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Обзор ископаемых муравьев рода Myrmica из янтаря

"Insectes sociaux", Муравьиные НОВОСТИ: Виды-2007 и 2008, Статьи-2007 и 2008

Добавлено 27-02-2009

Обзор ископаемых муравьев рода Myrmica из янтаря

Alexander Radchenko1, Gennady Dlussky2 and Graham W. Elmes3
E-mail: rad@izan.kiev.ua

1 Museum and Institute of Zoology, Polish Academy of Sciences, 64, Wilcza str., 00-679, Warsaw,
2 Moscow State University, Vorob'evy gory, 119899, Moscow, Russia
3 Centre for Ecology and Hydrology, Benson Lane, Crowmarsh Gifford, Wallingford, Oxfordshire OX10 888, United Kingdom

The ants of the genus Myrmica (Hymenoptera, Formicidae) from Baltic and Saxonian amber (Late Eocene)

"Journal of Paleontology", 2008, Volume 81 (6), Pages 1494-1501 (November 2007)


      Обзор ископаемых муравьев рода Myrmica ( подсемейство Myrmicinae) из Балтийского и Саксонского янтаря из Позднего Эоцена с описанием новых видов: Myrmica paradoxa n. sp. (Германия) и Myrmica eocenica n. sp. (Россия, Калининградская обл.) Даны новые комбинации видов: Myrmica rudis (Mayr, 1868b) new combination и Myrmica intermedia (Wheeler, 1915) new combination.


      Проводя ревизию муравьев рода Myrmica Latreille, 1804, авторы обнаружили в коллекциях Российской Академии Наук и Польской Академии Наук кусочки Балтийского янтаря (Поздний Эоцен, около 40 млн.лет; возможно до Олигоцена, Wheeler, 1915; Larsson, 1978; Bolton, 1995; Dlussky, 1997), каждый из которых включал ископаемые остатки экземпляров Myrmica. Также удачно были исследованы кусочки Саксонского янтаря (того же примерно возраста, что и Балтийский янтарь) из коллекции Manfred Kutscher, которые содержали 3 экз. мирмик. Четыре экземпляра принадлежат двум новым видам: Myrmica paradoxa n. sp. и Myrmica eocenica n. sp. Приводятся их описания и сравнения с другими мирмиками, дается определитель 5 вымерших видов настоящих представителей рода Myrmica.

      До сих пор было описано 17 вымерших видов в роде Myrmica (Heer, 1850, 1867; Mayr, 1868b; Novak, 1878; Meunier, 1915): 4 вида из Позднего Эоцена и Олигоцена (около 40–35 млн.лет) и 13 видов из Миоцена (около 25 млн.лет). Однако, только один из них - M. longispinosa Mayr, 1868b - найден в Балтийском янтаре и бесспорно принадлежит к этому роду. Все остальные ископаемые якобы "Myrmica" виды принадлежат к другим родам муравьев. The high number of misidentifications is not surprising because most species were described a long time ago from impressions in rock. Such fossils are very difficult to interpret and identify correctly, even with considerable experience and a good knowledge of recent ant taxonomy. Три вида "мирмик", M. pusilla Heer, 1850, M. duisburgi Mayr, 1868b и M. rugiceps Heer, 1850 были формально перемещены в другие рода следующими авторами: Handlirsch (1907), Wheeler (1915) и Assmann (1970) соответственно (Mayr, 1867) ... ...

      Американский мирмеколог Wheeler (1915) также описал род Nothomyrmica emphasizing its similarity to Myrmica, from which it differs by only a single feature (considered very important at that time)—the lack of spurs on the middle and hind tibiae of workers. It contained only the four species placed there by Wheeler: N. intermedia Wheeler, N. rudis (Mayr, 1868b), N. petiolata (Mayr, 1868b) and N. rugosostriata (Mayr, 1868b); the latter three were first described in the genus Macromischa Roger, 1863 (now a synonym of Temnothorax Mayr, 1861; see Bolton, 1995, 2003).

