Courtship and mating in the whip spider Phrynichus orientalis Weygoldt, 1998 (Chelicerata: Amblypygi)

Mating behaviour and spermatophore morphology of the south-east Asian whip spider Phrynichus orientalis is described. During courtship there is less bodily contact between mates than in other studied species of the same genus. However, a peculiarity for the species is that the pair often pauses for minutes, sitting front to front without any movement. For spermatophore formation, the male shifts his body under that of the female like in many other species. However, after spermatophore formation, the male lures the female to the spermatophore instead of pulling her as in most other species studied. The spermatophore differs from those of most other species of the genus by the fact that the cushions in front of the sperm masses are replaced by a hard, transparent mass which has the same function as the cushions. Mating behaviour and spermatophore morphology of P. orientalis thus show peculiarities that so far have not been described for other species of the genus. It is not clear whether these characters should be considered plesiomorphic or apomorphic. But it does clearly show that there is more diversity within this genus than expected.     

Reproductive Biology of two Species of Phrynichus, P. exophthalmus Whittick, 1940 and P. deflersi arabicus nov. ssp. (Chelicerata: Amblypygi)

Courtship, mating and spermatophores of Phrynichus exophthalmus and P. deflersi arabicus nov. ssp. are described. Courtship of P. exophthalmus is more complex than in other amblypygids, with close contact between mates nearly throughout the mating dance. After spermatophore formation, the female is lured to the spermatophore and then firmly pressed onto it. Mating behaviour and spermatophore confirm the placement of this species in the Phrynichus ceylonicus group and show that the similarities with Euphrynichus, like long and thin pedipalps with bacilliform apophyses instead of spines, are the result of convergent evolution. Courtship of P. d. arabicus is conservative, similar to that of P. dhofarensis and many other amblypygids. After spermatophore formation, the female is lured to the spermatophore, then pulled over and pressed onto it. The results suggest that the P. ceylonicus group and the P. deflersi group and sister groups, but the low number of investigated species precludes a definite conclusion.     

Spermatophores, female genitalia, and courtship behaviour of two whip spider species, Charinus africanus and Damon tibialis (Chelicerata: Amblypygi)

This paper describes courtship behaviour, spermatophore morphology, and the female genitalia of the African whip spiders Charinus africanus Hansen, 1921 (Charinidae) and Damon tibialis (Simon, 1876) (Phrynichidae). In C. africanus, only the first part of courtship behaviour, up to spermatophore formation, could be observed; though different in detail, it is similar to that of many other species. The small spermatophore of C. africanus contains one large median sperm package. Charinus africanus is one of the Charinus species with thin finger-like gonopods and the first species with such gonopods of which the spermatophore is known. Spermatophores and female genitalia of D. tibialis are similar, though different in details, to those of Trichodamon and Musicodamon. They thus suggest that these two genera are correctly included in the Damoninae.     

Kampf,Paarungsverhalten, Spermatophorenmorphologie und weibliche Genitalien bei neotropischen Geißelspinnen (Amblypygi,Arachnida)

Zusammenfassung Agonistisches Verhalten, Samenübertragung und Spermatophorenmorphologie vonTarantula (Phrynus) palmata (Herbst) undHeterophrynus alces Pocock sind ähnlich wie bei anderen Arten der Familie Tarantulidae (Phrynidae). Anders ist das agonistische Verhalten vonTrichodamon froesi Mello Leitao. Die Palpen tragen an ihren Spitzen kleine Scheren und sind beim Männchen stark verlängert. Beim Kampf versucht jeder, die Tibien des 3. Beinpaares des Gegners zu ergreifen und dann nach hinten und hochzustemmen. Die Balz besteht aus Serien von vibrierenden und trillernden Bewegungen der Fühlerbeine und gelegent lichen Eingreifen des Weibchens; das Weibchen antwortet darauf mit charakteristischen Fühlerbeinbewegungen. Die komplizierte Spermatophore wird dadurch entleert, daß das Weibchen auf zwei Hebel drückt und dadurch die Samenpakete hochschnellen läßt. Diese werden dann in die Receptacula eingeführt und mit den hakenartigen Strukturen an den Gonopoden ergriffen und abgerissen. Ein Vergleich der bisher untersuchten Arten zeigt, daß das Paarungsverhalten keine charakteristischen Unterschiede bei verschiedenen Familien oder Gattungen zeigt. Aber die Spermatophoren zeigen familientypische Baueigenschaften.

