Phylogenetic relationships within the Phyllopoda (Crustacea,Branchiopoda) based on mitochondrial and nuclear markers

For several decades the relationships within the Branchiopoda (Anostraca + Phyllopoda) have been a matter of controversy. Interpretations of plesiomorphic or apomorphic character states are a difficult venture, in particular in the Phyllopoda. We explore the relationships within the Phyllopoda at the level of nucleotid comparisons of the two genes 12S rDNA (mitochondrial) and EF1alpha (nuclear), and at a higher molecular level based on introns found in the gene EF1alpha. Within the Phyllopoda our explorations show further evidence for a non-monophyletic Conchostraca (Spinicaudata + Cyclestherida + Laevicaudata). The monotypic Cyclestherida is more closely related to the Cladocera, both together forming the Cladoceromorpha. The Spinicaudata (Leptestheriidae, Limnadiidae, and Cyzicidae) is well supported. Spinicaudata and Cladoceromorpha form a monophylum. The position of the Laevicaudata remains unclear but we find neither support for a sister group relationship to the Spinicaudata nor for a close relationship of Laevicaudata and Cladocera. Within the Cladocera, we favour the Gymnomera concept with the monotypic Haplopoda being the sister group to the monophyletic Onychopoda. The Ctenopoda seems to be the sister group to the Gymnomera, which contradicts the common view of a more basal position of the Ctenopoda.     

The Cladocera (Crustacea: Branchiopoda) as a relict group

According to the fossil record and DNA data, the Cladocera is an ancient crustacean group. Recent revisions make their taxonomy amenable to zoogeographical analysis. A bipolar (antitropical) disjunct distribution of faunal complexes and taxa ( Daphnia , Daphniopsis , Pleuroxus , Tretocephala , etc.), the wide ranges of some species and narrow restriction of others, the presence of isolated populations and the concentration of endemics in the warm temperate – subtropical zone of both hemispheres are traits of cladoceran zoogeography. These enable us to compare them with better studied (both living and fossil) plants, invertebrates and vertebrates, and to analyse their faunal formation by the modern version of the concept of 'ejected relicts' instead of vicariance. This reveals the extant Cladocera as a relict group, whose taxa were widely distributed in the past. Tertiary climatic changes, primarily within the present tropical and boreal latitudes, resulted in mass extinction of their biotas, while the warm temperate – subtropical regions remained comparatively unchanged. Although most recent Cladocera have relict status, others such as the D. pulex and D. longispina species groups and the subgenus Eubosmina are evolutionary young and show recent speciation.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 109–124.     

Chorusing Behaviour,a Density-dependent Alternative Mating Strategy in Male Common Toads (Bufo bufo)

Male chorusing behaviour was studied in a population of common toads (Bufo bufo) on the island of Öland south Sweden, and the functional role of male advertisement calling in this species was experimentally examined. Calling males were larger and heavier than non-calling males (t = 2.41, p < 0.025 and t = 2.36, p < 0.025, respectively). However, small males were also found to call. This indicates that large males persisted in calling for longer and/or called more often. The proportion of calling males decreased as population size increased during the breeding season, indicating that calling is a low density strategy. Females responded more readily to calls than males. There were insufficient data to determine if the dominant frequencies of advertisement calls were inversely correlated with male body size, however, this relationship was found for the similar release calls. Females were found not to discriminate between high and low frequency calls, but when given a choice between two calls of different sound pressure levels (SPL), females were attracted to the louder calls. Thus, the function of chorusing is to advertise the position of males to mate-seeking females when the population density is low.     

The occurrence of artemocyanin in Branchiopoda (Crustacea)

Artemocyanin, the extracellular hemolymph biliprotein of Artemia, is demonstrated in the fairy shrimp Streptocephalus, the clam shrimp Leptestheria and the water flea Daphnia. Artemocyanins can be purified from hemolymph as intact polypeptides (M_r 170–190,000), but are degraded upon homogenization of the whole animal by partial proteolysis to polypeptides with M_r 102,000 and 85,000. The aminoterminal sequence of the intact artemocyanin polypeptide was determined, but no clear-cut relationships with arthropod biliproteins or other protein families could be demonstrated.     