      Since many extant Myrmica species are known to have reduced tibial spurs, or even completely lack them, the feature of reduced tibial spurs is considered much less diagnostic nowadays (Bolton, 1988a; Radchenko and Elmes, 2003). In our opinion, genus Nothomyrmica is quite heterogenic and artificially joins unrelated species (including some Myrmica) having one common feature—the absence of spurs on the hind and middle tibia. Nothomyrmica intermedia and N. rudis are very similar to M. longispinosa and all three closely resemble living species from the M. ritae-group (Radchenko, 1994; Radchenko and Elmes, 2001b). Therefore, we formally transfer N. intermedia and N. rudis to the genus Myrmica; N. rugosostriata is transferred to the newly described extinct genus Eocenomyrma Dlussky and Radchenko, 2006, and N. petiolata is transferred to the genus Temnothorax Mayr (Dlussky and Radchenko, 2006).


      Myrmica Latreille, 1804

      Myrmica Latreille, 1804, p. 179; type species—Formica rubra Linnaeus, 1758, p. 580, by subsequent designation of Latreille, 1810, p. 437.

      Nothomyrmica WHEELER, 1915, p. 60; type species — Macromischa rudis Mayr, 1868b, p. 85, fig. 85, by original designation; RADCHENKO, 1994, p. 48; BOLTON, 1995, p. 292, n. syn.

      Notes. For the diagnosis and full synonymy of the genus Myrmica see Bolton (1988a, 1995, 2003), Radchenko and Elmes (2003). Here, we have transferred two of the four Nothomyrmica species to the genus Myrmica, including type species of Nothomyrmica.

      Myrmica paradoxa sp. n.

      Diagnosis. Propodeum with two denticles that are short, straight, wide, blunt at the apex, directed upward and slightly backward. Frontal carinae short, very weakly curved to merge with the rugae, which surround antennal sockets. Only anterior (lower) part of frons with longitudinal rugae, remaining part of head dorsum with dense but not coarse reticulation. Whole alitrunk and waist with reticulation. Hind tibiae with conspicuous pectinate spur.

      Description. Head longer than broad, with convex sides and occipital margin, and broadly rounded, barely marked occipital corners. Eyes of moderate size, not prominent laterally, situated approximately at the midlength of lateral margins of head. Anterior clypeal margin very broadly rounded, not prominent, shallowly notched medially, with pair of long median setae and fringe of shorter setae on the anterior margin (similar to that of other Myrmica species). Antennae twelve-segmented and with distinct three-segmented club, second to ninth funicular joints distinctly longer than broad. Antennal scape relatively long, at least reaching or slightly surpassing the occipital margin of head, gradually curved at the base, without any trace of lobe or carina.

      Alitrunk short, wide, and robust. Promesonotum feebly convex (in lateral view) and in dorsal view with broadly rounded anterio-lateral corners, distinctly narrowing posteriorly (approximately twice wider anteriorly than posteriorly), without any trace of a promesonotal suture. Metanotal groove distinct, abrupt but not very deep. Dorsal surface of propodeum subequal to its declivity (= posterior surface), propodeal lobes not sharply pointed, at most bluntly angulated at the apex. Petiole with relatively short but distinct peduncle, its node quite massive, with rounded dorsum. Postpetiole subglobular and lower than petiole (in lateral view).

      Body with rather abundant, suberect to erect, moderately long, thin, pointed hairs. Antennal scape with subdecumbent to suberect hairs; hind tibiae and femora with short subdecumbent hairs, but with longer suberect hairs on the inner margin of femora.

      Etymology. The species named for its paradoxical combination of features.

      Material examined. Three workers (holotype and paratypes in the same piece of amber), GPMHU, No F-199, Germany, Saxonian Amber.

      Measurements (mm) and indices. Holotype: HW 1.12, FW 0.42, FLW 0.50, SL 0.99, AL 1.57, PNW 0.95; FI 0.38, FLI 1.20, SI2 0.89. Paratype specimen one: HL 1.27, SL 0.92, HTL 0.87; SI1 0.73. Paratype specimen two: PL 0.46. Body length is ca. 6 mm.