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Agonistic and mating behaviour, spermatophore morphology, and female genitalia in neotropical whip spiders (Amblypygi, Arachnida)

Summary Agonistic and mating beaviour and Spermatophore morphology ofTarantula (Phrynus) palmata (Herbst) andHeterophrynus alces Pocock are similar to those of other species of the Tarantulidae (Phrynidae). Males ofTrichodamon froesi Mello Leitao perform a different agonistic behaviour. Their pedipalps bear small apical chelae and are extremely elongate. During fighting, each male grasps the tibiae of the opponent's third legs and pushes them upward and backward. During mating, the male performs vibrating and tapping movements with its antenniform legs and occasionally grasps the female. The female responds by performing characteristic movements of her antenniform legs. The spermatophore is a complicated structure. It is emptied by the female's pressing down two levers which causes two sperm packages to become elevated. These are then grasped by the female's seminal receptacles and by two hook-like structures of her gonopods and finally pulled off the spermatophore. Comparison of the species so far studied shows that the mating behaviour, although it varies in different species, does not exhibit differences useful for taxonomic research. But the different spermatophores show family characteristic structural properties.

     

Reproductive Biology of Phrynichus dhofarensis Weygoldt, Pohl & Polak, 2002 (Chelicerata, Amblypygi, Phrynichidae)

The Phrynichus deflersi species group is a well-defined group inhabiting the Arabian deserts and adjacent areas. The relationship of this group to other species of Phrynichus (the P. ceylonicus group) is obscure. The study of mating behaviour and spermatophore morphology of a member of the Phrynichus deflersi species group allows comparison with other species of Phrynichus and with Euphrynichus and evidence concerning the interrelationships of these three groups. Features of morphology, mating behaviour and spermatophore morphology of Phrynichus dhofarensis suggest that the P. deflersi species group and the P. ceylonicus species group are sister groups and that both (the genus Phrynichus) comprise the sister group of Euphrynichus. This conclusion remains tentative given that only two members of the P. ceylonicus species group and one of the P. deflersi species group have been studied. Mating success is quite variable, suggesting sexual selection.     

The morphology of juvenile stages of Crotonia pulcher (Beck, 1962) (Acari: Oribatida: Crotonioidea) with a redescription of the adult

The morphology of juvenile and adult stages of the Neotropical oribatid mite Crotonia pulcher (Beck, 1962) from Ecuador is described and illustrated. This species seems to be a member of the cophinaria group of Crotonia.     

Identification and breeding biology of the diving petrels Pelecanoides georgicus and P. urinatrix exsul at South Georgia

Abstract

The biology of two species of diving petrel (Pelecanoides georgicus and P. urinatrix exsul) was studied at Bird Island, South Georgia. Existing criteria (bill shape and morphology, wing length) for distinguishing these species are reviewed, and several new characters are recognised. For adults bill depth, the colour of the posterior part of the tarsus, and vocalisations are distinctive; chicks have down of different colours (pale grey in P. u. exsul, dark grey in P. georgicus). At South Georgia the species breed in different habitats and at different times - P. u. exsul nests in steep tussock slopes in very early summer, P. georgicus in fine scree slopes in midsummer. P. georgicus lays a proportionately larger egg, and has an incubation period of 46 days (c.54 days in P. u. exsul) and a chick fledging period of 46 days. The fledging period of P. u. exsul is 54 days, very similar to recorded values for P. u. urinatrix (53.5 days) and P. u. chathamensis (56 days). Data on feeding frequency and feed size were derived from daily chick weighings and from twice-daily weighing during 30 days preceding fledging. In both species chicks are fed every night, and often by both parents. In P. georgicus true mean chick feed size is c.37 g; in P. u. exsul it may be slightly less. Analysis of chick stomach contents suggests that P. u. exsul feeds extensively on copepods, whereas P. georgicus largely takes krill (Euphausia superba). P. u. exsul breeding adults commence moulting before their chicks have fledged; P. georgicus moults exclusively in the non-breeding season. Ectoparasites were collected, and the feather louse Pelmatocerandra setosa was found to be restricted to P. u. exsul, and Pelmatocerandra enderleini to P. georgicus. P. georgicus, which breeds later and whose chicks fledge faster, is suggested to be better adapted to the climatic and marine environmental conditions than P. u. exsul, which may be the more recent colonist of these high latitudes.     

