Genus Cardiocondyla Research Articles

Cardiocondyla hashemi sp. n., a new species of the C. batesii species-group (Hymenoptera: Formicidae) from Saudi Arabia, with a key to the Saudi species

A new species, Cardiocondyla hashemi sp. n. is described and illustrated using a Scanning Electron Microscope (SEM) based on the worker caste. Cardiocondyla hashemi is a member of the C. batesii species-group with a resemblance to C. tenuifrons Seifert, 2003 from Jordan. Cardiocondyla hashemi can be readily distinguished by the uniform yellow body, the rare or distinctly scattered foveolae on the posterior third of cephalic surface, the broadly medially concave anterior clypeal margin, the sculptured mesosomal dorsum except for the mesonotum which is shallowly sculptured with faint longitudinal rugae, and petiolar node distinctly densely microreticulate. Ecological and biological remarks on the type locality are given. A key to the Saudi fauna of the genus Cardiocondyla is presented. http://www.zoobank.org/urn:lsid:zoobank.org:pub:1DB79558-0CE0-413B-A6EB-9E0ABB884882

Genetic Diversity in Pioneer Ants: The Cardiocondyla shuckardi Group

The myrmicine ant genus Cardiocondyla consists of around 80 species of small, inconspicuous ants, which live mainly in subtropical and tropical Africa, Eurasia, and Australia. Several species have been accidentally introduced to America and have also invaded numerous originally ant-free islands around the world. The diversity of life histories in this genus, with lethally fighting wingless males, dominance hierarchies among queens, and considerable variation in the sociogenetic organization of colonies across species, has made it an interesting model to investigate the evolution and behavioral ecology of reproductive strategies. Taxa of the African Cardiocondyla shuckardi group are of particular interest, as in a phylogeny they lie between facultatively polygynous species with fatal male competition and monogynous species with mutually tolerant males. Studies on male behavior and sociogenetics in C. “venustula,” a widespread member of the C. shuckardi group, showed that males defend small territories inside their subterraneous nests. At the same time, they revealed a surprisingly large variability in mtDNA haplotypes both within and between populations in South Africa, Angola, and Côte d’Ivoire. As the different lineages are very similar in morphology and nuclear markers, more research is needed to clarify the very confused taxonomic situation in this fascinating group of ants.

A revision of the Palaearctic species of the ant genus Cardiocondyla Emery 1869 (Hymenoptera: Formicidae).

A synopsis of the Palaearctic species of the ant genus Cardiocondyla Emery 1869 is provided. The four species groups which are of Palaearctic origin or which are restricted in their distribution to this faunal zone, the C. elegans, C. ulianini, C. batesii and C. stambuloffii group-consisting of 25 recognized species-are taxonomically revised. Further nine species belonging to species groups of Ethiopic and Oriental origin which may penetrate into and build up true outdoor populations in the southern Palaearctic are only treated in the determination key. The main working rationale of this revision is Numeric Morphology-Based Alpha-Taxonomy (NUMOBAT) with formation of species hypotheses largely based on exploratory data analyses and checking these hypotheses by discriminant analysis. NUMOBAT data of the species considered comprise 727 worker samples with 1555 individuals and 23,300 primary data. Including high-resolution photos of surface microstructures, all species are depicted by z-stack imaging in four standard visual positions. Numeric data on 19 phenotypical characters are presented in comparative tables and supplementary verbal descriptions are given. In contrast to species groups with Ethiopian, Oriental and Australasian origin, no member of the four Palaearctic species groups has developed a tramp species potential to spread globally. Four cryptic species are described as new: Cardiocondyla dalmaticoides n. sp., C. caspiense n. sp., C. verdensis n. sp. and C. rolandi n. sp. Confirmed were the synonymies of Cardiocondyla elegans santschii Forel 1905, C. provincialis Bernard 1956 and Xenometra gallica Bernard 1957 with C. elegans Emery 1869, that of C. elegans eleonorae Forel 1911 with C. bulgarica Forel 1892, that of C. elegans torretassoi Finzi 1936 with C. nigra Forel 1905, and that of C. bogdanovi Ruzsky 1905, C. montandoni Santschi 1912 and C. stambuloffii taurica Karavajev 1927 with C. stambuloffii Forel 1892. Cardiocondyla bicoronata Seifert 2003 was newly synonymized with C. nigra Forel 1905.