      Discussion. Myrmica paradoxa n. sp. well differs from any other known modern or fossil Myrmica species, especially by the very short propodeal denticles: such a shape of propodeal armature is peculiar, unknown in any modern or other fossil Myrmica species. It is paradoxical because on one hand it is characterized by features that are quite normal for the genus Myrmica, but on the other hand, others that are not known from any other Myrmica species. Thus, while it has a sculpture and shape of the head, pilosity of the body, shape of petiole and postpetiole and a distinct pectinate spurs on the hind tibiae that are typically Myrmica, its very well marked three-jointed antennal club and the generally short and robust alitrunk with the short blunt propodeal denticles are atypical. No other extant Myrmica species has such a short dorsal surface of the propodeum relative to the posterior surface (the propodeal declivity) and only the unusual queens of Myrmica luteola Kupyanskaya, 1990 have short propodeal denticles. Unfortunately, the maxillary and labial palpes, which can be diagnostic of certain genera, are invisible. We consider that most probably, M. paradoxa represents an extinct lineage within the genus.

      Myrmica eocenica sp. n.

      Diagnosis. Propodeum with very long, straight propodeal spines, that are directed backwards, strongly divergent (in dorsal view). Frontal carinae very weakly curved, not curving outwards, merging with the rugae, which run backwards to the occipital margin of the head. Body with coarse sculpture. Hind tibiae with conspicuous pectinate spur.

      Description. Head distinctly longer than broad, with convex sides, straight occipital margin, and narrowly rounded occipital corners. Eyes relatively small, convex, situated approximately at the midlength of lateral margins of head. Anterior clypeal margin narrowly convex, but not prominent, without notch, with pair of long median setae and fringe of shorter setae on the anterior margin (similar to that of other Myrmica species). Frons wide; frontal lobes slightly extended, quite narrow and pointed at the apex. Antennae twelve-segmented and with three (possibly four?)-segmented club, funicular joints second to eighth distinctly longer than broad. Antennal scape long, surpassing occipital margin of head, very feebly curved at the base, without any trace of lobe or carina. Frons between frontal carinae with only three coarse longitudinal, slightly sinuous rugae, which reach occipital margin; its lower (anterior) part between frontal lobes with four additional short longitudinal rugae, which extend only to the level of the eyes; upper (rear) part of frons and remainder part of head dorsum with coarse reticulation. Antennal sockets deep and surround by at least one coarse ruga. Clypeus with coarse longitudinal rugosity.

      Alitrunk relatively long and slender. Promesonotum somewhat flattened (in lateral view), with narrowly rounded anterio-lateral corners and with trace of promesonotal suture (in dorsal view). Metanotal groove distinct, but not very deep. Dorsal surface of propodeum somewhat longer than posterior one, propodeal lobes sharply pointed at the apex. Petiole with long peduncle, its node low and long, with weakly convex dorsum, postpetiole fig-shaped (in dorsal view). Whole alitrunk with coarse reticulation, only propodeal dorsum seems to be also having short longitudinal rugae (the last feature is not clearly visible in the investigated specimen). Petiolar node with longitudinally-concentric rugae, its dorsum also with moderate (not coarse) reticulation, postpetiole in dorsal view with longitudinal rugae.

      Body with a few, suberect to erect, thin, pointed hairs. Antennal scape, hind tibiae and femora with sparse subdecumbent, quite long hairs; inner margin of femora has longer suberect hairs.

      Etymology. The species named after geological epoch when the type specimen was trapped in amber.

      Material examined. One worker, holotype, PIN, 964/479, Kaliningrad Province of Russia, Baltic Amber.

      Measurements (mm) and indices. HL 1.11, HW 0.95, FW 0.41, FLW 0.52, SL 1.06, AL 1.34, ESL 0.63, ESD 0.62, HTL 0.91; CI 1.16, FI 0.43, FLI 1.28, SI1 0.96, SI2 1.12, ESLI 0.66, ESDI 0.98. Body length ca. 6.5 mm.

      Discussion. Myrmica eocenica n. sp. is similar to all the previously described Myrmica species from Baltic Amber. Unfortunately, types of all others appear to be lost (see above), so this comparison is based solely on Mayr’s and Wheeler’s original descriptions and drawings.

