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Systematics of the Mesynodites sensu lato genus complex

(Coleoptera: Histeridae)

Alexey K. Tishechkin and Christopher E. Carlton

A project supported by the National Science Foundation

PRODUCTS TO DATE

Carlton, C. E., M. Dean, & A. K. Tishechkin. Diversity of two beetle taxa at a western Amazonian Locality (Staphylinidae: Pselaphinae and Histeridae: Hetaeriinae).

Carlton, C. E., & M. Dean. New species of Barroeuplectoides from Central and South America (Coleoptera: Staphylinidae: Pselaphinae).

PROJECT SUMMARY

The Histeridae is a diverse family of approximately 4,000 mainly predatory beetle species that occupy a wide variety of ecological niches. All species within the subfamily Hetaeriinae are guests of ant or termite colonies and have evolved in association with a wide variety of hosts. This subfamily is one of the most diverse myrmecophilous and termitophilous monophyletic insect taxa, with approximately 315 described species. The proposed research will result in a systematic revision of the ant-associated genus complex Mesynodites sensu lato, with about 55 described species and at least twice that number of undescribed species. This project will lead to a mongraphic treatment of Mesynodites. Important steps towards its successful completion include fieldwork in poorly known areas of the neotropics and visits to museums that house major collections of hetaeriines and other inquilinous taxa.

Cladistics-based systematic treatment at the species and genus levels within Mesynodites will provide information that is essential to gaining insight into the phylogenetic relationships of the subfamily Hetaeriinae. Species within Mesynodites are associated with an unusually wide range of hosts for a single genus, including numerous species of army ants and leafcutter ants. The genus is polyphyletic and consists of a mixture of lineages with unclear interrelations, so this apparent wide host range probably is an artifact of poor phylogenetic resolution. A proper understanding of guest-host coevolution within the Hetaeriinae cannot be achieved until this polyphyletic assemblage has been resolved into component monophyletic genera. By identifying the component lineages within Mesynodites and superimposing their ant hosts onto the obtained phylogeny, we will analyze patterns of evolution that have resulted in the adaptation of species to their respective social insect hosts. Social insect inquilinism has evolved independently in many different lineages of insects, so this study will provide information that is important in discovering general patterns in the evolution of behavioral and morphological adaptations across a broad range of inquilinous taxa and hosts.

We will bring together all distributional and host information on the genus complex and make it available to interested researchers via this project website while we conduct systematic and coevolutionary analyses. This database will be important for taxonomic and biodiversity studies, and may also be used as a source of information by researchers conducting studies of the evolution of ant and termite associations in beetles. Field expeditions to poorly collected regions of South America will be conducted to procure specimens for phylogenetic study and to fill gaps in distributional and host data. Additional inquilinous insects and their host data will be collected during the fieldwork and made available for the systematics community. In addition to currently used mass-collecting techniques, we will employ observational and hand collecting methods that were successfully used by early 20th century naturalists to obtain most of the data we currently possess about the host associations and behaviors of many inquilinous species.

OBJECTIVES

We propose a monographic study of the genus Mesynodites s.l. and closely related genera of neotropical Hetaeriinae (Coleoptera: Histeridae) with the following goals:

1. Revision of all material available in collections Worldwide for validation of described species-level taxa, recognition of synonyms, and description of new species.

2. Phylogenetic cladistic analysis of all species under consideration to elucidate monophyletic species-group lineages and develop modern generic concepts for the included species.

3. Preparation of identification keys for all the genera and species treated.

4. Analysis of the host associations within phylogenetic lineages of Mesynodites s.l. and allies with attention to patterns of guest evolution and host-guest co-evolution.

5. Accumulation and dissemination of data on host associations and geographical distribution of neotropical Hetaeriinae and other collateral contributions related to beetles from targeted habitats. .

INTRODUCTION AND BACKGROUND

The beetle family Histeridae includes about 4,000 described species distributed worldwide with the exception of Arctic and Antarctic regions (Mazur 1997). Histerids are small to medium-sized beetles (0.7-25 mm) and occur in many habitats, from dense forests to deserts and dunes, where most are predators of beetle, fly and flea larvae. They live in a variety of decomposing organic materials (dung, carrion, rotten wood, seaweed, and forest litter), under loose bark of woody plants, in galleries of wood-boring insects, vertebrate nests, rhizospheres of desert plants, and in nests of social insects (ants and termites). There are also several specialized soil and cave-dwelling species (Kryzhanovkij & Reichardt, 1976; DÈgallier & Gomy, 1983).

