Termitariophily: expanding the concept of termitophily in a physogastric rove beetle (Coleoptera: Staphylinidae)

1. Termitophily in some rove beetles is commonly attributed to the striking termite worker resemblance that is provided by the beetles' hypertrophic (‘physogastric’) abdomen. However, a termite nest may offer to a termitophile additional benefits, such as a continuously repaired shelter. 2. This could apply to Corotoca melantho (Aleocharinae: Corotocini), a viviparous obligatory termitophile staphylinid beetle species. While conferring morphological congruence to its host worker termites, its physogastry may impair mobility, leading to vulnerability and the need for a secure environment. It seems plausible to hypothesise that physogastry in C. melantho would imply in interactions between this termitophile and its host termites as well as its host termitarium. 3. This study provides evidence to build such a hypothesis by inspecting the morpho‐anatomical reproductive traits of this termitophile. It was found that a gradient of growth stages of embryos and larvae in the oviduct explains physogastry in females while pointing to iteroparity. The asynchronous development of oocytes in females, combined with a full developing sequence of sperm cells indicative of continuous spermatogenesis in males, suggests frequent matings. 4. While improving guest–host similarity, physogastry and flightlessness should confer vulnerability to pregnant females, forcing C. melantho to seek close and sheltered environments. These could facilitate the frequent male–female contacts demanded by iteroparity. It is possible, therefore, that physogastry in C. melantho is not only associated with the termites themselves but also with the physical structure of the termitarium. Thus, C. melantho can be hypothesised to be a termitariophile in addition to being a termitophile.     

The rove beetle Drusilla sparsa (Coleoptera: Staphylinidae) is a myrmecophilous species associated with a myrmicine ant,Crematogaster osakensis (Hymenoptera: Formicidae)

The rove beetle genus Drusilla includes some myrmecophilous species. The Japanese species Drusilla sparsa (Sharp, 1874) has been regarded as a non‐myrmecophilous beetle. In Kagawa Prefecture, Shikoku Island, western Japan, however, we often observed that D. sparsa adults were walking in the vicinity of foraging workers of the myrmicine ant Crematogaster osakensis Forel, 1990. The body color of the beetle is similar to C. osakensis as in other myrmecophilous beetles found near the trails of the host ants. To examine whether D. sparsa is myrmecophilous, we investigated the distribution of D. sparsa and C. osakensis in the field, as well as their behavior including prey preference of the beetle in the laboratory. Drusilla sparsa beetles were collected only in sites where C. osakensis ants occurred. When the beetles encountered the ant workers, they bent the abdominal tip toward the ants. The ants licked the abdominal tip, and then the beetles usually walked away. Such behavioral reaction of the ants was not observed when the beetles encountered workers of the formicine ant Nylanderia flavipes (Smith, 1874) that continuously attacked the beetles. Drusilla sparsa preferred to feed on dead workers of C. osakensis even when other ants were available as food, indicating that D. sparsa is a myrmecophilous species associated with C. osakensis. Crematogaster osakensis was frequently found in the stomach in the ant predator, the Japanese treefrog Hyla japonica Günther, 1859. Thus, the significance of body color similarity between the host ants and beetles is not a case of Batesian mimicry.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

Multilocus phylogeny defines a new classification of Staphylininae (Coleoptera,Staphylinidae), a rove beetle group with high lineage diversity