      It differs from M. longispinosa by its sharply pointed apically propodeal lobes, by the strongly divergent propodeal spines (compare Figs. 1.1 and 4, 5), and by its somewhat longer body ( 6 mm vs. ca. 5 mm). Compared to the former ‘‘Nothomyrmica’’ species, it differs by its well developed pectinate spurs on the hind tibiae; though this feature may be not so distinctive at the generic level (see above), it is usually more constant at the species level. It clearly differs from M. rudis by the shape of the propodeal spines and petiole: in M. rudis the spines are somewhat curved and directed backward and upward while its petiole has a much shorter peduncle and subtriangular node (compare Figs. 1.2 and 4, 5). Myrmica eocenica most resembles M. intermedia by the shape and sculpture of its body but, apart from the tibial spurs, it also differs by its straight and strongly divergent propodeal spines (compare Figs. 1.3 and 4, 5), and body length ( 6 mm vs. 4.7 mm). Generally, M. eocenica is very similar to living species of the ritae-group of Myrmica (see Radchenko, 1994; Radchenko and Elmes, 1998, 2001b) and almost certainly should be placed in this group.

      Myrmica longispinosa
Mayr, 1868b new comb.
      Occurrence. Kaliningrad Province of Russia, Baltic Amber.

      Myrmica rudis
(Mayr, 1868b) new comb.
=Macromischa rudis MAYR, 1868b,
=Nothomyrmica rudis (Mayr, 1868b).

      Occurrence. Kaliningrad Province of Russia and northwestern Poland, Baltic Amber.

      Myrmica intermedia
(Wheeler, 1915) new comb.
=Nothomyrmica intermedia WHEELER, 1915

      Occurrence. Kaliningrad Province of Russia, Baltic Amber.


      Большинство учёных рассматривают трибу Myrmicini (включающую роды Myrmica, Manica Jurine, 1807, Eutetramorium Emery, 1899, Hylomyrma Forel, 1912, Pogonomyrmex Mayr, 1868a и Secostruma Bolton, 1988b) в качестве наиболее морфологически базальной в подсемействе Myrmicinae, based on the plesiomorphic state of features, such as the structure of the mesosoma with well defined sutures, six-jointed maxillary and four-jointed labial palpes, and the presence of pectinate spurs on the mid and hind tibia. A recent molecular phylogeny of ants (Moreau et al., 2006) suggests that subfamily Myrmicinae arose more than 100 Ma prior to a huge diversification of ants that the authors believe was related to the evolution of angiosperm plants. By the Late Cretaceous (about 80 Ma) the subfamily already comprised five or so, distinct (‘‘old’’) lineages and most of the modern genera were established during the next 35 million years before the start of the Eocene. Interestingly, they found that the three studied representatives from tribe Myrmicini fall in two of the ‘‘old’’ lineages: Myrmica and Eutetramorium are in one, while Pogonomyrmex is in another that includes Messor Forel, 1890 and Aphaenogaster (both latter genera belong to the tribe Pheidolini). When an even more detailed molecular phylogeny is made, the tribe Myrmicini will probably have to be revised.

      Мирмекологи Radchenko and Elmes (2001b) считают, что анцестральные Myrmicini вероятно были наиболее сходны с современными Myrmica, или возможно Manica, because the other four genera possess many derived features (see Bolton, 2003 and literature cited therein). They had considered that any resemblance between Messor and Pogonomyrmex was solely due to convergent evolution, but the Moreau et al. study (loc. cit.) suggests otherwise. Despite Manica species appearing to have more plesiomorphic features, e.g., a full complement of mesosomal sutures with some large specimens having a rudimentary scutellum, filiform antennal funiculus in males, Radchenko and Elmes (2001b) believed Manica and Myrmica most probably derived from a common ancestor at about the same time. Unfortunately Moreau et al. (2006) did not sample Manica species, but the general thrust of their results support that idea. Furthermore, if both genera arose 60–70 Ma during a burst of ant diversification and before North America was completely separated from Eurasia, it would explain both the modern distribution of Manica (restricted to North America except for two species) and the general dissimilarity between North American and Eurasian Myrmica species-groups.

