The Micro-caddisflies of Lord Howe Island (Hydroptilidae: Trichoptera: Insecta)

Hydroptilid species are described from Lord Howe Island for the first time. All four species are in the endemic Australian genus Orphninotrichia, and all are endemic to Lord Howe Island. Modifications exhibited by these new species to the normally conservative Orphninotrichia features are suggestive of sympatric speciation and, for several, of a shift in larval niche.     

Charr truth: sympatric differentiation in Salvelinus species

Charrs, Salvelinus species, are characteristic fishes of northern freshwater lakes and rivers. They are highly variable in almost every aspect of their behaviour, morphology and life history. Several possible explanations have been proposed to account for this variability and resolve the taxonomic confusion in this genus. I propose that sympatric trophic polymorphism, in the ecological context of these species, can make sense of this variability. I use examples from charr in Canada and Iceland to construct an evolutionary scenario for this genus.     

Sexual Dimorphism and Inter-Generic Variation in Proboscidean Tusks: Multivariate Assessment of American Mastodons (Mammut americanum) and Extant African Elephants

Characteristics of social structure, mating strategies, and parental investment can be inferred for mammalian species based on degree of sexual dimorphism, especially when males are substantially larger than females. American mastodons (Mammut americanum) and extant African elephants (Loxodonta africana, Loxodonta cyclotis) both exhibit marked dimorphism in tusk size. To evaluate the hypothesis that this dimorphism might be indicative of similar, and possibly conserved, behavioral patterns in each lineage, we undertook a detailed evaluation of the pattern of tusk growth in these two genera. Separate discriminant function analyses (DFA) of 21 adult mastodon tusks of inferred sex and 48 adult elephant tusks of known sex show that patterns of ontogenetic change in tusk circumference, regardless of genus, effectively discriminate between sexes. Canonical variates analysis (CVA) of tusks from male and female mastodons and male and female elephants shows that male tusks in both genera are larger than female tusks across all measurements, especially for maximum tusk circumference and pulp cavity depth. CVA’s emphasis of inter-group differences in tusk morphology also shows that mastodon tusks are more robust than elephant tusks. Overall, this study illustrates that there is a characteristic male and a characteristic female tusk form shared by elephants and mastodons. This shows that elephants and mastodons display a shared syndrome of traits beyond sex-linked differences in tusk size, supporting the hypothesis that mastodons exhibited behaviors similar to those we observe today in African elephants.     

Phylogeny, speciation and species turnover. The case of the Mediterranean gastropods of genus Cyclope Risso, 1826

The genus Cyclope Risso, 1826 (family Nassariidae) has appeared in the fossil record since the Pliocene. Although it is still found today, the teleoconch morphology has never undergone modification, despite the fact that the protoconch morphologies of fossils (multispiral) and living forms (paucispiral) are different. They vary in their embryological and larval development and, hence, are two different species: C. migliorinii (Bevilacqua, 1928), the fossil species, and C. neritea (Linnaeus, 1758), the living species. We discuss the morphologic modifications in the evolution of this genus: the speciation that leads to its appearance and the speciation driving the Pliocene species to the living one. The order and the direction of these changes are based on phylogenetic analysis. No intermediate forms have been found showing a gradual morphological change that could have been worked by natural selection. Our analysis takes as the origin of the morphological novelties the genetic modifications in the ontogenetic processes which resulted in rapid and important phenotypic changes. Both speciation processes are sympatric cladogenetic. The changes that determine the appearance of the genus affect only the teleoconch, not the larval development. The modifications that lead from one species to the other, within the genus Cycope, affect the larval development exclusively. This points to a certain disconnection between the development of the embryo-larval phase and the young-adult formation, such that evolutionary processes could have occurred independently in different ontogenetic stages. The influence of larval ecology in relation to extinction of the ancestor and persistence of the derived species is also analysed. We hypothesize that climatic fluctuations may have affected the planktonic larvae of the fossil species, driving it to extinction. The living species, developing without the planktonic phase, would have resisted these climatic changes. We consider that the mechanisms described as drivers of the evolution of this genus can be of more general validity in prosobranch gastropods.     