We provide the first multilocus molecular phylogeny of a group corresponding to the former subfamily Staphylininae. Results are corroborated by the morphological, biogeographical and palaeobiological evidence to serve as a baseline for an updated suprageneric classification. The former subfamily Staphylininae is proven to be a lineage sister to the monophyletic Paederinae and reclassified according to a robust phylogeny resolving a number of long-standing controversies. The subfamily Xantholininae (revised status) is reinstated to contain the tribes Xantholinini, Othiini, Maorothiini and Diochini. Subfamily Platyprosopinae (revised status) is reinstated for the tribes Platyprosopinini, Arrowinini and †Thayeralinini. For a highly peculiar genus Coomania Cameron, formerly in Diochini, a new subfamily Coomaniinae subfam.n. is established and the composition of Diochini (revised status) is changed accordingly. The subfamily Staphylininae (revised status) is reduced to contain the former tribe Staphylinini only. Elevating this mega-diverse tribe to the subfamily rank opened up an opportunity for its more fractional classification by raising several subtribes to the tribal level as follows: Acylophorini, Afroquediini, Amblyopinini, Antimerini, †Baltognathini, Cyrtoquediini, Erichsoniini, Hyptiomini, Indoquediini, Quediini and Tanygnathinini (revised status for all). As a result, the most species-rich tribe Staphylinini (revised status) is reduced to the more homogeneous lineage containing the subtribes Algonina, Anisolinina, Philonthina, Philothalpina, Staphylinina and Xanthopygina. Morphological synapomorphies and diagnostic characters supporting all newly defined higher taxa are provided. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:DED8B042-83C9-4D10-B0CB-B50372B067A9 .     

Structure and energy pathways of spermatozoa of the rove beetle Aleochara bilineata (Coleoptera, Staphylinidae)

The morphology of mature spermatozoa of the rove beetle Aleochara bilineata was examined by using scanning and transmission electron microscopy. They are about 1000 mum long and filiform. The acrosome and the nucleus are elongate and each about 20 mum long. A well-developed centriole adjunct region connects the nucleus with the sperm tail. The axoneme reveals the 9 + 9 + 2 pattern of the pterygote sperm flagellum. Two accessory bodies and two mitochondrial derivatives with paracrystalline inclusions are present. Cristae are reduced to the cortical zone of the derivatives. Cytochrome-c oxidase activity was detected within the cristae by DAB-reaction. The energy metabolism of the spermatozoa was investigated by using different inhibitors affecting the mitochondrial and cytoplasmic metabolic pathways. Sperm movement was used as an indicator for the utilization of ATP by the axoneme. In control experiments, the duration of motility was longer than 45 min. In the presence of atractyloside or potassium cyanide the motility duration was not affected. On the other hand, iodoacetic acid in the medium stopped sperm motility within 15 min. This indicates that sperm energy metabolism mainly depends on the glycolytic pathway.     

Food preference of the rove beetle, Atheta coriaria Kraatz （Coleoptera： Staphylinidae） under laboratory conditions

A study, involving laboratory choice tests, was conducted to determine the feeding behavior, based on food preference, of the adult and larval stage of the rove beetle, Atheta coriaria Kraatz when presented with both fresh moistened oatmeal and second instar fungus gnat, Bradysia sp. nr. coprophila （Lintner） larvae in Petri dishes. Rove beetles used in this study came from a laboratory-reared colony. A rating scale from 1 to 5, based on percent missing （1 = 0 to 10%, 2 = 11 to 30%, 3 = 31 to 50%, 4 = 51 to 75%, and 5 = 76 to 100%）, was used to objectively assess the amount of oatmeal and number of fungus gnat larvae consumed by each rove beetle adult and larva. In all the choice tests, A. coriaria adults and larvae preferred to feed on fungus gnat larvae （78% and 69%, respectively） significantly more so than oatmeal （9% and 5%, respectively） based on the amount of oatmeal and number of fungus gnat larvae consumed after 4 and 6 hours. There were relatively minimal differences in the amount of food consumed for both adults and larvae after 4 and 6 hours. The results of this study indicate that oatmeal may be an inexpensive supplemental food source, during the rearing process, which will not inhibit the effectiveness of rove beetles to control fungus gnat larvae when released into greenhouses.     