A phylogenetic analysis of the Conchostraca and Cladocera (Crustacea, Branchiopoda, Diplostraca)

A computer-assisted cladistic analysis on morphological characters of the Diplostraca (Conchostraca and Cladocera) has been undertaken for the first time. The morphological information has been obtained from literature and transformed into 56 suitable characters. The analysis included 47 ingroup taxa, comprising five conchostracan taxa (four families of the Spinicaudata and the Laevicaudata) and 42 genera of the Cladocera. A detailed character discussion is presented which will be a useful working base for future phylogenetic studies on the group. A number of systematic groups were, with differing degrees of certainty, supported in all 218 equally short trees. These are the Diplostraca, Cladocera, Gymnomera (Onychopoda and Haplopoda), Onychopoda, Podonidae, Cercopagididae, Anomopoda, Daphniidae, Moininae, Scapholeberinae, Chydoridae, Chydorinae and Sididae. The Spinicaudata were only supported on some of the 218 equally short trees while no support was found for the Conchostraca. Two taxa—the Macrothricidae and Aloninae—were relatively strongly indicated to be paraphyletic. A suggested classificatory hierarchy, without indication of absolute rank, is presented.     

Wind-borne short-range egg dispersal in anostracans (Crustacea: Branchiopoda)

Evidence is provided for the first time that at least part of the egg bank of the southern African fairy shrimp Branchipodopsis wolfi is dispersed over short distances by wind. A total of 423 sticky surfaces were mounted around and between individual basins at three rock pool sites, to trap any dispersing eggs during a 3 day period. Eight viable eggs were found in different egg traps (1.9% of the total), seven of which were located at one site with shallow basins. Given the proximity of vast egg banks to the egg traps, the low observed dispersal rate cautions against overestimating the importance of wind dispersal for population genetic processes. By generating egg banks to hedge against drought catastrophes, and producing egg types with different potential dispersibility, B. wolfi is a strong bet-hedger which has established a means of escaping temporal and spatial stress. These life history functions enable populations to persist in small desert rock-pools, which are the most extreme of temporary habitats.     

Distribution,diversity and conservation of Anostraca (Crustacea: Branchiopoda) in southern Africa

Southern Africa is defined here as Africa south of the Zambezi and Kunene Rivers. Here, annual average rainfall, rainfall season, and effective temperature are climatic factors which influence anostracan distribution. The type of temporary pool also appears to have some effect on distribution. Areas of high species richness are not confined to any particular climatic region and such areas occur in arid, montane and subtropical regions of southern Africa. Forty-six anostracan species have been recorded from the subcontinent and 80% of these are endemic. IUCN Red Data assessment of species revealed that two species can be considered 'Critically Endangered', while a further four are 'Endangered', and three are 'Vulnerable'. Many parts of southern Africa have been inadequately sampled but in South Africa, where the anostracans are relatively well known, temporary pool inhabitants are threatened mainly by agriculture, urbanization, pollution and pesticides. This revised version was published online in August 2006 with corrections to the Cover Date.     

Feeding Mechanisms and Comparative Functional Morphology of Macrothricidae (Crustacea : Cladocera)

Feeding of the Macrothricidae is accomplished b y two principal methods: active collection of food from the substrate surface and filtration of suspended particles. As differentiated from Ilyocryptus sordidus and Lathonura rectirostris, Ophryoxus gracilis is equipped with the most universal set of limbs allowing catching of food along with filtratory feeding. They are described in detail. Lathonura rectirostris is a highly specialized species, an inhabitant of plant thickets, since i t adapted to feeding on epiphytic algae and particles at the plant surface. Feeding is accomplished only by scraping or catching food by means of complexes of setae of the II pair of limbs. Filtratory feeding and appropriate structures are lacking. The structure and action of the limb apparatus of Ilyocryptus sordidus rule out active catching of food particles.     

The status of Anostraca,Notostraca and Conchostraca (Crustacea: Branchiopoda) in Yugoslavia

The investigations of Anostraca, Notostraca and Conchostraca in Yugoslavia began after a great delay compared with other European countries. Intensive systematic investigations date back only to the 1970s.To date, 19 species have been recorded. All species, with the exception of Artemia salina, are faunal elements of the Pannonian region. Seven species, including Artemia salina, Branchinecta orientalis, Branchipus serbicus, Leptestheria dahalacensis, Limnadia lenticularis, Imnadia cristata and Imnadia panonica are known from single localities. Six species including Branchinecta ferox, Chirocephalus brevipalpis, Chirocephalus carnuntanus, Lepidurus apus, Eoleptestheria spinosa and Imnadia banatica have restricted distributions. Streptocephalus torvicornis and Cyzicus tetracerus are known from several localities, while Branchipus schaefferi, Chirocephalus diaphanus, Triops cancriformis and Leptestheria saetosaare common. Large branchiopods are mainly confined to anthropogenic landscapes, especially in the northern part of the country. Rare species inhabiting a single pond, or those with rather restricted distributions, are the most endangered and such species account for about 70% of the fauna. Limnadia lenticularis, Imnadia cristata and I. panonica have not been found for more than 20 years and they are probably extinct in Yugoslavia. Branchipus serbicus, Imnadia cristata and panonica have not been documented since being originally described. Eoleptestheria spinosa is currently the most endangered species since it only appears irregularly in small populations. At present, Branchinecta ferox, Chirocephalus carnuntanus and Imnadia banatica are the safest of the rare species as they regularly appear in large numbers. There are no large branchiopods on the List of protected animals in Yugoslavia. This revised version was published online in August 2006 with corrections to the Cover Date.     