The role of head shape and trophic variation in the diversification of the genus Herichthys in sympatry and allopatry

In the present study we evaluated the putative cases of sympatric speciation in the genus Herichthys by studying the variation in head shape using principal component analysis, phylomorphospace and reconstructions of the ancestral states of feeding preferences. Herichthys includes both allopatric and sympatric sister species, as well as sympatric unrelated species and thus offers great potential for evolutionary studies of putatively sympatric speciation. Herichthys is the northernmost group of cichlids in America and one of the most ecologically disparate genera within Middle American cichlids. Fifteen anatomical points were recorded on the heads of 293 specimens of the 11 species recognized within the genus. The results show that in spite of having wide variation in consumed diets, most species of Herichthys are close in morphospace. However, morphological variation was great among the two pairs of sympatric sister species in agreement with the suggested sympatric model of speciation.     

The effects of mode of development on phylogeography and population structure of North Atlantic Crepidula (Gastropoda: Calyptraeidae)

The mode of development of marine invertebrates is thought to influence levels of population structure and the location of species range endpoints via differences in dispersal ability. To examine these effects, populations of three sympatric clades of sedentary, marine gastropods in the genus Crepidula were sampled along the Atlantic and Gulf coasts of North America. A haplotype tree was constructed for each clade based on 640 bp sequences of mitochondrial cytochrome oxidase c subunit I. Examination of the tree topology, and AMOVA analysis show that species with direct development (those hatching as benthic juveniles) have higher levels of population structure than do species with planktonic development. Both species in the direct-developing C. convexa clade have high levels of geographical differentiation, with most populations representing a discrete clade of haplotypes. The planktotrophic species C. fornicata contains two major haplotype clades, both of which include samples from throughout the Atlantic coast. In this species there is no geographical differentiation among haplotypes but AMOVA analysis detects a small but statistically significant level of geographical structure. The population structure within the C. plana species complex appears also to vary with mode of development: C. atrasolea, a direct-developing species, has higher levels of population structure than does C. depressa, a sympatric planktotrophic species. The coincident occurrence of range endpoints and genetic breaks along the east coast of Florida in both direct-developing species and species with planktonic development indicates that this biogeographic break is not due to development-specific mechanisms such as hydrographic effects on larval recruitment.     

Sibling species complexes in sympatric populations of Petrotilapia Trewavas (Cichlidae, Lake Malawi)

The endemic Lake Malawi cichlid, Petrotilapia tridentiger Trewavas, was found to be a complex of sibling species. The various sympatric species show considerable homogeneity in body form and dentition but are separable on the basis of differences in adult coloration, size at maturity, depth distribution, size and degree of overlap of territories, aggression and courtship. Colour is considered to be important for specific mate recognition in this genus. Examination of wild populations is a valuable, perhaps essential, adjunct to classical taxonomy when dealing with the systematics of sibling species in Lake Malawi.     

Alsodes Australis,A New Species of Leptodactylid Frog from the Temperate Nothofagus Forest of Southern Chile and Argentina

A new species of frog, Alsodes australis, is described from the temperate Nothofagus forest of southern Chile and Argentina, based on adults and tadpoles. Up to now, no sympatric occurrence of another congeneric species is known from the type locality. The genus comprises three species groups. From a karyological point of view this species is included in the monticola group, whose species have 26 chromosomes.     

Interspecific competition and the evolution of plumage-colour variation in three closely related old world flycatchers Ficedula spp

When closely-related species are sympatric, there will be a coevolutionary selection pressure for species-specific characteristics to be developed, e.g. in plumage-colour. Because the three European Ficedula flycatchers have similar ecological requirements, we expected that some kind of interspecific signalling system had been evolved in sympatric regions of species of this genus. By using museum specimens we studied the seasonal and geographical variations in plumage-colour of the males of the three European species of the genus, and found that in areas of sympatry with F. albicollis , the subdominant species, F. hypoleuca , was lighter coloured than in allopatric areas. The plumage-colour of F. hypoleuca gradually changes from light in sympatric areas to dark in allopatric areas. A similar pattern was found for the eastern species F. semitorquata . The results of this paper, therefore, support the character displacement hypothesis.     