Biology and distribution of the littoral rove beetle Psamathobledius punctatissimus (Le Conte) (Coleoptera: Staphylinidae)

The intertidal macrofauna on a small sheltered marine beach at Pawley's Island, South Carolina, was dominated by insects, primarily Psamathobledius punctatissimus, a small beetle which attained densities of up to 2 260 adults m⁻². Both adult and larval beetles remained buried in the sand when covered by the tide, then emerged to form mole-like surface trails and feed upon diatoms when exposed. Eggs and young larvae were maintained within special maternal burrows, while larger larvae and non-breeding adults occupied their own individual burrows. Both adults and larvae became comatose when in direct contact with water, but recovered after as long as six hours submergence. In the field, burrows retained air during tidal coverage, allowing the beetles to avoid direct contact with sea water. Seaward extension of the population is probably limited by time available for feeding, while longshore distribution is restricted by sand texture and diatom concentration, or by sediment mobility in more exposed locations. Although very locally distributed, P. punctatissimus has achieved considerable success in a habitat poorly exploited by competitors of either marine or terrestrial origin. Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research     

The ecology of the marine rove beetle,Bryothinusa spp. (Coleoptera: Staphylinidae) in Hong Kong

Bryothinusa spp. are common marine insect in Hong Kong. They occur in the intertidal zone of sandy shores between 0.6 and 1.2 m tide level. They emerge when the tide recedes, possibly to mate and feed, then burrow again at the advance of the incoming tide.Bryothinusa has a special respiratory apparatus for long submersions and therefore is able to inhabit the changeable intertidal zone where the competition with other living organisms is comparatively low.     

Morphology and ultrastructure of paired prototergal glands in the adult rove beetle Philonthus varians (Coleoptera, Staphylinidae)

Philonthus and other genera of Philonthina possess a pair of prototergal glands located in the first abdominal tergum and hidden at rest by hind wings and elytra. In Philonthus varians they occupy the whole length of the tergum and form a pouch-like invaginated reservoir with a scaly glandular zone and a smooth outlet. A grille of long setae covers the opening of each gland. The fine structure of these glands is given for the first time. Three types of cells are found in the glandular epithelium. Epidermal cells underlie the cuticular scales, numerous class 1 secretory cells open in the centre of calyces made of finger-like processes of the cuticle, and class 3 cells are connected to pored tubercles. A cytological comparison is made with the diverse class 1 cells described to date in Coleoptera. In these cells different evolutionary trends are shown in the structure of the cuticular apparatus, particularly in the number, size and position of the cuticular apertures as well as in the length and abundance of epicuticular filaments. A possible defensive function of the prototergal glands against pathogens and their interest for the phylogenetic study of Staphylininae are discussed.     

Diversity of pselaphine beetles (Coleoptera: Staphylinidae: Pselaphinae) in eastern Thailand

Pselaphine beetles (Coleoptera: Staphylinidae: Pselaphinae) are cosmopolitan, species‐rich, and yet poorly studied, particularly in the tropics. We sampled beetles in three types of primary forest and two types of disturbed forest habitats in eastern Thailand to assess the utility of pselaphine beetles as bioindicators of forest disturbance. We simultaneously measured leaf litter mass, soil moisture, soil acidity and canopy cover at each site to infer which environmental factors affect pselaphine beetle diversity and abundance. At each site, pselaphine beetles were extracted from ten 1 m² samples of leaf litter and soil with Tullgren funnels. We sampled 1867 adult beetles representing six supertribes, 51 genera and 114 morphospecies; 7% of the genera and 92% of the species were undescribed. Forest types differed significantly in species richness, abundance, diversity and evenness. Primary forest had greater numbers of species and individuals, and higher diversity indices (H′). Teak plantation and secondary forest had substantially fewer individuals and species of pselaphine beetles. Species composition differed between primary and degraded forests. Canopy cover, soil moisture, and leaf litter mass positively correlated with beetle species richness and abundance. Leaf litter mass and soil moisture were the two most important factors affecting the diversity of pselaphine beetle assemblages. Among the 114 morphospecies collected, 43 morphospecies were specific to two or three habitats and 64 morphospecies were found only in a single habitat. Thus pselaphine beetles appear to have rather narrow habitat requirements and their presence/absence was correlated with environmental differences. These traits make pselaphine beetles a suitable bioindicator taxon for assessing forest litter diversity and monitoring habitat change.     

