Evolutionary convergence of body shape and trophic morphology in cichlids from Lake Tanganyika

A recent phylogenetic analysis of mitochondrial DNA sequences from eretmodine cichlids from Lake Tanganyika indicated independent origins of strikingly similar trophic specializations, such as dentition characters. Because genetic lineages with similar trophic morphologies were not monophyletic, but instead were grouped with lineages with different trophic phenotypes, raises the question of whether trophic morphology covaries with additional morphological characters. Here, we quantified morphological variation in body shape and trophically associated traits among eretmodine cichlids using linear measurements, meristic counts and landmark‐based geometric morphometrics. A canonical variates analysis (CVA) delineated groups consistent with dentition characters. Multivariate regression and partial least squares analyses indicated that body shape was significantly associated with trophic morphology. When the phylogenetic relationships among taxa were taken into account using comparative methods, the covariation of body shape and trophic morphology persisted, indicating that phylogenetic relationships were not wholly responsible for the observed pattern. We hypothesize that trophic ecology may be a key factor promoting morphological differentiation, and postulate that similar body shape and feeding structures have evolved multiple times in independent lineages, enabling taxa to invade similar adaptive zones.     

Flat-faced longhorn beetles (Coleoptera: Cerambycidae: Lamiinae) from the Neotropical region: new species from Ecuador,nomenclatural changes and notes

Summary

The differences between Anisopodus White, 1853 and Hyperplatys Haldeman, 1847 are discussed and the gender of the latter is corrected. Notes on Hyperplatys pusillus (Bates, 1863) are provided. Hyperplatys nigrisparsus is considered a new rank for H. pusillus nigrisparsus (Bates, 1885). This species is newly recorded from Brazil (Amazonas). Alcidion laetulum Bates, 1880 (currently Nealcidion laetulum), Alcidion costatum Monné & Martins, 1976 (currently Nealcidion costatum), and Nealcidion murinum Monné, 1998, are synonymized with Alcidion bispinum Bates, 1863 (currently Nealcidion bispinum). Anisopodus melzeri Gilmour, 1965 is transferred to Hyperplatys with a new record from the Brazilian state of Mato Grosso do Sul. Notes on a second specimen of Confluentia colombiana (Gilmour, 1950) are provided. Three new species are described from Ecuador: Nealcidion kayi n. sp. (Acanthocinini), Hyperplatys pichinchensis n. sp. (Acanthocinini) and Confluentia quijos n. sp. (Colobotheini).     

Ontogenic diet shifts and scale-eating in Roeboides dayi, a Neotropical characid

Scale feeding (lepidophagy) has been documented for a variety of Neotropical fish taxa, including the characid genus Roeboides. Ontogenetic differentiation of jaws and snout teeth allow larger Roeboides to remove scales, however, the less specialized tooth/jaw structure in Roeboides dayi, indicates that it may be a facultative scale feeder. Population dynamics and diets of R. dayi in a Venezuelan lowland swamp/creek and a piedmont stream were compared over an annual cycle. Juvenile R. dayi consumed aquatic insect larvae and microcrustacea, and although spawning was year-round at both sites, most reproduction occurred during the wet season when the availability of these resources was greatest for juveniles. At both sites, larger R. dayi fed on a combination of invertebrate prey and fish scales, the former being more important at the piedmont site, and the latter being especially important during initial low water conditions at both sites. In the lowland stream, the reduction of aquatic habitat during the early dry season created higher fish densities and a more profitable environment for scale-feeders. Insectivory probably was less profitable during this early low water period due to interspecific competition for reduced aquatic insect stocks.     

