A unique case of commensalism: The beaver beetle Platypsyllus castoris (Leiodidae,Coleoptera) and its morphological adaptations

Platypsyllus castoris is closely associated with beavers and displays a unique set of structural specializations. We document the morphology of adults with modern techniques, and interpret evolutionary changes linked with the specific life style. The small subfamily Platypsyllinae has evolved an entire suite of features correlated with a more or less close association with mammals, for instance a flattened body, a dorsal cephalic shield, flightlessness, eye reduction, and depigmentation. Within this small group, Platypsyllus displays numerous autapomorphic features, correlated with a close association with the beaver. Essential is a combination of mechanical stabilization and firm anchorage on the host, and efficient forward movement in the fur. Exo- and endoskeletal structures of the head and thorax are reinforced by vertical cuticular columns and by an array of internal ridges. The antennae are shortened and strongly modified, the mandibles distinctly reduced and flattened, unsuitable for cutting, scraping or grinding. The musculature of the mouthparts is simplified, whereas an enhanced set of prepharyngeal and pharyngeal dilators forms an efficient sucking pump. The prothoracic musculature is strongly developed. In contrast, the pterothoracic muscle system is distinctly simplified, even though leg muscles are strongly developed. Using the legs, the flattened beetles move sideways through the dense fur of the beaver, using posteriorly directed groups of setae and ctenidia to prevent being pushed backwards by the densely arranged hairs. In contrast to the anterior body, the cuticle of the abdomen is thin, and the entire tagma flexible, with thin layers of segmental muscles. The hind gut is not connected with the mid gut. The beetles probably consume liquid, possibly with emulgated minute skin debris. As the morphology of the mouthparts excludes damage to the skin of the host, the association should not be addressed as ectoparasitic but as commensalism.     

First cavernicolous trechine beetle discovered in Guilin karst,northeastern Guangxi (Coleoptera,Carabidae, Trechinae)

A new subgenus and new species of anophthalmic trechine beetles, Oodinotrechus (Pingleotrechus) yinae subgen. n., sp. n., is described and illustrated from a limestone cave called Chaotianyan in southern part of Guilin karst, northeastern Guangxi Zhuang Autonomous Region. The new taxon is very different from the Maolan-Mulun congeners belonging to the nominate subgenus Oodinotrechus (s. str.) Uéno, 1998, in several important character states including pronotal structure, elytral chaetotaxy and male genitalia. It is the first record of a cavernicolous trechine beetle in Guilin karst, and in the eastern part of Guangxi. In addition, a distribution map for the genus Oodinotrechus Uéno, 1998, is provided.     

The cephalic morphology of free-living and cave-dwelling species of trechine ground beetles from China (Coleoptera,Carabidae)

Morphological adaptations of cave-dwelling organisms including different groups of Coleoptera have fascinated researchers since a long time. Nevertheless, very few detailed documentations of the anatomy of cave beetles using modern techniques are available. In this study, we describe and illustrate external and internal cephalic features of free-living and cave-dwelling trechine carabid beetles using digital microscopy, SEM, micro-CT, and computer-based 3D reconstruction. Morphological characteristics found in three selected species with different habitat preferences are compared. The following derived features distinguish a troglobite species (Sinaphaenops wangorum Ueno et Ran 1998) from a species living in the entrance of caves (Trechiotes perroti Jeannel 1954) and from a fully epigean species (Bembidion sp.): (1) the optic lobes are completely missing; (2) the head capsule, mouth parts, and antennae are distinctly elongated; (3) some muscle attachment areas are shifted; and (4) an additional dilator of the pharynx is present. Despite of conspicuous differences likely related with subterranean habits, the link of some cephalic features to this specific habitat preference remains uncertain.     