Species diversity and community structure of rove beetles (Coleoptera: Staphylinidae) attracted to dung of sika deer in coniferous forests of southwest Japan

Insect communities of mammal dung have been known as excellent model ecosystems for scientific study. Ecological surveys of diversity and seasonal patterns of coprophilous rove beetles in relation to wild mammals have rarely been conducted, although the high potential species diversity and abundance of the rove beetles are known. In order to investigate biodiversity of these beetles, we analyzed species composition, abundance, feeding guild and seasonality of rove beetles that were attracted to sika deer Cervus nippon dung by using dung‐baited pitfall traps for a 1.5‐year study in two plantations (cypress, cedar) and one secondary natural forest (pine) in Fukuoka Prefecture, southwest Japan. Consequently, saprophagous Anotylus sp. (Oxytelinae) was dominant in all forests. Analyses of feeding guild structure showed the number of individuals were dominated by saprophagous beetles, but the number of species were dominated by predatory beetles. Seasonal effects suggested that the species richness and abundance of rove beetles are possibly regulated by scarabaeoid dung beetles. These findings feature one example of a coprophilous rove beetle community.     

Hollow oaks and beetle functional diversity: Significance of surroundings extends beyond taxonomy

Veteran hollow oaks (Quercus spp.) are keystone structures hosting high insect diversity but are declining in numbers due to intensification of land use and the abandonment of traditional management. The loss of this vital habitat is resulting in a reduction of biodiversity, and this likely has consequences for ecosystem functioning, especially if functional diversity is reduced. A considerable amount of research has been done on predictors of beetle taxonomic diversity in veteran oaks, but predictors of functional diversity have remained largely unexplored. The aim of this study was to establish whether the features and surroundings of veteran oaks are related to functional diversity within three functional groups of beetles (decomposers, predators, and flower visitors) and determine whether species richness and functional diversity within the groups are dependent on the same predictors. Sampling was carried out intermittently between 2004 and 2011 on 61 veteran oaks in Southern Norway. Of the 876 beetle species that were collected, 359 were determined to be decomposers, 284 were predators, and 85 were flower visitors. Species richness and functional diversity in all groups were consistently higher in traps mounted on veteran oaks in forests than in open landscapes. However, additional predictors differed between groups, and for species richness and functional diversity. Decomposer species richness responded to tree vitality, while functional diversity responded to habitat connectivity, predator species richness responded to regrowth of shrubs while functional diversity responded to tree circumference, and flower visitor richness and functional diversity did not respond to any additional predictors. Previous studies have found that the features and surroundings of veteran oaks are important for conservation of taxonomic diversity, and the results from this study indicate that they are also important for functional diversity within multiple functional groups.     

Phylogenetic placement of the austral rove beetle genus Hyperomma triggers changes in classification of Paederinae (Coleoptera: Staphylinidae)

Sister‐group relationships are resolved for the systematically and biogeographically puzzling austral rove beetle genus Hyperomma by means of phylogenetic analysis of five gene markers (one mitochondrial and four nuclear protein‐coding) for 25 taxa broadly representing the subfamily Paederinae, and six outgroup taxa from Staphylininae and Pseudopsinae. As a result, the new subtribe Dicaxina subtrib. nov. is established for Hyperomma and five other Southern Hemisphere genera previously classified in Cryptobiina. Based on the molecular phylogeny and the discussion of several adult and larval morphological characters, the concept of the tribe Paederini is changed as follows: Paederini sensu novo is reduced to include Paederina, Cryptobiina, Dolicaonina and Dicaxina only, while Lathrobiini sensu novo is established for Lathrobiina, Scopaeina, Astenina, Stilicopsina, Medonina, Stilicina and Echiasterina. The tribe Cylindroxystini stat. resurr. is reinstated for the Paederini subtribe Cylindroxystina because of its very peculiar morphology not fitting either Paederini or Lathrobiini in new sense. The tribe Pinophilini was resolved as sister to Lathrobiini sensu novo, and its status remains unchanged. Morphological diagnoses and other relevant systematic information are provided for all newly established taxa. The taxonomic history of the higher‐level systematics of the subfamily Paederinae is summarized.     