Description of the male reproductive system of Paguristes eremita (Anomura,Diogenidae) and its placement in a phylogeny of diogenid species based on spermatozoal and spermatophore ultrastructure

The male gonopores, male reproductive apparatus, spermatophore and spermatozoa of the Mediterranean hermit crab Paguristes eremita are described, using interference phase microscopy, scanning electron microscopy and transmission electron microscopy. A correlation is made between the gonopore morphology and the different kinds of setae accompanying them, and the reproductive biology of these crabs. Each testes merges into a tubular duct made up of four zones: (1) the collecting tubule with free spermatozoa; (2) the proximal zone, where the ampulla of the spermatophores starts to be formed; (3) the medial zone, where the ampulla is completed, the stalk lengthens and the pedestal is formed; (4) the distal zone, where the mature spermatophores are stored. The sizes of the different parts of the spermatophore and of the sperm are given and their exterior morphology and ultrastructure described and compared to congeners. The morphology of the gonopore, male reproductive system, spermatophore and spermatozoa of P. eremita are species-specific, clearly distinguishing the species from the other members of the family. The available spermatozoal and spermatophore data is used to place P. eremita within a sperm phylogeny of the hermit crab family Diogenidae.     

Two new species of Pergalumna (Acari,Oribatida, Galumnidae) from Costa Rica,including a key to all species of the genus from the Neotropical region

Two new species of oribatid mites of the genus Pergalumna (Oribatida, Galumnidae), P. elongatiporosa sp. n. and P. striatiprodorsum sp. n., are described from leaf litter of a secondary forest in Costa Rica. Pergalumna elongatiporosa sp. n. is most similar morphologically to P. horvathorum P. Balogh, 1997 and P. sura P. Balogh, 1997, however, it differs from both by the body size, body surface ornamentation and morphology of notogastral porose areas A1 and A3. Pergalumna striatiprodorsum sp. n. is most similar morphologically to P. hawaiiensis hawaiiensis (Jacot, 1934) and P. strigulata Mahunka, 1978, however, it differs from P. hawaiiensis by the length of interlamellar setae and surface ornamentation of the prodorsum; from P. strigulata by the surface of ornamentation of the notogaster, length of interlamellar setae and morphology of bothridial setae. An identification key to known species of Pergalumna from the Neotropical region is given.     

Unusual Pre-copulatory Behaviour in the Hermaphroditic Planarian Flatworm,Dugesia gonocephala (Tricladida,Paludicola)

Hermaphrodite mating behaviour is poorly understood, particularly in species with internal fertilization and mutual penis insertion. We collected detailed recordings on the frequency, timing and outcome of the mating sequence of the simultaneous hermaphroditic flatworm Dugesia gonocephala (Tricladida, Paludicola). Mating in this species differs from that in other planarians in that receptive partners show a stereotypic pre-copulatory behaviour which results in copulation in only 17% (n = 342) of all attempts. It includes an unusual phase during which one partner glides on top of the other and first the lower, then both individuals, spread out and flatten. Genital intromission is not possible in this position. All observed copulations (n = 57) were preceded by this behaviour and most mating sequences were interrupted at this stage. Pairs were seen starting new mating sequences up to 11 times. Which animal lay on top varied randomly within a series of attempts. Cocoon production was a good predictor of mating activity, and copulations mainly occurred during the night. Copulations lasted between 24 min and 9.3 h (mean = 4.7 ± 2.3 h) and penises were mutually inserted. In the field, 99% (n = 70) of all matings involved reciprocal spermatophore exchange, compared with only 50% in the laboratory (n = 26). The likelihood of reciprocal spermatophore transfer increased with increasing duration of copulation. Duration was independent of the events that took place during courtship. We discuss the possibility that partners interrupt a copulation when there is no guarantee of reciprocal spermatophore transfer.     

Morphological and morphometric appraisal of the spermatophore of the southern hermit crab Isocheles sawayai (Anomura: Diogenidae), with comments on gonopores in both sexes

The spermatophore morphology of the hermit crab Isocheles sawayai from southwestern Atlantic (Brazil) is described. The spermatophores show similarities with those described for other members of the family Diogenidae, especially with the recently described Loxopagurus loxochelis. The spermatophore is composed of three major regions: a sperm filled head or ampulla, a columnar stalk and a foot or pedestal. The spermatophores show specific morphology in having a circular ampulla, and a constriction or neck between the ampulla (100 μm) and the thin (27 μm), long stalk (500 μm). The stalk penetrates less than half way into the spermatophore head. Most spermatophores show one of the small posterior projections on the underside of the ampulla as being bigger than the other, making it asymmetrical. The size of the spermatophore is related to hermit crab size with direct relationships found between spermatophore ampulla width, total length, and peduncle length with shield length of the hermit crab. The morphological characteristics of the spermatophore of I. sawayai are species-specific distinguishing it from other members of the family, and are useful to infer further phylogenetic relationships.     