The thoracic anatomy of Pselaphus heisei (Pselaphinae,Staphylinidae, Coleoptera)

We document external and internal thoracic structures of the free-living pselaphine beetle Pselaphus heisei (Pselaphitae) using a set of traditional and modern techniques. Like in the specialized myrmecophile Claviger testaceus (Clavigeritae), the skeletal elements of the pro- and pterothorax are highly compact, with largely reduced inter- and intrasegmental sutures. Features previously listed as synapomorphies of staphylinid subgroups, to which Pselaphinae belong, are confirmed for P. heisei. The only previously proposed thoracic synapomorphy of Pselaphinae, the mesoventral foveae, is likely transformed in P. heisei: we assume that the concavities are directed mesad and internally fused, thus forming a broad channel (“perforation”) extending through the keel-like median region of the mesoventrite. The prothoracic foveal system is strongly reduced, with only one pair of pits present in front of the procoxae. Their internal invaginations form a transverse ventral endoskeletal bar that stabilizes the prothorax. The condition observed in the free-living P. heisei is in contrast with previous hypotheses linking the reduction of the foveal system with myrmecophily. Moreover, traces of the mesoventral foveae are even preserved in the highly specialized inquiline C. testaceus. Gland cells are associated with areas of hyaline squamose setae on different body regions, suggesting release of secretions on the ventral side of the head, pro- and mesothorax, and abdominal base. Similar specialized setal patches are common in Pselaphini and related groups within Pselaphitae. The prothoracic musculature in P. heisei is more complex than that in the myrmecophilous C. testaceus and the free-living, unspecialized Creophilus maxillosus (Staphylininae). The metathoracic muscle system is strongly simplified, demonstrating that P. heisei cannot fly, even though wings, some skeletal elements of the flight apparatus, and some small direct flight muscles are preserved. It cannot be fully excluded that indirect flight muscles and thus a functional flight apparatus is preserved in a certain percentage of individuals.     

Parallels between two geographically and ecologically disparate cave invasions by the same species,Asellus aquaticus (Isopoda,Crustacea)

Caves are long‐known examples of evolutionary replications where similar morphologies (troglomorphies) evolve independently as the result of strong natural selection of the extreme environment. Recently, this paradigm has been challenged based on observations that troglomorphies are inconsistent across taxa and different subterranean habitats. We investigated the degree of replicated phenotypic change in two independent cave invasions by the freshwater isopod Asellus aquaticus; the first in a sulphidic aquifer in Romania, the second in a sinking river in the Dinaric Karst in Slovenia. Both ancestral surface populations still live alongside the subterranean ones. Phylogenetic analyses show independence of the two colonization events, and microsatellite analysis shows no evidence of ongoing genetic exchange between surface and subterranean ecomorphs. The overall morphology has changed dramatically at both sites (50 of 62 morphometric traits). The amount of phenotypic change did not reflect differences in genetic diversity between the two ancestral populations. Multivariate analyses revealed divergent evolution in caves, not parallel or convergent as predicted by the current paradigm. Still, 18 traits changed in a parallel fashion, including eye and pigment loss and antennal elongation. These changes might be a consequence of darkness as the only common ecological feature, because Romanian caves are chemoautotrophic and rich in food, whereas Slovenian caves are not. Overall, these results show that morphologically alike surface populations can diverge after invading different subterranean habitats, and that only about one‐third of all changing traits behave as troglomorphies in the traditional sense.     

Parallel morphological evolution and habitat‐dependent sexual dimorphism in cave‐ vs. surface populations of the Asellus aquaticus (Crustacea: Isopoda: Asellidae) species complex

Studying parallel evolution (repeated, independent evolution of similar phenotypes in similar environments) is a powerful tool to understand environment‐dependent selective forces. Surface‐dwelling species that repeatedly and independently colonized caves provide unique models for such studies. The primarily surface‐dwelling Asellus aquaticus species complex is a good candidate to carry out such research, because it colonized several caves in Europe. By comparing 17 functional morphological traits between six cave and nine surface populations of the A. aquaticus species complex, we investigated population divergence in morphology and sexual dimorphism. We found habitat‐dependent population divergence in 10 out of 17 traits, likely reflecting habitat‐driven changes in selection acting on sensory systems, feeding, grooming, and antipredator mechanisms. Sexual dimorphism was present in 15 traits, explained by sexual selection acting on male traits important in male–male agonistic behavior or mate guarding and fecundity selection acting on female traits affecting offspring number and nursing. In eight traits, the degree of sexual dimorphism was habitat dependent. We conclude that cave‐related morphological changes are highly trait‐ and function‐specific and that the strength of sexual/fecundity selection strongly differs between cave and surface habitats. The considerable population variation within habitat type warrants further studies to reveal cave‐specific adaptations besides the parallel patterns.     