Phylogenetic relationships of Middle American cichlids (Cichlidae, Heroini) based on combined evidence from nuclear genes, mtDNA, and morphology

Heroine cichlids are the second largest and very diverse tribe of Neotropical cichlids, and the only cichlid group that inhabits Mesoamerica. The taxonomy of heroines is complex because monophyly of most genera has never been demonstrated, and many species groups are without applicable generic names after their removal from the catch-all genus Cichlasoma (sensu Regan, 1905). Hence, a robust phylogeny for the group is largely wanting. A rather complete heroine phylogeny based on cytb sequence data is available [Concheiro Pérez, G.A., Říčan O., Ortí G., Bermingham, E., Doadrio, I., Zardoya, R. 2007. Phylogeny and biogeography of 91 species of heroine cichlids (Teleostei: Cichlidae) based on sequences of the cytochrome b gene. Mol. Phylogenet. Evol. 43, 91–110], and in the present study, we have added and analyzed independent data sets (nuclear and morphological) to further confirm and strengthen the cytb-phylogenetic hypothesis. We have analyzed a combined cytb-nuclear (RAG1 and two S7 introns) data set of 48 species representing main heroine lineages to achieve further resolution of heroine higher taxonomic levels and a combined cytb-morphological data set of 92 species to stabilize generic taxonomy. The recovered phylogenies supported the circumamazonian—CAM—Heroini (sensu Concheiro Peréz et al., 2007) as a monophyletic group, that could be divided into six main clades: (1) australoheroines (the southernmost heroine genus Australoheros), (2) nandopsines (the Antillean genus Nandopsis), (3) caquetaines (including the north western Amazonian genera Caquetaia and Heroina), (4) astatheroines (including Astatheros, Herotilapia and Rocio), (5) amphilophines (including Amphilophus and related genera), and (6) herichthyines (including Herichthyis and related genera). Nuclear and mitochondrial data partitions arrived at highly congruent topologies. Suprageneric relationships were influenced mainly by the nuclear signal, as well as the most basal phylogenetic position of Australoheros within CAM heroines. The new phylogeny of the tribe Heroini provides robust framework to stabilize the taxonomy of the group and for future comparative studies on these morphologically and ecologically diverse freshwater fishes. Morphology was mostly informative at the genus level and aid in determining the monophyly and composition of heroine genera. Upon acceptance of all putative genera, as recovered in this study, the Heroini would be with 35 genera the most genus-rich clade of Neotropical cichlids.     

Evolutionary conservatism and convergence both lead to striking similarity in ecology,morphology and performance across continents in frogs

Many clades contain ecologically and phenotypically similar species across continents, yet the processes generating this similarity are largely unstudied, leaving fundamental questions unanswered. Is similarity in morphology and performance across assemblages caused by evolutionary convergence or by biogeographic dispersal of evolutionarily conserved ecotypes? Does convergence to new ecological conditions erase evidence of past adaptation? Here, we analyse ecology, morphology and performance in frog assemblages from three continents (Asia, Australia and South America), assessing the importance of dispersal and convergent evolution in explaining similarity across regions. We find three striking results. First, species using the same microhabitat type are highly similar in morphology and performance across both clades and continents. Second, some species on different continents owe their similarity to dispersal and evolutionary conservatism (rather than evolutionary convergence), even over vast temporal and spatial scales. Third, in one case, an ecologically specialized ancestor radiated into diverse ecotypes that have converged with those on other continents, largely erasing traces of past adaptation to their ancestral ecology. Overall, our study highlights the roles of both evolutionary conservatism and convergence in explaining similarity in species traits over large spatial and temporal scales and demonstrates a statistical framework for addressing these questions in other systems.     

Spore morphology and ultrastructure of Cyathea (Cyatheaceae,Pteridophyta) species from southern South America

Spore sculpture and wall structure of eight Cyathea (Cyatheaceae) species from southern South America were studied using light microscopy (LM), scanning electron microscopy (SEM) and transmission electron microscopy (TEM) techniques. Two layers, i.e. an inner and an outer layer, were observed in the perispore. The inner layer has two strata: the inner stratum is attached to the exospore and composed of rodlets tangentially oriented to the spore surface and randomly intermixed; the outer stratum consists of a three-dimensional network of rodlets with either free or fused distal edges forming spinules. The outer layer is thin, darkly contrasted and covers the rodlets. In most cases, the exospore has two layers and a pitted surface. In Cyathea atrovirens, the exospore surface is smooth, while in C. delgadii and C. myriotricha it is verrucate. The homogeneity of perispore features within the genus Cyathea is evident, while exospore features are heterogeneous. The exospore has different kinds of surface-structures that are of potential interest for assessing evolutionary trends within the group.     

