Paulo, Raiz da Serra (CASENT0909768, CASENT0922450) differ radically in the curvature of anterior clypeal margin and in having a much larger absolute size. Head width CW is 499 and 512 µm in the Tapinoma breviscapum types whereas it is in T. jandai sp. nov. 423 ± 21 [393, 456] µm. Furthermore, the T. breviscapum types show a much smaller ratio MW/SL of 0.693 and 0.689 which is in T. jandai sp. nov. 0.798 ± 0.025 [0.758, 0.855].
I wish to thank Christophe Galkowski, Milan Janda, John Longino, Wouter Dekoninck, Raymond Delarze, Beatrice Viussoz and Brian Fisher for sending collection material. The equipment for the imaging was co-financed by tax money on the basis of the state budget passed by the Sächsischer Landtag according to the Antragsnummer 100590787 of the Sächsische Aufbaubank issued 3 August 2021.
Coordinates and morphometric data in millimeters
Data type: xlsx
Explanation note: Occurrence, morphometric data: primary measurements in millimeters, geographic coordinates and altitude (LAT, LON, ALT).