Repeated visitations of spermatophores and polyandry in females of eriophyoid mites

Eriophyoid females store sperm either asymmetrically in one spermatheca, or symmetrically in both spermathecae. Previous studies have suggested that species in which females store sperm asymmetrically pick up sperm from only one spermatophore, while those with symmetrical sperm storage pick up sperm from two or more spermatophores during their lifetime. The aim of this study was to examine spermatophore visitation behaviour and symmetry of sperm storage in Aculops allotrichus from the black locust tree and Cecidophyopsis hendersoni from the yucca. This would indicate monandry or polyandry in these species. In both eriophyoids, the spermatophore visitation consisted of three phases: mounting, lying on the spermatophore and dismounting. Aculops allotrichus stored sperm asymmetrically. However, nearly one-third of the observed females visited two spermatophores, rather than only one in their lives. When A. allotrichus females visited two spermatophores they spent a similar amount of time at the first and at the second visitation. Also, the times of visitation of the first of the two spermatophores and the single spermatophore in a female lifetime did not differ significantly. This would suggest that apart from monandry, double insemination also occurs in this species. By contrast, C. hendersoni females were polyandrous. They stored sperm symmetrically and visited several spermatophores, on average 1.54 (max 6) per day, and up to 33 spermatophores in their lives. The benefits of repeated spermatophore visitation and the possible mechanisms of sperm storage in both species are discussed.     

Mating effort function of the spermatophore in the bushcricket Poecilimon veluchianus (Orthoptera, Phanero[pteridae): support from a comparison of the mating behaviour of two subspecies.

To analyse spcrmatophore function, various aspects of the mating behaviour (e.g. spermatophore mass, duration of sperm transfer, mating frequency) were compared in two subspecies of the bushcricket P. veluchianus. Body mass was significantly different in both subspecies and had a strong effect on spermatophore mass, resulting in a large difference in absolute and relative (percentage of male body mass) spermatophore mass in both subspecies. After copulation the small P. v. minor spermatophores were consumed much faster by the female than the larger ones of P. v. veluchianus. The time necessary for sperm transfer from the spermatophore to the female spermatheca was much shorter in the subspecies with small spermatophores than in that with large spermatophores, and in both subspecies similar to the time required to consume the spermatophores. Mating frequency varied also between the subspecies and was murh higher in P. v. minor than in P. v. veluchianus. Differences in body mass between the two subspecies therefore resulted in changes in several aspects of mating behaviour. However, despite large differences in the mass of the spermatophore, its main function seems to be to ensure sperm transfer. This can be concluded from the similarity between the duration of sperm transfer and the time period necessary for spermatophore consumption in each of the subspecies. The spermatophore is thus considered to be male mating effort.     

A new species of Chiasmocleis (Microhylidae,Gastrophryninae) from the Atlantic Forest of Espírito Santo State,Brazil

Among Neotropical microhylids, the genus Chiasmocleis is exceptionally diverse. Most species of Chiasmocleis were described in recent years based on external morphology, but recent studies using molecular data did not support the monophyly of the species groups clustered based on feet webbing. Furthermore, a phylogeographic study of C. lacrimae estimated high genetic divergence and low gene flow among populations across small geographic ranges. Increasing the molecular and geographic sampling, and incorporating morphological data, we identified new cryptic species. Herein, we used novel genetic and morphological data to describe a new species of Chiasmocleis.     

Pterandra pyroidea: a case of pollination shift within neotropical Malpighiaceae

Background and Aims

Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species.

Methods

Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods.

Key Results

Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera.

Conclusions

Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.     

A new Planthopper (Insecta: Hemiptera: Nogodinidae) from Chiapas amber, middle Miocene of Mexico

A new species of Planthopper, Nogodina chiapaneca sp. nov. from the Mexican Chiapas amber (southeast Mexico) is described. The new species belongs to Nogodinidae: Nogodinini: Nogodinina. This is the second nogodinid species described from Mexican amber. Both species belong to Nogodinina, a taxon with a Neotropical distribution. The only recent species of Nogodina, N. reticulata, ranges from Honduras to Brazil and is unknown from Mexico yet.     