The resting eggs of the Ctenopoda (Crustacea: Branchiopoda): a review

The resting eggs of the Ctenopoda have been studied since the second half of the 19th century. By now they are known in 21 species of the group but only in 4 species (Sida crystallina, Diaphanosoma brachyurum, Penilia avistoris and Holopedium gibberum) have they been investigated thoroughly. Gross and internal structure of eggs are characterized by considerable diversity. In general their structure has much in common with resting eggs of the Anostraca, Notostraca, Conchostraca and Onychopoda as well as with that of some other remotely related groups of freshwater invertebrates (Rotifera, Hydra). The taxonomic value of resting eggs is discussed. Probably they play a significant role in determining geographical distribution. Waterfowl are the main agent of their dispersal.     

Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater

With about 500 known species worldwide, the large brachiopods are a relatively small group of primitive crustaceans. With few exceptions they live in temporary aquatic systems that are most abundant in arid and semi arid areas. As many regions remain unexplored and as especially the number of species in clam shrimps and tadpole shrimps is underestimated due to difficult identification, the species list will increase with future surveys. The Branchiopoda are monophyletic, but inter-ordinal relationships, as well as many evolutionary relationships at lower taxonomic levels are still unclear. Ongoing molecular studies will more accurately depict species diversity and phylogenetic patterns. With the exception of some anostracan families, most families are not restricted to the northern or southern hemisphere or specific zoogeographical regions. Large branchiopods are used for the assessment of the quality and function of temporary wetlands. Due to the reduction in number and quality of temporary wetlands, several species became endangered and are red listed by the IUCN. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Redescription of Branchinecta granulosa Daday, 1902 from Argentina (Crustacea: Branchiopoda)

The species Branchinecta granulosa Daday 1902, is redescribed on the basis of adult material from near Facundo (Chubut Province, Argentina). Its relationship to its regional congeners is outlined.     

The Origin of Cladocera (Crustacea,Branchiopoda): A New Understanding of an Old Hypothesis

Comparative analysis of the ontogeny of representatives of two sister taxa (Cladocera and Cyclestherida) showed that the paedomorphic morphology of cladocerans (the small number of thoracic segments and segments of branches of antennae II, and the reduction of the carapace) was caused by the cessation of development of the somatic structures at early larval stages of ontogeny. It is demonstrated that this stop is not associated with the accelerated development of the reproductive system (progenesis), since it takes place long prior to the beginning of reproduction. In accordance with this fact, the past hypotheses that cladocerans evolved from the reproducing larvae of the ancestral form or that they are early maturing metanauplii should be recognized as erroneous. Cyclestheria. The origin of Cladocera from a Cyclestheria-like ancestor should be regarded as neotenic, taking into consideration the extended interpretation of this term.     

Functional morphology of amplexus (clasping) in spinicaudatan clam shrimps (Crustacea,Branchiopoda) and its evolution in bivalved branchiopods: A video‐based analysis

Male clam shrimps (Crustacea: Branchiopoda: Laevicaudata, Spinicaudata, and Cyclestherida) have their first one or two trunk limb pairs modified as “claspers,” which are used to hold the female during mating and mate guarding. Clasper morphology has traditionally been important for clam shrimp taxonomy and classification, but little is known about how the males actually use the claspers during amplexus (clasping). Homologies of the various clasper parts (“movable finger,” “large palp,” “palm,” “gripping area,” and “small palp”) have long been discussed between the three clam shrimp taxa, and studies have shown that only some structures are homologous while others are convergent (“partial homology”). We studied the clasper functionality in four spinicaudatan species using video recordings and scanning electron microscopy, and compared our results with other clam shrimp groups. General mating behavior and carapace morphology was also studied. Generally, spinicaudatan and laevicaudatan claspers function similarly despite some parts being nonhomologous. We mapped clasper morphology and functionality aspects on a branchiopod phylogeny. We suggest that the claspers of the three groups were adapted from an original, simpler clasper, each for a “stronger” grip on the female's carapace margin: 1) Spinicaudata have two clasper pairs bearing an elongated apical club/gripping area with one setal type; 2); Cyclestherida have one clasper pair with clusters of molariform setae on the gripping area and at the movable finger apex; and 3) Laevicaudata have one clasper pair, but have incorporated an additional limb portion into the clasper palm and bear a diverse set of setae. J. Morphol. 278:523–546, 2017. © 2017 Wiley Periodicals, Inc.     