The Ant Genus Cardiocondyla (Hymenoptera: Formicidae): The Species Groups with Oriental and Australasian Origin

The world-wide fauna of the genus Cardiocondyla is assessed to have 79 currently recognized species and is subdivided into 14 informal species groups plus three monotypical clades. It is argued that the phylogenetic placement of Cardiocondyla next to the genus Ocymyrmex Emery, 1886 as proposed by Blaimer et al. (2018) is doubtful. The main body of the paper is a species-level taxonomic revision of those six species groups and the one monotypical clade which have their centers of origin in the Oriental and Australasian regions. The fauna of this area comprises 26 species eleven of which are described as new: Cardiocondyla schulzi n.sp., C. argentea n.sp., C. semiargentea n.sp., C. argyrotricha n.sp., C. latifrons n.sp., C. micropila n.sp., C. allonivalis n.sp., C. excavata n.sp., C. goroka n.sp., C. subspina n.sp. and C. sulcata n.sp. Recognized as junior synonyms were: Cardiocondyla tsukuyomi Terayama, 1999 and C. breviscapus Seifert, 2003 of C. minutior Forel, 1899 and C. brevispinosa (Donisthorpe, 1948) of C. paradoxa Emery, 1897. Each confirmed species is characterized by verbal descriptions, 18 numerically recorded phenotypic characters, z-stack images in three viewing positions, the key, a differential diagnosis and, if available, a brief life history. Exploratory and hypothesis-driven data analyses of phenotypic characters support the separation of the cryptic species Cardiocondyla minutior Forel, 1899, C. goa Seifert, 2003 and C. tjibodana Karavajev, 1935, two of which have dispersed as tramp species to the New World. Missing development of postzygotic isolation due to very strong prezygotic barriers is indicated in the closely related species Cardiocondyla latifrons n.sp. and C. micropila n.sp. which produce fertile hybrids in laboratory rearings but apparently do not hybridize in a natural context even if nesting in close neighborhood. The strong prezygotic isolation is supposed to be generated by the prevalence of intranidal mating and the difficulty for a mated heterospecific gyne to become accepted in a foreign colony.

First Record of Cardiocondyla obscurior Wheeler, 1929 (Hymenoptera: Formicidae: Myrmicinae) for Pakistan

Ants of the genus Cardiocondyla are considered omnivorous in nature. This genus is native to Afrotropical, Australasia, Indomalaya, Malagasy, Oceania, and Palearctic regions. A total of 72 valid species and 2 valid subspecies has been described worldwide. However, little is known about these ants in Pakistan. As a result of a survey of different sites in the Gatwala park of district Faisalabad during 2018, we collected individuals of the genus Cardiocondyla. Collected specimens were identified using the most recent and available literature. Prior to current work, only two species of this genus were reported for Pakistan. In this study we added the first record of Cardiocondyla obscurior for Pakistan, followed by a brief description, distribution, and identification key.

Courtship with two spoons-Anatomy and presumed function of the bizarre antennae of Cardiocondyla zoserka ant males.