A new cave-dwelling and endemic species of the genus Pholeuonopsis (Coleoptera, Leiodidae) from Serbia

In the present study, a new species of a cavernicolous beetle is described from Serbia: Pholeuonopsis (Pholeuonopsis) zlatiborensis sp. n. (Leptodirini, Cholevinae, Leiodidae). All important taxonomic features have been thoroughly analyzed and illustrated. The new species probably belongs to an old phyletic lineage of mesogeid origin, like other known Pholeuonopsis species inhabiting cave and endogean habitats in the western part of the Balkan Peninsula. Therefore, the species is both relict and endemic inhabitant of cave habitats in Serbia.     

Stevewoodia minutum, a new genus and species of Scolytidae (Coleoptera) from the West Indies. Studies on West Indian Scolytidae (Coleoptera) 6

A new genus of Scolytidae (Coleoptera), Stevewoodia, from St. Lucia in the Lesser Antilles, is herein named and described. The type species, Stevewoodia minutumsp. n. is also named. The genus is named in honor of the late Steven L. Wood for his many contributions to the systematics of the Scolytidae.     

Phylogeny of the Agathidiini Westwood (Coleoptera: Leiodidae) and implications for classification and contractile morphology

A cladistic analysis of the tribe Agathidiini Westwood is presented. Agathidiines are slime mould specialists and they are hypothesized to be a monophyletic group consisting of 12 genera (Afroagathidium Angelini & Peck, Agathidium Panzer, Anisotoma Panzer, Besuchetionella Angelini & Peck, Cyrtoplastus Reitter, Decuria Miller & Wheeler, Gelae Miller & Wheeler, Liodopria Reitter, Pseudoagathidium Angelini, Sphaeroliodes Portevin, and Stetholiodes Fall), based on three synapomorphies: epipleuron present to apical third, mesoventrite without longitudinal carina and longitudinal setal lines present on the tibiae. The dataset for phylogenetic analysis comprised 72 characters representing 198 character states derived from adult morphology. These data were analysed using equal weighting and implied weighting (k = 1–6) and supported the monophyly of the tribe based on three unique characters (epipleuron present to apical third, mesoventrite without longitudinal carina, longitudinal setal lines present on tibia) and two homoplastic characters [antennomeres 7–10 (or 6–9) asymmetrical, apical shape of terminal antennomere abruptly tapered]. The topology of IW trees with k = 4–6 was identical with one of three EW trees. Decuria was sister group to the remaining agathidiine genera whereas the following groups were resolved as monophyletic: Anisotoma, Gelae + Liodopria, and Pseudoagathidium (Afroagathidium + Besuchetionella). The clade [Sphaeroliodes rufescens (Agathidium bockshini, Agathidium subcostatum)] was supported in all analyses except for the IW (k = 1) cladogram. The monophyly of Agathidium was not supported at all and was rendered paraphyletic by the placements of Sphaeroliodes, Stetholiodes and the Pseudoagathidium (Afroagathidium + Besuchetionella) clade. Sphaeroliodes is synonymized with Agathidium ( syn.n. ) resulting in two new combinations [A. acuminatus (?vec) and A. rufescens (Portevin)]. Contractability is a complex character composed of several morphological features that have evolved independently within the agathidiine tree. Conglobation (the ability to roll the body into a ball) has arisen at least twice in Agathidiini.     