Spernathecal morphology,sperm transfer and a novel mechanism of sperm displacement in the rove beetle,Aleochara curtula (Coleoptera,Staphylinidae)

Summary The mechanism by which sperm are transferred from the male's spermatophore to the female's storing cage is described for the rove beetle Aleochara curtula, emphasizing a novel mechanism of sperm displacement by competing males. The cuticular, U-shaped spermatheca is equipped with a valve structure and two sclerotized teeth. The tube of the spermatophore extends into the spermathecal duct through the guidance of the flagellum of the male endophallus. Further elongation of the spermatophore tube, however, occurs only after separation of the pair. A primary tube bursts at its tip after passing through the valve. Within the lumen of the primary tube, a second tube passes through the valve and continues to extend up to the apical bulb of the spermatheca, doubles back on itself and swells to form a balloon filling most of the spermatheca. The balloon of the spermatophore is pierced within the spermatheca by tooth-like structures pressed against the spermatophore through contraction of the spermathecal muscle. The same process of spermatophore growing and swelling is also observed in mated females. Sperm from previous copulations are backflushed through the valve and the spermathecal duct, indicative of last-male sperm predominance.Abbreviations ad adhesive secretion covering the sperm - sac am amorphous secretion of the spermatophore - as ascending portion of the spermatophore - ds descending portion of the spermatophore - end parts of the male endophallus - ext extended tube - f flagellum - gs genital segment - lt large tooth - m muscle of the spermatheca - nsc non sclerotized cuticle - op opening of the spermathecal gland - pt primary tube - sc sclerotized cuticle - sd spermathecal duct - se secretion of the spermathecal gland - sf secretion flowing out of the primary tube - sg spermathecal gland - sm sperm - smt small tooth - sp spermatheca - ss sperm sac - st secondary tube - vm vaginal muscle     

Morphogenia: a new genus of the Neotropical tribe Jubini (Coleoptera,Staphylinidae, Pselaphinae) from the Brazilian Amazon

A new genus and species of the large Neotropical pselaphine tribe Jubini is described from Manaus, Brazil, based on material preserved in the Natural History Museum, London. Morphogenia struhli gen. et sp. n. represents the possible sister taxon of the abundant and speciose genus Barrojuba Park, sharing with it the putatively derived condition of anterolaterally shifted vertexal foveae, producing a smoothly convex vertex devoid of fovea or sulci. However, unlike Barrojuba, Morphogenia retains a plesiomorphic antebasal sulcus on the pronotum in both sexes, and additionally lacks elaborate abdominal fovea-like pockets and teeth on the lateral margins of the pronotum that are typical of Barrojuba. The genus is also unusual among jubine genera in lacking the characteristic V- or Y-shaped gular carina. In contrast to the commonly-collected Barrojuba, specimens of Morphogenia are absent in extensive jubine collections housed in museums in the United States, indicating that the new taxon may be relatively scarce or localised.     

Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification

The rove beetle tribe Staphylinini (Staphylinidae: Staphylininae) is a monophyletic lineage of over 5500 relatively large and charismatic species, yet its higher classification remains deeply rooted in historical concepts. Despite recent progress toward inferring phylogenetic relationships within this group using morphological and molecular datasets, relationships among taxa that were united under a polyphyletic “Quediina” remain largely unknown. To infer these relationships, we analysed a six‐gene dataset (4370 bp) using parsimony and model‐based analyses and the results were placed in the context of morphology. While all genes contributed synapomorphies for major lineages or relationships between them, carbamoyl synthetase (CAD), topoisomerase I (TP) and wingless (Wg) were the most informative. TP was generally most informative at the level of subtribe, Wg above this level and CAD throughout the tree. The monophyly of Staphylinini was strongly supported and analyses support seven clades that correspond to higher taxonomic levels, four of which are formally described as subtribes here: Acylophorina stat. rev., Cyrtoquediina new subtribe, Erichsoniina new subtribe and Indoquediina new subtribe. The majority of Staphylinini taxa were recovered within a well‐supported “northern hemisphere clade” that is weakly represented in the southern hemisphere. The composition and morphological diagnosis of the “Staphylinini propria” clade are revised, and the pronotum shape historically associated with this group is shown to have evolved multiple times elsewhere in Staphylinini. The genus Stevensia is moved from Staphylinina to Acylophorina based on morphological evidence. Cyrtoquedius stat. nov., previously a subgenus of Quedius, is raised to the genus level. The following 32 new combinations (from Quedius) are proposed: Cyrtoquedius anthracinus (Solsky); C. arrogans (Sharp); C. basiventris (Sharp); C. bolivianus (Sharp); C. bruchi (Bernhauer); C. clypealis (Sharp); C. concolor (Sharp); C. flavicaudus (Sharp); C. flavinasis (Bernhauer); C. frenatus (Erichson); C. graciliventris (Sharp); C. jacobi (Scheerpeltz); C. jocosus (Sharp); C. labiatus (Erichson); C. laeviventris (Bernhauer); C. mexicanus (Sharp); C. ochropygus (Bernhauer); C. ogloblini (Bernhauer); C. ornatocollis (Bierig); C. protensus (Sharp); C. rufinasus (Sharp); C. verecundus (Sharp); C. verres (Smetana); Indoquedius borneensis (Cameron); I. dispersepunctatus (Scheerpeltz); I. javanus (Cameron); I. malaisei (Scheerpeltz); I. micantiventris (Scheerpeltz); I. parallelicollis (Scheerpeltz); I. philippinus (Cameron); I. recticollis (Scheerpeltz); and I. sanguinipennis (Scheerpeltz). Cyrtoquedius verres is recorded from the state of Georgia (USA) for the first time, which, together with its transfer from Quedius, extends the distribution of the Cyrtoquediina significantly northward into the Nearctic.     

A new genus and species of the rove beetle tribe Mesoporini from Baltic amber (Coleoptera: Staphylinidae: Aleocharinae)

A new extinct rove beetle, Palaeomesoporus electiricus gen. et sp.n., is described from a single specimen of Eocene Baltic amber. This fossil beetle is placed in the tribe Mesoporini, a ‘basal’ group of the mega-diverse subfamily Aleocharinae. Palaeomesoporus is easily discriminated from other mesoporine genera due to its less-developed antennal club, longer and slender elytron lacking a sinuate posterior margin, and prominently shorter mesotarsus. This finding sheds light on the paleodiversity and evolutionary history of the tribe and ‘basal’ Aleocharinae.     

Carabid beetle (Coleoptera: Carabidae) richness and functional traits in relation to differently managed grasslands in the Alps

Species richness, composition, and functional traits of carabid beetle assemblages (Coleoptera: Carabidae) were studied in relation to different grassland management. Carabid beetles were sampled during the summers 2008 and 2009 by 165 traps located in 11 sites in the central-eastern Italian Alps. Using mixed effect models to account for potential spatial bias, we found that mown grasslands had significantly more species, a lower proportion of wingless species and a lower proportion of species with long larval development than grazed and natural grasslands. Within grazed and mown grasslands, neither cattle density nor number of cuts had any significant effect neither on species richness nor on any of the traits. The influence of grassland management can be summarised as follows: (1) grazing does not change community structure and functional traits compared to natural grasslands; (2) mowing negatively affects the carabid beetle assemblages; (3) the intensity of grazing and of cutting may not affect the structure of species assemblages of ground beetles. Our results support the hypothesis that agroecosystem practices in alpine grasslands influence carabid beetle communities. Specifically, the species with traits typical of undisturbed habitats (low dispersal abilities and long larval development) are more sensitive to perturbations (e.g. cutting). Our suggestion for agricultural and environmental planning and for conservation schemes is that the preservation of natural grasslands (e.g. forest gaps) and the implementation of grazing should be promoted during the planning of agroecosystem mosaics.