Identifying mesopredator release in multi-predator systems: a review of evidence from North America

1. The mesopredator release hypothesis, defined as the change in distribution, abundance or behaviour of a middle-ranking predator in response to a decrease in density or distribution of an apex predator, is an increasingly popular topic in ecology. Terrestrial mesopredators have been reported as being released in multiple systems globally, particularly in North America, over the past century.
2. We reviewed 2687 scientific articles, of which we determined that 38 met our criteria for investigating mesopredator release (MR) in terrestrial North American mammalian predators.
3. We observed no support or mixed support for MR in 46% of all relevant studies, including conflicting evidence between measures (mesopredator distribution, abundance or behaviour) within a given study and among studies of the same community in different settings.
4. To advance the study of MR, we provide a conceptual framework that 1) highlights the multiple spatial, temporal and ecological scales at which mesopredator responses can occur; 2) suggests the relative weight of evidence for MR that is provided by measures of mesopredator responses at each scale; and 3) clearly defines the threshold for determining when MR is occurring.
5. In increasingly reshuffled predator communities with declining apex predators, there is a need for future studies to assess in more detail the contexts in which mesopredator behavioural responses scale up to the population-level processes and species-level distribution changes needed to identify these responses as MR.

     

Annual and seasonal changes in diets of martens: evidence from stable isotope analysis

Theory predicts that generalist predators will switch to alternative prey when preferred foods are not readily available. Studies on the feeding ecology of the American marten (Martes americana) throughout North America suggest that this mustelid is a generalist predator feeding largely on voles (Microtus sp.; Clethrionomys sp.). We investigated seasonal and annual changes in diets of martens in response to the changing abundance of small rodents (Peromyscus keeni, and Microtus longicaudus) on Chichagof Island, Southeast Alaska, using stable isotope analysis. We hypothesized that martens would feed primarily on small rodents during years with high abundance of these prey species, whereas during years of low abundance of prey, martens would switch to feed primarily on the seasonally available carcasses of salmon. We also hypothesized that home-range location on the landscape (i.e., access to salmon streams) would determine the type of food consumed by martens, and martens feeding on preferred prey would exhibit better body condition than those feeding on other foods. We live-captured 75 martens repeatedly, from mid-February to mid-December 1992–1994. We also obtained marten carcasses from trappers during late autumn 1991 and 1992, from which we randomly sub-sampled 165 individuals. Using stable isotope ratios and a multiple-source mixing model, we inferred that salmon carcasses composed a large portion of the diet of martens in autumn during years of low abundance of rodents (1991 and 1992). When small rodents were available in high numbers (1993 and 1994), they composed the bulk of the diet of martens in autumn, despite salmon carcasses being equally available in all years. Selection for small rodents occurred only in seasons in which abundance of small rodents was low. Logistic regression revealed that individuals with access to salmon streams were more likely to incorporate salmon carcasses in their diet during years of low abundance of small rodents. Using stable isotope analysis on repeated samples from the same individuals, we explored some of the factors underlying feeding habits of individuals under variable ecological conditions. We were unable to demonstrate that body weights of live-captured male and female martens differed significantly between individuals feeding on marine-derived or terrestrial diets. Therefore, martens, as true generalist predators, switched to alternative prey when their principal food was not readily available on a seasonal or annual basis. Although salmon carcasses were not a preferred food for martens, they provided a suitable alternative to maintain body condition during years when small rodents were not readily available. Received: 1 May 1996 / Accepted: 24 February 1997     

Trophic position and seasonal changes in the diet of the red wood ant Formica aquilonia as indicated by stable isotope analysis