Limiting similarity and species assemblages in the short-tailed fruit bats

We analyzed the morphometrics of seven taxa of short-tailed fruit bats Chiroptera : Phyllostomidae : Carollia , ( Carollia benkeithi , Carollia brevicauda , Carollia castanea , Carollia perspicillata azteca , Carollia perspicillata perspicillata , Carollia sowelli and Carollia subrufa ) to assess within-taxon variation and to address the possibility that morphological distinctions facilitate sympatry within this genus of ecologically similar bats. Whereas morphological similarities were found among species in certain combinations of allopatric taxa, members of sympatric assemblages differ significantly, indicating that there is a consistent representation of morphotypes within various assemblages of Carollia despite changes in species composition across the range of the genus. Furthermore, the differences in the skull size and shape among taxa correspond to potential differences in food-handling abilities and point to dietary differentiation among species as a factor in the facilitation of coexistence.     

Description of a new long‐snouted beaked whale from the Late Miocene of Denmark: evolution of suction feeding and sexual dimorphism in the Ziphiidae (Cetacea: Odontoceti)

A new genus and species of Ziphiidae, Dagonodum mojnum gen. nov., sp. nov., from the upper Miocene Gram Formation (c. 9.9–7.2 Ma) represents the first occurrence of the family in Denmark. This long‐snouted ziphiid is characterized by two pairs of mandibular tusks, the Eustachian outlet that approximately levels with the dorsalmost margin of the posterior portion of the involucrum, and the left trapezoid nasal with a posteromedial projection into the frontal. A phylogenetic analysis including 25 species and 69 characters was conducted. Dagonodum mojnum is placed in a basal ziphiid clade as the sister taxon of Messapicetus. The specimen is probably a male, because it has enlarged tusks. Alternatively, females could also be involved in fights and develop erupted tusks as in the extant Berardius. Although less well supported, this interpretation proposes that aggressive interactions were not restricted to males in stem‐ziphiids. With a thickened thyrohyal and the presence of a precoronoid crest, D. mojnum was able to use suction feeding, but was less specialized to it compared to extant ziphiids. The elongated neck of D. mojnum less optimized to perform deep dives, and the shallow depth at which the Gram Formation was deposited corroborates the hypothesis that at least part of the stem‐ziphiids were not regular deep divers.     

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ABSTRACT

Five distinct song types of the genus Mecopoda were found in Southern India. Four were morphologically indistinguishable. Some of them were both sympatric and had synchronous breeding seasons. The songs of these five song types ranged from simple short chirps to highly complex songs with multiple components. The temporal patterns of the songs of the five song types were very distinctive, whereas their spectral features were similar. Component elements of the different songs were distinct despite overall similarity. The song types possibly represent sibling species.     

Evolutionary systematics of the Australian Cyzicidae (Crustacea,Branchiopoda, Spinicaudata) with the description of a new genus