      Четыре из пяти известных ископаемых видов рода Myrmica очень сходны с современны ми представителями группы ritae-group. These species are usually considered as basal in relation to the extant forms (Radchenko, 1994; Radchenko and Elmes, 2001b; and above). They are very poorly represented in collections and were considered to be unusual and rare until recent studies showed that they are relatively numerous and common in certain subtropical forests on mountains in the extreme southeastern part of the Palaearctic and in the north of the Oriental Regions (Radchenko and Elmes, 1998, 1999, 2001a, 2001b; Radchenko et al., 2001). Some species are superficially very similar to Aphaenogaster (tribe Pheidolini) and are frequently misidentified as such (see Radchenko and Elmes, 1998) but we believe that this is genuine convergence similar to, for example, that between Cataglyphis FoЁrster, 1850 (tribe Formicini) and Myrmecocystus Wesmael, 1838 (tribe Lasiini). Weber (1950) also noted that M. ritae Emery, 1889 and related species have similar characteristics to some tropical species of Aphaenogaster and therefore suggested that they might have similar habits, such as a more arboreal way of life than is normal for Myrmica. Indeed, recent field studies of some ritae-group species (G. W. Elmes, A. Schulz, and K. Eguchi, personal commun., 2003) suggest that some species might live and forage entirely on trees.

      We suggest that the putative ancestor of modern Myrmica lived in the warm early Eocene forests and by the late Eocene had given rise to M. ritae-like species and other unusual forms, such as the extinct M. paradoxa n. sp. or extant M. mirabilis Elmes and Radchenko, 1998. While it is possible that the ancestors of most other modern Myrmica species-groups had by this time already developed a greater cold tolerance and had penetrated the more northerly temperate forest zone, it is unlikely because there is a total absence of Myrmica specimens in the Oligocene and Miocene deposits of Eurasia that are associated with temperate conditions, despite the presence of quite a rich ant fauna, including several Aphaenogaster species that are morphologically and ecologically similar to Myrmica (Theґobald, 1937; Dlussky, 1981; Zhang, 1989). More probably, ancestor(s) of the different modern species- groups remained restricted to the warmer Eocene forests, coexisting with M. ritae-like species. Given our current understanding of the available fossil records and the paucity of fossil Myrmica, represented by only a few specimens found in amber deposits, we suggest that at the end of the Eocene, Myrmica species might have been generally rare with restricted distributions. Yet, some of the early Myrmica species survived the ecological trauma of the post-Eocene cooling and adapted to the cooler habitats.

      Unfortunately, there are no data on the Pliocene ant fauna of Eurasia to indicate how adaptive radiation proceeded to the extent that Myrmica is now one of the dominant ant genera of the temperate Holarctic, whether measured in terms of the number of species, colonies or individuals. Indubitably, isolation due to alternating glacial ages of the last millions of years played an important role in subsequent speciation. On the other hand, typical ritae-group species and a few other basal forms, for example M. mirabilis (rather similar to Manica in the general shape of the body, but in other features clearly Myrmica), were confined to forests on high mountains in subtropical South and Southeast Asia, where many species of plant and animal found in late Eocene amber still persist (Larsson, 1978).

      Интересно, что большинство современных видов Myrmica с предположительно базальными морфологическими чертами в основном обнаружены в южных горных системах. There are numerous unusual and endemic Myrmica species, including presumably basal forms, in the southwestern Himalayas, which were isolated from the rest of the Myrmica fauna at some time in the late Miocene – early Pliocene. We have found some similar atypical Myrmica in the mountains of Turkey (Elmes et al., 2002) and few unusual species have been described from the southern mountains of the USA (see also Radchenko and Elmes 2001b). Almost certainly, more rare, basal forms of Myrmica remain to be discovered in the southern mountains of North America. It would not surprise us if species similar to M. ritae were discovered on high mountains in Mexico, since Tertiary flora and fauna have persisted in the Americas because migration in response to climate change is facilitated by the northsouth orientation of mountain systems.

      Морфология ископаемы Myrmica, discussed above, suggest that female castes of the ancestral species probably had long, gently curved antennal scape, with no thickening or lobes. A short scape in males is plesiomorphic evolutionary condition for all ants (Dlussky, 1983; Dlussky and Fedoseeva, 1988); therefore the males of the ancestral Myrmica species probably had short antennal scapes. This is the case in most ritae-group species for which this character is known (Radchenko and Elmes, 1998). The two morphological features that are most important in defining modern species-groups (sharply angled antennal scape of the female castes and longer antennal scape among males, see Radchenko and Elmes, 2001b) probably derived from the ancestral form during the adaptive radiation of Myrmica following the post-Eocene cooling.


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