1. Red wood ants are among the most numerous generalist predators and strongly affect the composition of arthropod communities in forest ecosystems. However, their trophic position remains poorly understood. Stable isotope analysis was applied to study the trophic position of Formica aquilonia and reveal seasonal changes in its trophic links with both myrmecophilous aphids and other invertebrates in a mixed forest of western Siberia. 2. The δ¹⁵N values of F. aquilonia exceeded those of herbivores and aphids by approximately 3.5‰. Despite obligate trophobiotic relationships with aphids, F. aquilonia occupied the trophic position of first‐order predator. The higher content of ¹³C in the worker ants, compared with members of grazing food chains, was explained by their consumption of ¹³C‐enriched aphid honeydew. 3. Myrmecophilous tree‐dwelling aphids were enriched in ¹³C and ¹⁵N relative to grass‐inhabiting species, and the honeydew of tree‐dwelling aphids had higher δ¹³C values than those of the honeydew of grass‐inhabiting aphids. 4. The decrease in δ¹³C values of the worker ants from spring and summer to autumn apparently reflected the transition from the collection of tree sap and feeding on the aphid honeydew from trees with high ¹³C content in the spring and early summer to a more diverse liquid diet in late summer, which included ¹³C‐depleted honeydew of aphids from herbs. 5. The prevalence of the ¹⁵N‐depleted aphid honeydew in the ants' diet in the second half of the summer is discussed as one possible explanation for the seasonal decline in δ¹⁵N values of the worker ants.     

Characterizing the trophic niches of stocked and resident cyprinid fishes: consistency in partitioning over time,space and body sizes

Hatchery‐reared fish are commonly stocked into freshwaters to enhance recreational angling. As these fishes are often of high trophic position and attain relatively large sizes, they potentially interact with functionally similar resident fishes and modify food‐web structure. Hatchery‐reared barbel Barbus barbus are frequently stocked to enhance riverine cyprinid fish communities in Europe; these fish can survive for over 20 years and exceed 8 kg. Here, their trophic consequences for resident fish communities were tested using cohabitation studies, mainly involving chub Squalius cephalus, a similarly large‐bodied, omnivorous and long‐lived species. These studies were completed over three spatial scales: pond mesocosms, two streams and three lowland rivers, and used stable isotope analysis. Experiments in mesocosms over 100 days revealed rapid formation of dietary specializations and discrete trophic niches in juvenile B. barbus and S. cephalus. This niche partitioning between the species was also apparent in the streams over 2 years. In the lowland rivers, where fish were mature individuals within established populations, this pattern was also generally apparent in fishes of much larger body sizes. Thus, the stocking of these hatchery‐reared fish only incurred minor consequences for the trophic ecology of resident fish, with strong patterns of trophic niche partitioning and diet specialization. Application of these results to decision‐making frameworks should enable managers to make objective decisions on whether cyprinid fish should be stocked into lowland rivers according to ecological risk.     

Scale morphology and squamation patterns in cichlids (Teleostei, Perciformes): A comparative study

Problems in cichlid systematics call for new characteristics to be exploited. Scale surface morphology and squamation patterns could provide novel and useful information. A comparative study comprising 105 African, 10 American, and eight outgroup species was conducted to identify the most useful scale and squamation characters within the family and to clarify their phylogenetic significance. A great number of obviously genetically fixed characters were established. At least IS scale granulation types could be identified, proving to be ofparticular value for systematic purposes. Squamation patterns on the head, breast and fins were also found to be of interest in this context. Different character states are described and their distribution is reviewed, together with a discussion of their plesiomorphic status. From the results it can be concluded that for the Cichlidae scale and squamation studies can be valuable tools in investigating phylogenetic relationships.     