A new branchipodid genus and species (Crustacea: Branchiopoda: Anostraca) from South Africa

Rhinobranchipus martensi gen. et sp. nov. (Anostraca: Branchipodidae) was collected from a shallow temporary pool in the surroundings of Grahamstown (Cape Province, South Africa) (33^o18'S, 26^o32'E). Rhinobranchipus shares partial fusion of the chitinized basal parts of the male antennae, the so-called clypeus, with other Branchipodidae. The new genus is distinguished from confamilial genera by the morphology of the basal part of the penes. The new species, known from its type locality only, should be considered endangered, if it has not already become extinct as a result of recent modifications of its habitat.     

pH-related Hatching Success of Triops cancriformis (Crustacea: Branchiopoda: Notostraca)

Six different pH ranges were tested for the hatching rate of Triops cancriformis cysts. Cumulative hatching numbers approach to approximately 50% of cysts. The highest hatching rates, co-occurring with the shortest hatching times, were recorded in the neutral range. No hatching took place at pH values 9 and above. This corresponds with the most frequent habitats of this species, snowmelt pools of river flood plains as well as aestival pools caused by heavy rainfalls. Occurrences in sodic alkaline pans can be explained by relatively low pH values at the time of inundation.     

Male Mating Tactics in the Shrimp Palaemonetes pugio (Decapoda, Caridea): Precopulatory Mate Guarding vs. Pure Searching

The guarding of females approaching a limited period of sexual receptivity is a common mating tactic of males. In many decapod crustaceans, such as the shrimp Palaemonetes pugio , females can only copulate during a short period after a reproductive molt. It has been predicted that mate guarding by males (pre-copula) evolves in such species if sex ratios are not highly female-biased and if males can detect the molt stage of the female. The mating tactics of males were investigated in P. pugio . Time-lapse video observations were made on interactions among two males, a pre-molt female, and an inter-molt female (20 replicates). There was no evidence that males recognized a pre-molt female until 24 h before its molt. Significant numbers of male contacts with pre-molt females occurred 1 h before and after the female molt. Copulation took place within 1–3 min of the molt. No behavior commonly associated with mate guarding in decapods was observed – no clasping, agonistic behavior, or close association. It is concluded that the male's mating tactic is pure searching, wherein males haphazardly contact many females in order to find a receptive one. The high encounter rate in nature of these very mobile, aggregated shrimps is proposed as the factor responsible for the evolution of pure searching. It is hypothesized that pure searching is the male tactic of the many species of decapod shrimps with small males, sexually monomorphic cheliped weapons, and aggregated populations.     

Variation in Male Calling Behaviour and Relation to Male Mating Success in the Strawberry Poison Frog (Dendrobates pumilio)

Calling behaviour and the characteristics of the males’ call are important for mate attraction, female mate choice and male mating success in anurans. In this study I investigated the variation in calling activity and the variation in one spectral (dominant frequency) and four temporal properties (call rate, call duration, duty cycle and pulse rate) of the advertisement call of strawberry poison frogs, Dendrobates pumilio, in two Costa Rican populations during two study periods. Regarding all call properties, no differences were found between populations or years, but between‐male variation was significantly higher than within‐recording and within‐male variation. Dominant frequency was less variable within and among males than temporal call properties and I classified the former as static and the latter as intermediate properties of the call. No call property seemed to be strongly affected by body length or weight. Moreover, I used behavioural observations to relate male calling behaviour to mating success. Calling activity was more variable between males and accounted for more variability in mating success in multiple regression models than any call property. Some call properties (call rate, dominant frequency and pulse rate) were also related to male mating success in one or two study periods. These call properties are probably important for females during courtship to assess their prospective mate and/or for competitive male–male interactions. Furthermore, I compared call properties and mating success of some males for which data were available for both study periods. Pulse rate was lower in the second than in the first year, suggesting age‐related changes in this call property. Moreover, male D. pumilio increased their relative mating success from the first to the second study year. Together with other studies, the data of this study suggest that, in anurans, endurance rivalry contributes more to variation in male mating success than does female choice based on male characteristics.