Mating in ants often occurs on the wing during nuptial flights or on the ground when scattered female sexuals attract males by pheromones. In both scenarios, there is little opportunity for males to engage in prolonged aggressive competition or elaborate courtship displays. Male morphology is therefore adapted to locating female sexuals and mating, and it lacks specific weapons or other traits associated with courtship. In contrast, sexuals of the ant genus Cardiocondyla typically mate in their natal nests. As a consequence, in many species winged males have been replaced by wingless fighter or territorial males, which kill or expel rival males with their strong mandibles and show complex mating behavior. However, no wingless males are known from Cardiocondyla zoserka from West Africa, and instead, winged males have evolved a bizarre secondary sexual trait: uniquely shaped antennae with spoon‐like tips that show heavily sculptured ventral surfaces with numerous invaginations. We here report on the courtship behavior of C. zoserka males and describe antennal glands with class 3 gland cells, which presumably secrete a close range sex pheromone. Antennal glands have not yet been found in males of other ant species, including a close relative of C. zoserka, suggesting that in ants with intranidal mating sexual selection can rapidly lead to highly divergent adaptations and the evolution of novel structures.

Morphometry and colony structure of ants of the genus Cardiocondyla (Hymenoptera: Formicidae) from Georgia

The ant genus Cardiocondyla is characterised by a remarkable variability of reproductive life histories and colony structures. Little is known about the C. stambuloffii group, whose distribution extends from southeast Europe to central and eastern Asia. We here present data on the structure of Cardiocondyla colonies from southern Georgia. Morphometry and sequencing of the mitochondrial cytochrome c oxidase subunits I and II indicates that it is distinct from the other species of this group (C. gibbosa, C. koshewnikovi, C. stambuloffii, and C. tibetana). Microsatellite analysis revealed a high nestmate relatedness and frequent inbreeding, matching the observation of exclusively wingless males and the phylogenetic position of the group in the “Palaearctic clade,” in which colonies are typically single-queened.

Emeryia, Xenometra, zoserka: it's a boy, again! The misleading morphology of Cardiocondyla male ants

Wingless “ergatoid” males of the ant genus Cardiocondyla have repeatedly been described as females of novel genera of workerless social parasites (Emeryia and Xenometra), which after recognition of their real nature were synonymized with Cardiocondyla. Examination of ants newly collected from Comoe National Park in Cote d’Ivoire, West Africa revealed that winged Cardiocondyla males have sparked a similar misidentification: the “winged female sexuals” of the supposed inquiline Cardiocondyla zoserka Bolton 1982, known only from the type material from Nigeria, are in fact the winged males of their presumed host, a species of the Cardiocondyla shuckardi group (sensu Seifert 2002). The types of C. zoserka are immediately recognizable by their strangely modified antennal funiculi with cup-shaped apical segments, which do not resemble any other known ant antenna. More than twenty winged individuals from two Cardiocondyla colonies from the flood plain of Comoe River exhibited the same strange modifications of their antennae and were otherwise also similar to the C. zoserka type specimens. However, these winged ants turned out to be winged males rather than winged female sexuals. The inspection of the holotype and one paratype of C. zoserka verified that they also were males: like the winged males from Comoe N.P., they have concealed male genitals and ocelli, which are considerably larger than those of the female sexuals of Cardiocondyla. All female sexuals and workers found in the colonies with the bizarre winged males from Comoe N.P. had antennae with a three-segmented club as is typical for this genus. Sequences of the mitochondrial cytochrome c oxidase subunits I (CO I) and II (CO II) of a winged male and two nestmate workers showed that they are close to C. venustula and C. shuckardi but differ from them in about 10% of the base pairs. Other samples of Cardiocondyla collected in Comoe N.P. belong to C. venustula and related species. Their males were always ergatoid or “intermorphic” (i.e., ergatoid with rudimentary wings), as previously found in other populations of C. venustula, and never winged with modified antennae.