Pretarsal structures in Leiodidae and Agyrtidae (Coleoptera,Staphylinoidea)

We analysed pretarsal characters of 87 species of Leiodidae (including 10 cholevines and representatives of all tribes and ca. 60% of the genera of non‐cholevines), five species of Agyrtidae, and nine representatives of outgroup taxa (Hydraenidae, Staphylinidae, Hydrophilidae, and Histeridae) using scanning electron microscopy. We focused our observations on the architecture of the empodium (including the sclerites and associated setae), the shape and composition of the medial projection of the distal margin of the terminal tarsomere, and the armature of the claws, which were considered a promising source of information for delimiting supraspecific taxa in our previous study. We identified several diagnostic features and recognize potential synapomorphies at the tribal, subtribal and generic levels. The internal systematic arrangement and/or even the monophyletic status of most of the subfamilies of Leiodidae (Camiarinae, Catopocerinae, Leiodinae, and Platypsyllinae) are challenged. We identified potential synapomorphies for Camiarinae (Camiarini and Agyrtodini) and Leiodinae. The non‐monophyly of Cholevinae is possible because part of the tribe (Anemadini, Eucatopini, and Oritocatopini) shares potentially apomorphic features with Leiodinae (e.g., a triangular medial projection with a diagonal row of conical spines), whereas another part (Leptodirini and Ptomaphagini) shares a potentially apomorphic feature with Coloninae and Platypsyllinae (a typical medial projection with two distinct triangular projections).     

中国广东优树甲属一新种(鞘翅目:拟步甲科)

记述采自中国广东的优树甲属1新种:高氏优树甲Uenostrongylium gaoi Lin&Yuan sp.nov.,该种因鞘翅刻点上缘具小颗粒而与浙江的粗皱优树甲Uenostrongylium scaber Yuan&Ren,2018相似.文中提供了新种的整体图和特征图.     

Ultrastructure of the mycangium of the southern pine beetle,Dendroctonus frontalis (Coleoptera: Curculionidae,Scolytinae): complex morphology for complex interactions

Yuceer, C, Hsu, C.‐Y., Erbilgin, N and Klepzig, K.D. 2011. Ultrastructure of the mycangium of the southern pine beetle, Dendroctonus frontalis (Coleoptera: Curculionidae, Scolytinae): complex morphology for complex interactions. —Acta Zoologica (Stockholm) 92 : 216–224. The southern pine beetle (SPB) (Dendroctonus frontalis Zimmermann) is the most economically important pest of southern pine forests. Beetles carry fungal cells within specialised cuticular structures, called mycangia. Little is known about the mycangia ultrastructure or function. We used cryo‐fracturing and scanning electron microscopy to examine the ultrastructural features of SPB mycangia and surrounding tissues. Mycangia, one on each side of anterior portion of the prothorax, are terminated on the dorsal side at a ‘mycangial bridge’. This sclerotised mycangial bridge does not appear to provide a passage between the two mycangia, suggesting that each mycangium functions independently. Mycangia are surrounded by abundant tracheoles connecting the structures to the outside via openings within the prothorax. Previously unknown pits overlying the mycangial gland cells were also observed in both the inner wall and anterior fold of prothorax. We hypothesise that these openings and pits may play roles in determining which fungi enter, and grow within, the mycangium.     

中国优树甲属一新种记述（鞘翅目：拟步甲科）

记述采自中国浙江的优树甲属1新种:Uenostrongylium scaber YuanRen sp.nov.。提供了特征描述,整体图和特征图,另外附有优树甲属分种检索表。     

A new hypogean species of the genus Domene Fauvel, 1873 from Turkey (Coleoptera: Staphylinidae: Paederinae)

Domene bordonii sp. n. is described and illustrated from Geyik Mountains, Konya province, central southern Anatolia. This new hypogean species was collected with a subterranean pitfall trap in the mesovoid shallow substratum.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:72D06EC5-885F-4BCC-8066-A16B28016697     

A second species of the genus Sciaphyes (Coleoptera: Leiodidae: Cholevinae: Sciaphyini) from Japan

A second species of the genus Sciaphyes Jeannel is reported from Japan. The new species, S. kawaharai sp. n., can be easily distinguished from other Japanese Cholevine by the lack of eyes and hind wings. S. kawaharai sp. n. was collected in the nests of lesser white-toothed Shrew, Crocidura suaveolens (Pallas 1811) (Insectivora: Soricidae), therefore its habitat is pholeophile.     