Many histerids successfully exploit social insects (Kistner, 1982) living in their nests and relying on their hosts for food and protection. Numerous and diverse morphological and behavioral adaptations have evolved within the Histeridae, especially among ant and termite guests (Reichensperger, 1926; Akre, 1968). Our research focuses on a systematic revision of the ant-associated genus Mesynodites sensu lato, which includes about 55 described species and is the largest New World genus of the completely inquilinous (social insect-associated) subfamily Hetaeriinae. The proposed study will include systematic treatment at the species and genus levels and accumulation and analysis of zoogeographic information and host data. This research will fill an important gap in understanding phylogenetic relationships and evolution of host use patterns within the subfamily and will contribute considerably to understanding the relationships among subfamilies of Histeridae.

The Hetaeriinae is a diverse, almost exclusively neotropical subfamily, containing about one-third of the World and two-thirds of neotropical histerid genera and about 20% of all described neotropical species (Mazur, 1997). Approximately 100 genera with 240 described species occur in the neotropics. In addition, the subfamily contains 1 Holarctic genus with 26 species in North America and six in the Palearctic, and three nearly exclusively Mediterranean genera containing 49 species. Four genera are shared between neotropical and nearctic regions (Mazur, 1997; YÈlamos, 1997). At the present level of knowledge, the striking diversity of neotropical hetaeriine genera is characterized by the large proportion of genera containing few or single species. Only 8% of hetaeriine genera contain four or more species. Indeed, about 65% are monotypic (Mazur, 1997). The target of the proposed research, the genus complex Mesynodites s. l. (includes Mesynodites s.str., Alloiodites, Daitrosister, Metasynodites and Monotonodites), represents a unique situation within the subfamily because it contains 55 described species (Mazur, 1984). The next largest genus within the Hetaeriinae contains 13 species (Mazur, 1997).

Important descriptive and revisionary work on the Hetaeriinae was performed by Borgmeier (1948), Bruch (1923, 1926, 1933), and Reichensperger (1924, 1935a, 1938, 1939, 1958), but no phylogenetic analyses were attempted until Helava et al. (1985) conducted the first revision of New World Hetaeriinae. In this work, 77 genera were included (mainly one species per genus) and keyed, the male genitalia were used as diagnostic characters for the first time for the subfamily, and a phylogeny of the included genera was generated using parsimony analysis. Helava et al. (1985) demonstrated that the Hetaeriinae is a monophyletic taxon with five distinct supra-generic lineages (Fig. 1). Unfortunately, this study has several weak points. First, the authors did not study all the genera. Even within the genera studied, there were no attempts to examine material deposited outside North American institutions. Recent collections containing many undescribed neotropical taxa have underscored the inadequacy of the sample upon which their conclusions were based. As a result, the key and the phylogeny provided are of limited application. Further, no published works, or those currently being completed, deal with Mesynodites sensu lato (Helava et al., 1985; DÈgallier, pers. comm.), the largest genus of the subfamily (Lewis, 1893; Schmidt, 1893; Bruch, 1923, 1926, 1933; Reichensperger, 1923, 1924, 1925, 1931, 1933, 1935 a, b, 1938, 1939; Mann, 1925). Helava et al. (1985) discussed the complexity and probable polyphyly of the genus, made several taxonomic re-arrangements and illustrated male genitalia of 4 species. DÈgallier (pers. comm.) will only redescribe the type species of Mesynodites, M. schuppi (Schmidt), and will include only this species and other genera within Mesynodites s.l. into his phylogenetic analysis.

The coevolutionary aspect of this project involves analysis of the host associations of Mesynodites s. l. and related hetaeriine genera. The oldest representatives of recent genera of Hetaeriinae and army ants, a group that includes many host ant species, are from middle Tertiary Dominican amber (Wilson, 1985; P.Kovarik, pers. comm.). Termites and ants apparently originated in middle to lower Cretaceous (Krishna & Weesner, 1970; Carpenter & Hermann, 1979; Agosti et al., 1997). Because hetaeriines are obligate inquilines of social insects, the long association of both guests and hosts, presumably dating at least from the late Cretaceous or early Tertiary, provides an ideal system to study a broad range of coevolutioary patterns and processes. Complete integration into social insect colonies by inquilinous guests provides greater access to food and protection from predation (Kistner, 1979). But social insects have sophisticated recognition systems and treat uninvited guests with extreme hostility. So this integration required “breaking the code of social insects” (Wilson, 1971: 410). Mechanisms for accomplishing this have included the use of appeasement and adoption chemicals, behavioral cues, and tactile (or Wasmannian) mimicry, or their combinations (Kistner, 1979). Such host-guest systems might be driven, at least in part, by continually evolving patterns of host and guest diversification, co-speciation, host shifts, and evolutionary escalation (Futuyma & Slatkin, 1983; Mitter & Farrell, 1991; Vermeij, 1994).