Lapita diet and subsistence strategies on Watom Island,Papua New Guinea: New stable isotope evidence from humans and animals

Stable isotope ratios (δ¹³C and δ¹⁵N) were analyzed from the bone collagen of individuals (n = 8) from a Lapita burial ground (ca. 2800–2350 BP) on Watom Island, located off northeast New Britain in the Bismarck Archipelago. The aim of this study was to assess the diet and subsistence strategies of humans that lived during the later Lapita period in Near Oceania. To aid in the interpretation of the human diet we analyzed the stable isotope ratios of faunal material from the site (n = 27). We also aim to assess methods of animal husbandry at the site over time from an analysis of the stable isotope ratios (δ¹³C and δ¹⁵N) of pig bones (n = 22) from different temporal periods (Lapita, post‐Lapita, and late prehistoric). The protein diet of the humans consisted of marine organisms from the inshore environment and some deep‐water species, most likely marine turtle, in addition to higher trophic level terrestrial foods, likely pig and native animals (e.g., fruit bat, Cuscus and bandicoot). Although the sample sizes were small, females (n = 4) displayed more variable δ¹³C and δ¹⁵N values compared with males (n = 4), which may be associated with the movement of adult females to the island. The stable isotope analysis of the pig bones indicated that there were few differences between the diets of the pigs from the Lapita and post‐Lapita layers, suggesting that the method of pig husbandry was similar between these two periods and was likely relatively free‐range. Am J Phys Anthropol 157:30–41, 2015. © 2015 Wiley Periodicals, Inc.     

Revolution in food web analysis and trophic ecology: diet analysis by DNA and stable isotope analysis

Characterization of energy flow in ecosystems is one of the primary goals of ecology, and the analysis of trophic interactions and food web dynamics is key to quantifying energy flow. Predator‐prey interactions define the majority of trophic interactions and food web dynamics, and visual analysis of stomach, gut or fecal content composition is the technique traditionally used to quantify predator‐prey interactions. Unfortunately such techniques may be biased and inaccurate due to variation in digestion rates ( Sheppard & Hardwood 2005 ); however, those limitations can be largely overcome with new technology. In the last 20 years, the use of molecular genetic techniques in ecology has exploded ( King et al. 2008 ). The growing availability of molecular genetic methods and data has fostered the use of PCR‐based techniques to accurately distinguish and identify prey items in stomach, gut and fecal samples. In this month’s issue of Molecular Ecology Resources, Corse et al. (2010) describe and apply a new approach to quantifying predator‐prey relationships using an ecosystem‐level genetic characterization of available and consumed prey in European freshwater habitats ( Fig. 1a ). In this issue of Molecular Ecology, Hardy et al. (2010) marry the molecular genetic analysis of prey with a stable isotope (SI) analysis of trophic interactions in an Australian reservoir community ( Fig. 1b ). Both papers demonstrate novel and innovative approaches to an old problem – how do we effectively explore food webs and energy movement in ecosystems?

Figure 1 Open in figure viewer PowerPoint The aquatic habitats used for two studies of diet and trophic interactions that employed molecular genetic and stable isotope analyses. Panel a: Example of Rhone basin habitat (France) where fish diet was determined using PCR to classify prey to a series of ecological clades (photo by Emmanuel Corse). Panel b: A weir pool on the lower Murray River (Australia) where food web and prey use was evaluated using a combination of advanced molecular genetic and stable isotope analyses (photo credit: CSIRO).     

Body shape differences in a pair of closely related Malawi cichlids and their hybrids: Effects of genetic variation,phenotypic plasticity,and transgressive segregation

Phenotypic differences may have genetic and plastic components. Here, we investigated the contributions of both for differences in body shape in two species of Lake Malawi cichlids using wild‐caught specimens and a common garden experiment. We further hybridized the two species to investigate the mode of gene action influencing body shape differences and to examine the potential for transgressive segregation. We found that body shape differences between the two species observed in the field are maintained after more than 10 generations in a standardized environment. Nonetheless, both species experienced similar changes in the laboratory environment. Our hybrid cross experiment confirmed that substantial variation in body shape appears to be genetically determined. The data further suggest that the underlying mode of gene action is complex and cannot be explained by simple additive or additive‐dominance models. Transgressive phenotypes were found in the hybrid generations, as hybrids occupied significantly more morphospace than both parentals combined. Further, the body shapes of transgressive individuals resemble the body shapes observed in other Lake Malawi rock‐dwelling genera. Our findings indicate that body shape can respond to selection immediately, through plasticity, and over longer timescales through adaptation. In addition, our results suggest that hybridization may have played an important role in the diversification of Lake Malawi cichlids through creating new phenotypic variation.     