In Australia's arid and semi‐arid zone, most aquatic habitats are nonpermanent. Although approximately 70% of its land surface belongs to these zones, very little is known of the iconic ‘large branchiopods’ that inhabit these important and widespread habitats. In the present study, we investigated 737 Australian specimens of the spinicaudatan taxa Caenestheria and Caenestheriella with a combination of one mitochondrial (cytochrome oxidase subunit I; COI) and three nuclear (elongation factor 1α, internal transcribed spacer 2, and 28S) markers to assess the diversity of species, their phylogenetic relationships, and phylogeographical history. The initial species delimitation was based on COI employing a combination of phylogenetic analyses and two automated approaches to species delimitation (general mixed Yule coalescent model and Automated Barcode Gap Discovery). The outcome was tested by the nuclear markers and considered under differing species concepts. The number of delineated species ranged from 14–27, in no case being in full agreement with any of the two automated approaches. The lower numbers resulted if inferred reproductive isolation, as required for the biological or Hennigian species concept, was employed. Although nuclear markers did not indicate ongoing reproduction, the lack of sympatric co‐occurrences inhibited inferences of definitive reproductive isolation in several instances. If monophyly or an ‘independent evolutionary fate’ was employed, as required for the phylogenetic or evolutionary species concepts, the species' distribution was of no importance and up to 27 species could be delimitated. Because the Australian representatives of both studied genera could not be clearly separated from each other but constitute a single monophyletic clade separated from all available non‐Australian representatives of these genera, we describe a new spinicaudatan genus Ozestheria gen. nov. to accommodate these species. Populations revealed relatively small levels of genetic differentiation over large areas of central and eastern Australia. By far the most pronounced levels of genetic differentiation were observed towards the north‐eastern regions, a pattern possibly explainable by ecological conditions and the movement of nomadic water birds that disperse resting eggs. © 2015 The Linnean Society of London     

Complex copulatory behavior and the proximate effect of genital and body size differences on mechanical reproductive isolation in the millipede genus Parafontaria

The role of species-specific genitalia in reproductive isolation is unclear. Males of the millipede genus Parafontaria use gonopods (modified eighth legs) charged with sperm from the genital openings of the second legs as intromittent organs. Males perform both preliminary and true intromission during mating. During preliminary intromission, a male attempts to insert his gonopods into the female genitalia before charging the gonopods with sperm. If this intromission is completed, it is followed by the ejaculation of sperm to the gonopods and true intromission for insemination. In two sympatric species of Parafontaria that lack effective precopulatory isolation, copulation was terminated without insemination because of preliminary intromission failure caused by mismatched genital and body sizes. Thus, mechanical isolation between these sympatric species resulted from morphological differentiation mediated by the obligatory preliminary intromission. These findings demonstrate the proximate importance of genital and body size differences for reproductive isolation within this genus of millipede.     

Genetic differentiation among host-associated Alebra leafhoppers (Hemiptera: Cicadellidae)

The limited importance ascribed to sympatric speciation processes via host race formation is partially due to the few cases of host races that have been reported among host populations. This work sheds light on the taxonomy of Alebra leafhoppers and examines the possible existence of host races among host-associated populations. The species of this genus show varying degrees of host association with deciduous trees and shrubs and, frequently, host populations of uncertain taxonomic status coexist and occasionally become pests. Allozyme electrophoresis of 21 Greek populations including sympatric, local and geographically distant samples collected on 13 different plant species, show that they represent at least five species: A. albostriella Fallén, A. viridis (Rey) (sensu Gillham), A. wahlbergi Boheman and two new species. Of these, one is associated to Quercus frainetto and other is specific to Crataegus spp. Significant genetic differences among sympatric and local host populations were found only in A. albostriella, between populations on Turkey oak, beech and common alder. It is suggested that the last two of these host populations may represent different host races. The results show that both the host plant and geographical distance affect the patterns of differentiation in the genus. The formation of some species seems to have been the result of allopatric speciation events while, for others, their origin can be equally explained either by sympatric or allopatric speciation.     

Phylogeography of the finless porpoise and potential implications for the taxonomy of Neophocaena spp.