Antenna morphology and sensilla microstructure of the male and female scarab beetle,Lepidiota mansueta Burmeister (Coleoptera: Scarabaeidae)

Studies on Lepidiota mansueta Burmeister were conducted to describe and compare the external morphology of the antenna and its sensilla of male and females. Distinct variations were observed in antennal segments between the sexes with males possessing longer scape, pedicel, flagellum and lamellar club segments. Variation was also seen in the antennal sensilla, with males possessing three different types of sensilla while the females possessed seven types of sensilla. Sensilla trichodea (TRS), sensilla placodea (PLAS) type I and sensilla placodea (PLAS) type II were common in both sexes while sensilla chaetica (CHS), sensilla auricilica (AUS) type I, sensilla auricilica (AUS) type II and sensilla auricilica (AUS) type III were found only in females.     

Profound head modifications in Claviger testaceus (Pselaphinae,Staphylinidae, Coleoptera) facilitate integration into communities of ants

Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free-living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of Claviger testaceus, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking/‘sponging’ device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far-reaching cephalic modifications characterize C. testaceus, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.     

New species,new genera,and new records in Cerambycidae from Peru (Coleoptera)

Three new species and two new genera of longhorn beetles are described from Peru: Cosmoplatus polis n. sp. (Pteroplatini), Hemilocallia grishami n. gen., n. sp. (Calliini) and Arabela ramirezi n. gen., n. sp. (Hemilophini). Esmeralda laetifica Bates, 1869 (Prioninae), Stratone transversalis (Chevrolat, 1862) (Cerambycinae), Gymnocerina cratosomoides (Bates, 1862) (Lamiinae), and Trachysomus verrucosus (Olivier, 1795) (Lamiinae) are new records for Peru.     

Evolution of starvation resistance in an invasive insect species,Aethina tumida (Coleoptera: Nitidulidae)

Starvation resistance, or the ability to survive periods without food, can shed light on selection pressure imposed by food scarcity, including chances to invade new regions as a result of human transport. Surprisingly, little information is known about starvation resistance for invasive insect species. Given that native and invasive populations differ in starvation resistance, this would suggest different selection scenarios and adaptive shifts fostering invasion success. Here, we show striking differences in starvation resistance of adult small hive beetles Aethina tumida (SHB) between native and invasive populations. In the laboratory, starvation resistance of freshly emerged laboratory‐reared and field‐collected adult females and males was evaluated in the beetle's native African range and in their invasive North American range. SHB in their native African range survived longer than SHB in their invasive North American range. Across ranges, females survived longer than males. Field‐collected SHB survived in Africa longer than freshly emerged ones, but not in the invasive range. This suggests no selection for starvation resistance in the invasive range, possibly due to differences between African and European‐derived honey bee hosts facilitating a trade‐off scenario between reproduction and starvation resistance. The ability of adult females to survive up to two months without food appears to be one factor contributing to the invasion success of this species. Assuming food availability is usually high in the invasive ranges, and trade‐offs between starvation resistance and fecundity/reproduction are common, it seems as if selection for starvation resistance during transport could set up potential trade‐offs that enhance reproduction after invasion. It would be interesting to see if this is a possible general pattern for invasive insect species.     

Four new species of Cymatodera Gray from Mexico (Coleoptera,Cleridae, Tillinae)

Four new species of Cymatodera from Mexico: Cymatodera bogcioides sp. n.; Cymatodera pueblae sp. n.; Cymatodera mitae sp. n.; and Cymatodera lineata sp. n. are described based on adult male and female specimens. Male genitalia and other characters of taxonomic value are presented.