Population and colony structure of an ant with territorial males, Cardiocondyla venustula

BackgroundMany species of social insects have large-scale mating and dispersal flights and their populations are therefore often relatively homogenous. In contrast, dispersal on the wing appears to be uncommon in most species of the ant genus Cardiocondyla, because its males are wingless and the winged queens mate in their natal nests before dispersing on foot. Here we examine the population structure of C. venustula from South Africa. This species is of particular interest for the analysis of life history evolution in Cardiocondyla, as it occupies a phylogenetic position between tropical species with multi-queen (polygynous) colonies and fighting males and a Palearctic clade with single-queen colonies and mutually peaceful males. Males of C. venustula exhibit an intermediate strategy between lethal fighting and complete tolerance – they mostly engage in non-lethal fights and defend small territories inside their natal nests. We investigated how this reproductive behavior influences colony and population structure by analyzing samples on two geographic scales in South Africa: a small 40 × 40m2 plot and a larger area with distances up to 5 km between sampling sites in Rietvlei Nature Reserve near Pretoria.ResultsColonies were found to be facultatively polygynous and queens appear to mate only with a single male. The extraordinarily high inbreeding coefficient suggests regular sib-mating. Budding by workers and young queens is the predominant mode of colony-founding and leads to high population viscosity. In addition, some queens appear to found colonies independently or through adoption into foreign nests.ConclusionWhile C. venustula resembles tropical Cardiocondyla in queen number and mating frequency, it differs by the absence of winged disperser males. Dispersal by solitary, mated queens on foot or by short flights and their adoption by alien colonies might promote gene flow between colonies and counteract prolonged inbreeding. The abundance of suitable habitat and the high density of nests facilitate the spread of this species by budding and together with the apparent resistance against inbreeding make it a highly successful pioneer species and invader of degraded and man-made habitats.

Interruption points in the wing gene regulatory network underlying wing polyphenism evolved independently in male and female morphs in Cardiocondyla ants.

Wing polyphenism in ants, which produces a winged female queen caste and a wingless female worker caste, evolved approximately 150 million years ago and has been key to the remarkable success of ants. Approximately 20 million years ago, the myrmicine ant genus Cardiocondyla evolved an additional wing polyphenism among males producing two male morphs: wingless males that fight to enhance mating success and winged males that disperse. Here we show that interruption of rudimentary wing-disc development in larvae of the ant Cardiocondyla obscurior occurs further downstream in the network in wingless males as compared with wingless female workers. This pattern is corroborated in C. kagutsuchi, a species from a different clade within the genus, indicating that late interruption of wing development in males is conserved across Cardiocondyla. Therefore, our results show that the novel male wing polyphenism was not developmentally constrained by the pre-existing female wing polyphenism and evolved through independent alteration of interruption points in the wing gene network. Furthermore, a comparison of adult morphological characters in C. obscurior reveals that developmental trajectories lead to similar morphological trait integration between winged and wingless females, but dramatically different integration between winged and wingless males. This suggests that the alternative sex-specific developmental routes to achieve winglessness in the genus Cardiocondyla may have evolved through different selection regimes acting on wingless males and females.

Mandible morphology reflects the type of male antagonism in the ant genus Cardiocondyla (Hymenoptera: Formicidae)

The ant genus Cardiocondyla is characterized by wingless, ergatoid males, which in some species replace the typical winged male. Depending on species, ergatoid males engage in lethal fighting for access to sexual females, establish territories within their nests, or are mutually tolerant. Here we investigate, whether the morphology of ergatoid males and in particular the shape of their mandibles reflect phylogeny or male behaviour. In contrast to the worker mandible, which is relatively similar in shape in the ten species examined, mandibles of ergatoid males show a great variation in size and shape not only between the two clades of Cardiocondyla but also among species belonging to Cardiocondyla clade A. This diversity appears to reflect the diverse reproductive tactics of ergatoid males of different species, with extremely long, anteriorly oriented mandibles associated with indirect killing of adult competitors, while particularly short mandibles possibly constitute an adaptation to a tactic in which ergatoid males crush the cuticle of young rivals.