The finless porpoise (genus Neophocaena) is a poorly known cetacean of great conservation concern. Within its range, from western Pacific to northwestern Indian Ocean, there are currently two species recognized (N. asiaeorientalis and N. phocaenoides), thought to be reproductively isolated since last glacial maximum, with the only sympatric overlap zone in Taiwan Strait. However, the genetic variation across the genus’ distribution has not yet been extensively studied, especially in the Indian Ocean. We performed an exhaustive review of molecular data of the finless porpoise across its range. Neighbor-net networks analyses based on two mitochondrial loci (control region/CR and cytochrome b/cyt b) suggest that finless porpoises from the Indian and Pacific Ocean constitute two distinct clades, well-defined by fixed mutations at both loci. A molecular clock analyses indicate early split (CR: 13.1 Ma, cyt b: 12.9 Ma) between these two oceanic lineages, while spatial genetic analyses further suggest that in the Pacific the divergence was primarily due to the taxon from Japanese waters rather than inter-species divergence across the Taiwan Strait. As extinction risks can be substantially underestimated if threatened species are pooled together with non-threatened, especially in the absence of long-distance migration, we suggest that the present 2-species taxonomy of the genus Neophocaena should be given further examination, with concerted sampling effort in the Western Indian Ocean. More research effort and genomic information is needed before taxonomic revisions can be considered; such further studies are strongly recommended as they may affect the current status classification of the species constituting the genus Neophocaena. Most notably, the narrow-ridged finless porpoise off Japan merits urgent conservation attention.     

Increased floral divergence in sympatric monkeyflowers

Sympatric sister species are predicted to have greater divergence in reproductive traits than allopatric sister species, especially if mating system shifts, such as the evolution of self-fertilization, are more likely to originate within the geographic range of the outcrossing ancestor. We present evidence that supports this expectation-sympatric sister species in the monkeyflower genus, Mimulus, exhibit greater divergence in flower size than allopatric sister species. Additionally, we find that sympatric sister species are more likely to have one species with anthers that overtop their stigmas than allopatric sister species, suggesting that the evolution of automatic self-pollination may contribute to this pattern. Potential mechanisms underlying this pattern include reinforcement and a stepping stone model of parapatric speciation.     

A test of the sympatric host race formation hypothesis in Neodiprion (Hymenoptera: Diprionidae)

Theory suggests that sympatric speciation is possible; however, its prevalence in nature remains unknown. Because Neodiprion sawflies are host specialists and mate on their hosts, sympatric speciation via host shifts may be common in this genus. Here, we test this hypothesis using near-complete taxonomic sampling of a species group, comprehensive geographical and ecological data, and multiple comparative methods. Host-use data suggest that host shifts contributed to the evolution of reproductive isolation in Neodiprion and previous work has shown that gene flow accompanied divergence. However, geographical data provide surprisingly little support for the hypothesis that host shifts occurred in sympatry. While these data do not rule out sympatric host race formation in Neodiprion, they suggest that this speciation mode is uncommon in the genus and possibly in nature.     

Species boundaries, specialization, and the radiation of sponge-dwelling alpheid shrimp

Microevolutionary studies and natural history suggest that host-specialization has promoted the high diversity of tropical sponge-dwelling snapping shrimps (Decapoda, Alpheidae, Synalpheus ). Yet the taxonomic difficulty of this genus has precluded rigorous tests of this hypothesis. S. rathbunae Coutiére is among the most abundant invertebrates inhabiting the framework of sponges and dead coral that forms the floor of Caribbean coral reefs. Even within a small area S. rathbunae exhibits the apparently wide variation in size, color, and morphology that has long frustrated efforts to identify and define species boundaries within this large (> 100 described species) genus. Here I show that sympatric populations of this nominal species occupying different sponge hosts display clear, concordant differences in allozyme genotypes and in multivariate morphometries, confirming that the populations represent three distinct biological species. Moreover, careful field sampling revealed that the three S. rathbunce taxa and the closely related S. filidigitus Armstrong showed almost no overlap in the species of hosts occupied. Interestingly, while there was significant differentiation between Belizean and Panamanian populations of the one taxon that occurred at both sites (∼1500 km apart), these populations were recognizable as conspecific using both genetic and morphological characters. These results show that (1) diversity of Synalpheus , which is already among the most species-rich crustacean genera, is probably several times higher than currendy recognized, and (2) species of sponge-dwelling Synalpheus are highly host-specific, with related species distinctly segregated among hosts. Together with previous evidence of host race differentiation within shrimp species, these results suggest a primary role for resource specialization in the origin and/or maintenance of this group's characteristically high diversity.