Most information about patterns of host relationships of predatory inquilinous beetles is based on studies of the Staphylinidae, a family that contains a large number of inquilinous lineages associated with a wide variety of host taxa (Kistner, 1979, and references therein; Kistner & Jacobsen, 1990; Jacobsen & Kistner, 1991). Both myrmecophilous and termitophilous species exhibit a high degree of host specificity. In general, host specificity is higher among termitophiles, which often are restricted to a particular host species. Myrmecophiles are typically restricted to particular ant genera and may be found with several host species. As pointed out by Kistner (1979: 378), some apparent host specificity may be the result of sample bias rather than real pattern.

Phylogenetic analysis of Hetaeriinae genera will provide insight into the evolution of ant and termite host relationships. These genera display a wide spectrum of degrees of integration into host colonies. In Wasmann's (1903) classification they range from synechtrans (persecuted guests) to symphiles (welcome guests). In Kistner's (1979) classification poorly specialized non-integrated to completely integrated species are represented in the Hetaeriinae. Also, diverse host taxa are represented among the hetaeriines. Host termite genera in the Neotropics include Cornitermes, Nasutitermes, and Syntermes (Nasutitermitidae). Ant genera are more numerous as hosts of hetaeriinies, including Eciton, Labidus, Neivamyrmex, and Nomamyrmex within the Ecitoninae (army ants); Acromyrmex, Atta, Pheidole, and Solenopsis within the Myrmicinae, and Pachycondyla within the Ponerinae (Reichensperger, 1926; Helava et al., 1985). Army ants harbor the most diverse hetaeriine communities. Host specificity at the generic level is very high in the neotropical Hetariinae, with only one genus reported from more than one host genus (Helava et al., 1985). Ant genera that serve as hosts of hetaeriines outside the Neotropics include Tapinoma within the Dolichoderinae, Acantholepis, Formica and Lasius within the Formicinae, and Aphaenogaster and Tetramorium within the Myrmicinae (Kryzhanovskij & Reichardt, 1976; Helava et al., 1985; YÈlamos, 1992, 1995, 1997).

Several points are important in approaching questions related to inquilinous Hetaeriinae. First, they exhibit unusual and poorly understood morphological adaptations, but lack the adaptations most characteristic of inquilinous staphylinids such as physogastric and limuloid body shapes (Kistner, 1979, 1982). Second, existing data on host-guest associations and coevolution are rather poor, due to a paucity of phylogenetic information on Hetaeriinae (Helava et al., 1985), and data about the host range and specificity of hetaeriine species are incomplete. In fact, most information currently available about host relationships among hetaeriine species was gathered by 19th and early 20th century naturalists through direct collecting from host nests. Third, a hypothesis to explain the evolution of host associations was developed by Helava et al. (1985). They proposed that ancestors of the Hetaeriinae first invaded ant refuse deposits, then invaded ant and termite colonies. They predicted that sister taxa of the hetaeriines would be found among “facultative predators” in these deposits. Thus, the gradient from “facultative predators” to “symphiles” should be characterized by increases in guest species diversity and degrees of morphological and behavioral adaptations, and in host specificity.

Two obvious trends within the Hetaeriinae are rapid guest evolution and utilization of host lineages by multiple lineages of guests. Within coevolving monophyletic host-guest lineages there are many more species and genera of guests than hosts. Patchy distribution of colonies, rapid adaptation to host defenses, and low initial size of invading populations combined with limited genetic interchange with parent populations all may contribute to relatively rapid evolutionary changes in guests (Kistner, 1979; Helava et al., 1985). Also, a given host lineage may be utilized by several different lineages of guests. Multiple guest invasions within host lineages, host switches, including switches between ants and termites, have been hypothesized to explain these observed patterns (Seevers, 1965; Kistner, 1979; Helava et al., 1985; Jacobsen & Kistner, 1991). “Diffuse coevolution”, when host and guest clades have had a long but rather loose association and most shifts are between closely related hosts (Mitter et al., 1991), may account for these patterns of relationship. An analysis of such patterns within the Mesynodites genus complex is an important objective of the proposed research.