Endocranial morphology of Pygoscelis calderensis (Aves,Spheniscidae) from the Neogene of Chile and remarks on brain morphology in modern Pygoscelis

The endocranial anatomy of Pygoscelis calderensis, a fossil species from the Bahía Inglesa Formation (Middle Miocene–Pliocene) of Chile, South America, was described through CT scans. Reconstructions of the fossil P. calderensis and endocasts for the living Pygoscelis adeliae, and Pygoscelis papua are provided here for the first time. Comparisons with the extant congeneric species P. adeliae, Pygoscelisantarctica and P. papua indicate that the morphological pattern of the brain and inner ear of the extant pygoscelids has been present already in the Middle Miocene. The neurological morphology suggests that the paleobiology of the extinct form would have been similar to the extant species. It was probably true for diet, feeding behaviour and diving kinematics.     

Evolutionary history of the northern leopard frog: reconstruction of phylogeny, phylogeography, and historical changes in population demography from mitochondrial DNA

This study uses a combined methodological approach including phylogenetic, phylogeographic, and demographic analyses to understand the evolutionary history of the northern leopard frog, Rana pipiens. We tested hypotheses concerning how (or if) known geological events and key features of the species biology influenced the contemporary geographic and genetic distribution of R. pipiens. We assayed mitochondrial DNA variation from 389 individuals within 35 populations located throughout the species range. Our a priori expectations for patterns and processes influencing the current genetic structure of R. pipiens were supported by the data. However, our analyses revealed specific aspects of R. pipiens evolutionary history that were unexpected. The phylogenetic analysis indicated that R. pipiens is split into populations containing discrete eastern or western haplotypes, with the Mississippi River and Great Lakes region dividing the geographic ranges. Nested clade analysis indicated that the biological process most often invoked to explain the pattern of haplotype position is restricted gene flow with isolation by distance. Demographic analyses showed evidence of both historical bottlenecks and population expansions. Surprisingly, the genetic evidence indicated that the western haplotypes had significantly reduced levels of genetic diversity relative to the eastern haplotypes and that major range expansions occurred in both regions well before the most recent glacial retreat. This study provides a detailed history of how a widespread terrestrial vertebrate responded to episodic Pleistocene glacial events in North America. Moreover, this study illustrates how complementary methods of data analysis can be used to disentangle recent and ancient effects on the genetic structure of a species.     

Inter-species hybridization among Neotropical cats of the genus Leopardus, and evidence for an introgressive hybrid zone between L. geoffroyi and L. tigrinus in southern Brazil

Natural hybrid zones between distinct species have been reported for many taxa, but so far, few examples involve carnivores or Neotropical mammals in general. In this study, we employed mitochondrial DNA (mtDNA) sequences and nine microsatellite loci to identify and characterize a hybrid zone between two Neotropical felids, Leopardus geoffroyi and L. tigrinus, both of which are well-established species having diverged from each other c. 1 million years ago. These two felids are mostly allopatric throughout their ranges in South America, with a narrow contact zone that includes southern Brazil. We present strong evidence for the occurrence of hybridization between these species and identify at least 14 individuals (most of them originating from the geographical contact zone) exhibiting signs of interspecific genomic introgression. The genetic structure of Brazilian L. tigrinus populations seems to be affected by this introgression process, showing a gradient of differentiation from L. geoffroyi correlated with distance from the contact zone. We also corroborate and extend previous findings of hybridization between L. tigrinus and a third related felid, L. colocolo, leading to an unusual situation for a mammal, in which the former species contains introgressed mtDNA lineages from two distinct taxa in addition to its own.