Genital Morphology of Winged and Wingless Males in the Ant Genus Cardiocondyla (Formicidae, Myrmicinae)

The myrmicine ant genusCardiocondylais characterized by a peculiar male diphenism with winged disperser males and wingless, “ergatoid” fighter males. Here we describe and illustrate the morphology of the male external genitalia of 13 species of this genus. Several characters, especially the parossiculus of the volsella and the medial face of the paramere, vary between different species groups and might be useful to clarify the infrageneric taxonomy ofCardiocondyla. External genitalia between different phenotypes of a species differ primarily in size. All examined males, both winged and ergatoid, have a parameral hook that resembles a structure described inNesomyrmexmales but apparently is absent in other studied myrmicine ants. The presence and absence of this hook might be useful for future studies on the relationships within the myrmicine tribe Crematogastrini.

Mating and longevity in ant males.

Across multicellular organisms, the costs of reproduction and self‐maintenance result in a life history trade‐off between fecundity and longevity. Queens of perennial social Hymenoptera are both highly fertile and long‐lived, and thus, this fundamental trade‐off is lacking. Whether social insect males similarly evade the fecundity/longevity trade‐off remains largely unstudied. Wingless males of the ant genus Cardiocondyla stay in their natal colonies throughout their relatively long lives and mate with multiple female sexuals. Here, we show that Cardiocondyla obscurior males that were allowed to mate with large numbers of female sexuals had a shortened life span compared to males that mated at a low frequency or virgin males. Although frequent mating negatively affects longevity, males clearly benefit from a “live fast, die young strategy” by inseminating as many female sexuals as possible at a cost to their own survival.

First come, first served: the first-emerging queen monopolizes reproduction in the ant Cardiocondyla “argyrotricha”

While colonies of most tropical species of the ant genus Cardiocondyla regularly contain multiple egg-laying queens (polygyny), single-queening (monogyny) evolved convergently in a Palearctic clade of Cardiocondyla and in Southeast Asian C. “argyrotricha.” In the latter species, monogyny is probably an adaptation to patchily distributed but highly stable nest sites. In experimentally orphaned colonies of C. “argyrotricha,” the first emerging queen shed its wings, began to lay eggs, and stayed mostly on the brood pile. Queens that emerged later remained in the peripheral areas of the nest without dispersing but retained their wings even after mating in the nest. Aggressive interactions among queens and between workers and queens were occasionally observed, but it appears that the order of queen emergence determines which of them will become reproductive and inherit the nest. We conclude that young queens commonly compete for nest inheritance in some species of Cardiocondyla and that queen–queen antagonism does not necessarily involve lethal fighting.

Massacre of the innocents in Cardiocondyla thoracica: Manipulation by adult ant males incites workers to kill their immature rivals

AbstractWingless males of the ant genus Cardiocondyla exhibit a remarkable range of intrasexual competition, from mutual tolerance among males in the colonies of some species to regular lethal fighting in the nests of others. Here, we describe male–male antagonism in Cardiocondyla thoracica (Smith, 1859) from Northern Australia. Males appear to widely ignore adult competitors, but grab defenseless, young rivals during or immediately after emergence and daub them with hindgut secretion. This fluid elicits fatal worker aggression against the besmeared individual. Besmearing and subsequent worker aggression is known from other species of Cardiocondyla but is usually directed towards adult contenders, whereas males kill freshly eclosing males by biting or dismembering. The behavior of C. thoracica males thus constitutes a novel nuance of rival elimination.

Cryptic Lineages in the Cardiocondyla sl. kagutsuchi Terayama (Hymenoptera: Formicidae) Discovered by Phylogenetic and Morphological Approaches

The taxonomy of ant species in the genus Cardiocondyla is very confused due to the extreme difficulty in separating many species based on morphology alone. In Japan, one group of the species complex Cardiocondyla sl. kagutsuchi has both winged and wingless worker-like (ergatoid) males (dimorphic) whereas others have only ergatoid males (monomorphic). The presence of both groups prompted us to hypothesize that C. sl. kagutsuchi presumably includes several independent species with differences in their male wing morphologies. However, whether any species boundary actually exists between the male groups has remained unsolved over the 10+ years since the previous revision of this genus. In this study, using discriminant and phylogenetic analyses, we compared the worker caste morphology of this species complex among lineages detected by phylogenetic analyses. In addition, we examined the number of sexuals present in field colonies. Our results revealed the existence of at least three morphological and phylogenetic groups within this species complex.