Argulus japonicus: Sperm transfer by means of a spermatophore on Carassius auratus (L)

The process of sperm transfer is somewhat enigmatic in Argulus, even though copulation has been witnessed. A breeding colony of Argulus japonicus was kept under laboratory conditions in order to study reproduction in the species. Pairs in copula were removed and studied with histology and scanning electron microscopy to describe the mechanism of sperm transfer. Sections of copulating pairs revealed sperm on the accessory copulatory structures of the male’s swimming legs; and scanning electron microscopy showed that sperm transfer occurs in three phases which can be differentiated to 10 different stages. Sperm transfer occurs via a spermatophore which is extruded from the genital aperture of the male and is then transferred to the socket on the third pair of legs of the male, before being transferred into the spermathecae of the female via the spermathecal spines. This is the first observation of a spermatophore in Argulus.     

Are the microcyclic rust species Puccinia melampodii and Puccinia xanthii conspecific?

The taxonomic relationship between two microcyclic rust species, Puccinia melampodii and Puccinia xanthii, recorded from a number of Asteraceae hosts, was explored by comparing teliospore morphology, host specificity and sequence data. Teliospore morphology varied between and within individual rust accessions but, in general, teliospores of P. xanthii were larger than those of P. melampodii. Field observations and cross-inoculation greenhouse studies showed that P. melampodii accessions were highly specific to their original host species and unable to establish compatible interactions with related Asteraceae species. Though small amounts of genetic variation were detected between P. melampodii and P. xanthii accessions, the ITS and TEF phylogenetic analyses comprising several other rust species grouped these accessions within a well supported clade. Our data indicate that despite differences between P. xanthii and P. melampodii accessions these rusts cannot readily be separated at the species level, supporting earlier taxonomic conclusions and the recent proposal to group them into a morphospecies. Based on host-specificity data, the name Puccinia xanthii var. parthenii-hysterophorae is proposed to accommodate all Mesoamerican records of P. melampodii associated with the host Parthenium hysterophorus.     

Snake venomics of the pit vipers Porthidium nasutum, Porthidium ophryomegas, and Cerrophidion godmani from Costa Rica: toxicological and taxonomical insights

Within the Neotropical pit vipers, a lineage of primarily Middle American snake species referred to as the “Porthidium group” includes the genera Atropoides, Cerrophidion, and Porthidium. In this study, the venom proteomes of Porthidium nasutum, P. ophryomegas, and Cerrophidion godmani from Costa Rica were analyzed, and correlated to their toxic and enzymatic activities. Their HPLC profiles revealed a higher similarity between the two Porthidium species than between these and C. godmani. Proteins belonging to nine (P. nasutum), eight (P. ophryomegas), and nine (C. godmani) families were identified by mass spectrometry or N-terminal sequencing. Final cataloging of proteins and their relative abundances confirmed the close relationship between venoms of P. nasutum and P. ophryomegas, departing from that of C. godmani. Since the latter species had been taxonomically classified as Porthidium godmani previously, our venomic analyses agree with its current generic status. Venoms of P. nasutum and P. ophryomegas, despite containing abundant metalloproteinases and serine proteinases, lack procoagulant activity on human plasma, in contrast to venom of C. godmani. The latter induced strong myotoxicity in mice, which correlates with its high proportion of phospholipases A₂, whereas venoms from the two Porthidium species, containing lower amounts of these enzymes, induced only mild muscle damage.     

Ochoterenella esslingeri n. sp. (Nematoda: Onchocercidae: Waltonellinae) from Bokermannohyla luctuosa (Anura: Hylidae) in Minas Gerais,Brazil, with notes on Paraochoterenella Purnomo & Bangs, 1999

The waltonelline Ochoterenella esslingeri n. sp., a filarial parasite of the anuran Bokermannohyla luctuosa in Minas Gerais, Brazil is described. Several characters distinguish this new species from the 15 species presently included in the genus: the cuticular ornamentation of the female that is restricted to the posterior region of the body, the irregular arrangement of the small, rounded bosses, the postoesophageal vulva, the short glandular oesophagus, the size and shape of the microfilariae, the long left spicule and high spicular ratio. Irregularly arranged, tiny, rounded bosses are common in the monotypic genus Paraochoterenella from an Indonesian ranid, which is not well defined but likely valid. In the Neotropical Realm, the type hosts of the species of Ochoterenella are Hylidae (O. esslingeri n. sp.), Leptodactylidae (two species) and the remaining 13 species were described from the giant toad Rhinella marina (Bufonidae).