Determination of host relationships of Mesynodites s. l. and related genera is a model project that will set the stage for future research to analyze the guest-host relationships for the entire subfamily Hetaeriinae. The proposed project will allow us to test some of the assumptions and predictions of previous workers, and develop a more precise model of hetaeriine-host coevolution. Host-guest data are scattered through numerous publications (cited above) and museum specimen labels, some of which were summarized by Helava et al. (1985). All New World genera of army ants (Ecitoninae) except Cheliomyrmex are utilized as hosts by Mesynodites s.l. species. The general picture is the following: among species of Mesynodites s.l. with known host associations, 10-15 species are reportedly associated with Eciton, Labidus, and Neivamyrmex, six with Nomamyrmex, and four with Atta. Specialization at the level of host genera seems to be the typical pattern for Mesynodites s.l. species. Only five guest species (about 10%) are associated with two host genera.

Several important generalizations on these host-guest relations can be made. First, there are host relationships within Mesynodites s.l. with 2 distantly related ant taxa, Ecitoninae and Attini. Whether use of these two ant taxa as hosts occurs in discreet or scattered lineages can be tested phylogenetically. Second, ecitonine genera which have Mesynodites s.l. guests differ in their habits with respect to surface vs litter and diurnal vs nocturnal activities, and development of raiding behavior (Gottwald, 1982). These differences provide possibilities for divergent evolutionary trajectories within lineages associated with each ant genus. Third, no Mesynodites s.l. were found in Cheliomyrmex (no histerids or staphylinids at all were reported for this genus). This genus is considered to represent a separate lineage from the other three ecitonine genera and even to be a link between New and Old World army/driver ants (Wheeler, 1928; Gottwald, 1982). Searching for and discovering (or proving the absence of) hetaeriine guests in the nests of this genus would be an important contribution to understanding the evolution of Hetaeriinae. Resolving the Mesynodites s.l. phylogeny also may contribute to an understanding of relationships between army ant genera by optimizing host and guest phylogenies. All of the recent ant phylogenetic studies (Bolton, 1990; Baroni-Urbani et al., 1992; Shattuck, 1992) placed Ecitoninae among the most derived ants. But, none of these studies provided data on relationships among the genera. An attempt to incorporate phylogenetic information on staphylinid guests into predictions of relationships among army ant genera was recently done by Kistner and Jacobsen (1990), and the analysis of the Mesynodites s. l. host associations will provide a test of their conclusions and will contribute to a more complete understanding of host-ant generic relationships.

Geographical distribution of Mesynodites s.l. as well as most other New World hetaeriine genera is within the neotropical biogeographic realm from southern Mexico to northern Argentina, southern Brazil and Uruguay, but patterns of distribution within this region are unclear. The most substantial material on the subfamily from Central and South America was accumulated in 1920-30s by Bruch, Reichensperger and their local co-workers by direct collecting in the ant and termite nests in a few localities in the Argentinian provinces of Buenos Aires, Cordob·, and Misiones; Brazilian states of Goi·s, Paran·, Rio de Janeiro, and Santa Catarina; and Costa Rican provinces of San Jose and Limon. Since the 1950's, direct collecting from hetaeriines has been conducted in only one additional area, Barro Colorado Island in Panama (e.g., Akre, 1968; Akre & Rettenmeyer, 1968). Introduction of flight intercept traps (FIT's) in the 1970s (Peck & Davies, 1980) and their regular use in the neotropics since then has resulted in the accumulation of many hetaeriine specimens, particularly from Costa Rica and Panama. Modest samples from flight intercept traps and litter collecting in the 1980-90s are available from Argentina, Belize, Brazil, Ecuador, Mexico, Paraguay, and Peru. Few or single specimens/species are available from Bolivia, Guatemala, Guianas, Honduras, and Venezuela.

Two existing collection biases are particularly important. First, few hetaeriines have been collected in vast areas of the rainforests in the Amazon Basin which are considered to harbor the highest species diversity in the Neotropics (e.g., Prance, 1982). Small collections from eastern Ecuador and Peru are unique to those areas (Tishechkin, unpubl.), but are obviously insufficient. Information is lacking about the number of species of Hetaeriinae that occupy large ranges, and some records of apparently widely distributed species appear to be mistakes (e.g., DÈgallier, 1994). At the present level of knowledge, hetaeriine species diversity appears to be richest in peripheral areas of the tropics and even some subtropical areas. Second, abandonment of direct host nest collecting has resulted in a serious lack of data on host associations in hetaeriines, especially at the species level and across broad geographic scales (Helava et al., 1985). Thus, additional careful collecting in specific areas and from host nests, combined with accumulation, analysis and dissemination of precise species/host/distribution information is of considerable importance for fulfilling broad research goals dealing with the Hetaeriinae and promises to contribute substantially to our knowledge of inquilinous insects in general.

Inquilinous beetles of LaSelva, Costa Rica

Report on Fieldwork at Yasuni National Park, Ecuador

Histeridae of Louisiana

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