Polygyny, Inbreeding, and Wingless Males in the Malagasy Ant Cardiocondyla shuckardi Forel (Hymenoptera, Formicidae)

The ant genus Cardiocondyla exhibits a fascinating diversity of its reproductive biology, with winged and wingless males, long-winged and short-winged queens, strict monogyny and facultative polygyny with or without queen fighting. Here we report on the previously unstudied Malagasy ant C. shuckardi . We describe the nesting habits, male morphology and colony structure of this species. Furthermore, based on the genotypes from three microsatellite loci we document a very high incidence of sib-mating.

Virgin ant queens mate with their own sons to avoid failure at colony foundation

Mother-son mating (oedipal mating) is practically non-existent in social Hymenoptera, as queens typically avoid inbreeding, mate only early in life and do not mate again after having begun to lay eggs. In the ant genus Cardiocondyla mating occurs among sib in the natal nests. Sex ratios are extremely female-biased and young queens face the risk of remaining without mating partners. Here, we show that virgin queens of Cardiocondyla argyrotricha produce sons from their own unfertilized eggs and later mate with them to produce female offspring from fertilized eggs. Oedipal mating may allow C. argyrotricha queens to found new colonies when no mating partners are available and thus maintains their unusual life history combining monogyny, mating in the nest, and low male production. Our result indicates that a trait that sporadically occurs in solitary haplodiploid animals may evolve also in social Hymenoptera under appropriate ecological and social conditions.

Queen life-span and total reproductive success are positively associated in the ant Cardiocondyla cf. kagutsuchi

While reproduction and longevity are negatively correlated in a large number of animals, this fundamental trade-off appears to be absent from social insect queens. Here, we examine the distribution of the life-spans of queens and their mates and the age-trajectory of reproduction in experimental colonies of the Pacific tramp ant Cardiocondyla cf. kagutsuchi. Queens of Cardiocondyla are generally short-lived (0.5–2 years), which allows determining their lifetime reproductive success and establishing mortality tables. We show that the queen’s total number of sexual offspring is positively associated with its life-span. Regular counts of eggs, larvae, and pupae suggest that more fecund queens live longer than less fecund queens and that an early onset of sexual production does not negatively affect the queen’s life-span. This corroborates the view that the costs of reproduction are not born by the queen itself. The number of eggs present in colonies increased with queen’s age until shortly before death, indicating negligible reproductive senescence. Sex ratios were strongly affected by the origin of queens but neither by colony size nor total productivity. Several queens produced only males late in their lives, suggesting the occurrence of sperm depletion. Male life-span was not correlated with any other studied trait. Our study shows that the ant genus Cardiocondyla provides a suitable model system to investigate the evolution of life-span in social insect queens and males.

Cardiocondyla pirata sp. n. – a new Philippine ant with enigmatic pigmentation pattern (Hymenoptera, Formicidae)

A new species of the ant genus Cardiocondyla Emery, 1869 – Cardiocondyla pirata sp. n. – is described from the Philippines. The species belongs to an Indo-Malayan group of six species that is characterized by workers having a strongly bilobate postpetiolar sternite and a thickset mesosoma with strongly convex dorsal profile as well as wingless, ergatoid males with sickle-shaped mandibles. The female castes show a pigmentation pattern not known from any ant worldwide. If having any adaptive value, a possible function of this structure is supposed to be visual dissolution of body shape in order to irritate predators.